Physiology

BY

An

Instructional

This
published
visable

have

the

physiology

of
that

New

are

which

American

of

basis

Academy

joints

of this

of Orthopaedic

is drawn

Instructional

and to include
verified
or which

from
5,6,7#{149}

previously
It seems

certain
relatively
are controversial

ad-

recent
usually

an extensive

and

MacConaill

fibrous,

cartilaginous,

features,

and

In

a fibrous
tissue.

gomphosis,
as a third

joint,
The

the
type.

sometimes

two

joint

chief

boncs

joints

and

bone

a tooth

of cartilaginous

functions.

On the

be classified

a synarthrosis,

of fibrous
the

Cartilaginous

The

may

into

basis

three

of

main

Joints

called

types

between

joints

synovial.
Fibrous

fibrous

given

largely
Course

has

structural

are

Surgeons

Davies,

characteristic

references

MICHIGAN

however,

types:

a few

DETROIT,

; each,

most

Only

The

up-to-date
have not been

material

studies

omitted.

the

M.D.t,

bibliography
(see particularly
the recent
book by Barnett,
and the paper
by Putschar).
Joints
have
certain
common
structural
features
and
the

been

the

GARDNER,

Lecture,

accounts
bring

D.

Course

account
to

findings.

ERNEST

of Joints

joints

the

are

bones

sutures

are

and

of its socket,

united

by

syndesmoses.

is sometimes

A
classed

Joints

are

united

either

by

hyaline

cartilage

or by

fibrocartilage.
Hyaline

Cartilage

Hyaline

Joints

cartilage

sometimes

joints

synchondroses.

cartilaginous

skeleton,

that

An

it joins.

cartilage

joints

are
The

and

it serves

epiphyseal
are

Fibrocartilaginous

sometimes

hyaline

plate

obliterated,

called

cartilage

as a growth
is an example

that

is,

replaced

primary

cartilaginous

is a persistent

zone

for

part

one

joints

of the

or both

of this

type

of joint.

by

bone,

when

and

embryonic

of the

bones

Most

hyaline

growth

ceases.

Joints

Fibrocartilaginous
joints
are sometimes
called
secondary
cartilaginous
joints
and sometimes
amphiarthroses.
The term symphysis
is also applied
to it. The bones
are united
by fibrocartilage
during
some phase
of their
existence
; the fibrocartilage
is separated
from
the bones
by thin plates
of hyaline
cartilage.
Intervertebral
discs
and

the pubic
symphysis
In the intervertebral
aspects
of the
disc

lower
*

The personal

the
Institutes
from

investigations
reported
Michigan
Chapter
of the Arthritis
of Health,
United
States
Public

t I)epartment

of Anatomy,

College

the upper
and
the period
of

on in this paper
were supported
by research
grants
and Rheumatism
Foundation
and from the National
Health
Service,
A-532.
of Medicine,
Wayne
State
University,
1401 Rivard,

7, Michigan.

Detroit

VOL.

are examples
of fibrocartilaginous
joints.
disc, thin, hyaline
cartilage
plates
separate
from the adjacent
vertebral
bodies.
During

45-A,

NO.

5. JULY

1963

1061

1062

A.A.O.S.

growth

each

in height.

plate

provides

In a young

collagenous
bundles
hyaline
plates
and
cartilage.
The

one of the

adult,

nucleus

contains

pulposus

fibrous

chordal
elements.
and the differences
Few,
in

if any,
its

nerve

is white,

and

cartilaginous

supply

trunks

and

longitudinal

the

synovial
capsule

fibrous

which

vertebral

of a series

above
The

and below,
innermost

to the
lamellae

and

and

body
margins
contain

semigelatinous.

that

It

gradually

grows

of lamellae

of the
fibro-

and

replace

the

noto-

disc tends
to become
cartilaginous
and the annulus
are often
lost.
except
on its surface
or perhaps

sensory

fibers

from

ligament.

(recurrent

meningeal)

spinal

nerves

and

Similar

fibers

from

nerves

supply

the

Joints

joints,
also termed
diarthrodial
and by ligaments,
but are otherwise

joints,
the
separated

bones
are united
by a cavity.
The

by synovial
membrane,
which
and lubricates
the joint.

produces

the

by a
inner

synovial

There
may be intra-articular
ligaments
or tendons
which are usually
covered
synovial
membrane.
The
joint
cavity
is sometimes
partially
or completely
divided
by fibrous
or fibrocartilaginous
discs or menisci
; these
are not covered
membrane
of the bones

Types

of

Synoiial

Synovial
her

of

for

example,

more

joints

only

and

extended.

at their
periphery.
which
is usually

The
hyaline.

may

be classified

surfaces.

as simple

A simple

or compound

joint

has

one

according

pair

of

to the

interphalangeal
joint.
A compound
joint,
such as
pair of articulating
surfaces.
joints
may be classified
according
to axis of movement.

one

axis

of rotation,

Joints

can be flexed,
with
at least

articular

articulating

an
one

Synovial
have

except
to a certain
extent
are covered
with cartilage,

by
subby

Joints

articulating
than

of

amorcells. It

ligament.

aspect
of the capsule
is lined
fluid that
fills the joint
cavity

synovial
surfaces

of

consists

cells, cartilage
cells,
and contains
notochordal

longitudinal

sinuvertebral

the

consists

Synovial

In

from

elements

anterior

LECTURE

fibrosus

glistening,

Vasomotor

trunks

sympathetic

zones

fibers,
connective-tissue
At birth,
it is mucoid

lamellae.

sympathetic

two

annulus

With advancing
age, the entire
between
the nucleus
pulposus
endings
are present
in the disc

outermost

posterior

the

COURSE

that
are anchored,
to the ring epiphyses.

fine strands
of collagenous
phous
ground
substance.
also

INSTRUCTIONAL

with

extended,
three
axes,

two

for
axes

instance,

a hinge

of movement,

joint,

such

abducted,
and adducted
such as a ball-and-socket

which

the

can

as saddle

elbow,

has

A joint

may

only

and

but cannot
joint,
all

num-

surfaces,

be flexed

ellipsoid

be rotated.
movements

joints,
In joints
can take

place.
lar

Finally,
surfaces.

synovial
joints
may
The more
common

and-socket,

ellipsoidal,

Movements

at Synovial

Movements

at

pivot,

be classified
types
are

and

saddle

according
to the
plane,
hinge,
and
joints

are

less

shapes
condylar

of their articujoints.
Ball-

common.

Joints
synovial

joints

include

movements

occur

active,

passive,

and

accessory

move-

ments.
Three

types

of active

movements,
(2) angular
or about
an anteroposterior
ments
about
a longitudinal
Passive

movements

at synovial

joints:

movements
about
a horizontal
axis
axis (abduction
and adduction)
axis (medial
and lateral).
are

those

produced

by

an

external

(1) gliding
,

or slipping

(flexion
and extension)
and (3) rotatory
moveforce,

such

as

gravity

or an examiner.
THE

JOURNAL

OF

BONE

AND

JOINT

SURGERY

E.

Accessory
duced

by the

joint

but

the

these

GARDNER:

movements

only

wrist

D.

(often

classified

manipulation

of a subject,

muscle

PHYSIOLOGY

one

OF

as passive)

of an examiner.
can

bring

arrangements

about

are

such

1063

JOINTS

are

For

those

example,

a certain

amount

that

the

subject

and

Function

that

can

be

pro-

by manipulating
of gliding

cannot

and

the
rotation,

actively

carry

out

movements.
Structure

Synovial
are

friction

lubrication

joints
are
minimized
depends

(synovial

fluid)

Synovial

specialized
for more
or less free
by the lubrication
mechanisms

on a number

and

of factors,

incongruous

Membrane

and

Synovial

surface
surface

from
tissue

which
varies

cavity.

articular
acid,

cartilage.
sometimes

through
ground

cells

1 per

cent

the hyaluronic
by enzymatic
greater

viscosity

blood
plasma.
that
factors
weight

synovial

termed

mucin,

form

which

The

fluid

is produced

to the

by cells
are the
Solutions

of synovial

are

lymphatic

vessels

a relatively

smooth

pads project.
or fatty.

presence

The

sub-

of the

synovial

drops

membrane
in the
of more

acid

to that

of water

by enzymatic
hydrolysis,
varies
with temperature,
derived

chiefly

a number
of proteins
are present
shape
and charge
seem
important,

of

of hyaluronic

mucopolysaccharides
of hyaluronic
fluid

fluid

surface
of
network

of synovial
fluid (synovia),
and aids in the nutrition

entirely

just
as
tissues.

of synovial

However,
of molecular

form

formation
the joint

almost

viscosity

contains

and fat
areolar,

(by acid precipitation,

The viscosity
also

temperature.
constituents

lubricating

cartilage).

membrane

for the
lubricates

reactions,
connective

gels.

effects
of
Effective

that lines the inner

feature
is a capillary

also

of villi, folds,
may be fibrous,

is due

acid is removed
depolymerization)

at lower
The other

membrane

of

is responsible
fluid that

energy-requiring
substance
of other

than

Synovial

Its

a viscous

(articular

connective
tissue
morphological

a variable
number
in thickness
and

Synovial
membrane
is a highly
viscous

which

surfaces

The
joints.

Fluid

Synovial
membrane
is a vascular
the joint capsule.
Its most characteristic
adjacent
to the joint
and a few nerve
fibers.
The
connective-tissue

particularly

bearing

movement.
of these

by

if
or

being

dialysis

from

in concentrations
such
as well as molecular

#{176}.

Synovial
rived

from

Articular

fluid
the

also

lining

normally

contains

a few

cells,

mostly

mononuclear,

de-

tissue.

Cartilage

Articular
chondroitin
calcified.
chondral

cartilage

is usually

hyaline,

the

matrix

of which

consists

chiefly

of

sulphate
A and C. The
part
immediately
adjacent
to bone is usually
During
growth,
articular
cartilage
provides
the growth
zone for endoossification
in the epiphysis.
The cartilage
can also regenerate
and repair

defects.
When
growth
ceases,
articular
cartilage
loses
However,
some growth
is still possible
and
use can be replaced
Furthermore,
growth

repair.
normal

in the
stress
being
These
is the
VOL.

remodeling

of adult

joints

that

may

result

much
of its power
of growth
and
thus cartilage
that is lost during
of cartilage
is directly
involved
from

mechanical

or pathological

8,9

Cartilage
able
to
qualities
mechanism
45-A,

NO.

is a resilient
and elastic
tissue
that
absorb
synovial
fluid
which
diffuses
are

5, JULY

important
by which
1963

factors
nutritive

has the
through

spongelike
property
of
the cartilage
matrix.

in effective
lubrication.
The
materials
reach
the cartilage

diffusion
of fluid
cells. These
ma-

1064

A.A.O.S.

terials
in

are

the

derived

arterial

Lubrication

synovial

at the

fluid,

COURSE

from

periphery

LECTURE

epiphyseal

of the

vessels,

and

from

capillaries

joint.

Mechanisms

Joint
is much
to

from

circle

INSTRUCTIONAL

lubrication
lower

is extremely

than

that

of ice

effective,
sliding

so effective

on

ice.

Joint

be an example
of hydrodynamic
or fluid-film
presence
of incongruous
bearing
surfaces

During

range
of movement).
spaces
occupied
by

movement,

synovial

fluid

of some importance
in joint
been presented
to the effect
tion,
perhaps
lubrication,

with a contribution
the nature
of the

of prime
The

importance
shape
of the

and

coefficient

the

moving

lubrication.
that joint

adherent
particularly

such

as synovial
However,

are

considered

essential
do not

permits

these

condition
fit perfectly
the

spaces,

establish-

and

a pressure

moving
surfaces
separated
by a film of fluid.
Menisci,
intra-articular
discs, fat pads, and
fluid throughout
the joint and are therefore
Recently,
lubrication

experiments
is an example

of quasihydrodynamic
surfaces
and

the

and arguments
of boundary

lubrication.
quality
of the

In boundary
lubricant
are

surfaces

and

the

viscosity

lubricant

unimportant,

separated

by lubricant

studies

lend

of the

velocity

a few

molecules

only

to, or incorporated
into,
the
well adapted
for slow-moving,
joints.
still other

of the

is independent

movement.

thick,

surfaces.
heavily

Boundary
loaded,

to the

hydrodynamic

support

are
of

the

lubrication
may
Any mechanical

Some

wear-and-tear

normal

activity,

mon

result

to the

is the

extent

tear may
esses that

occur
under
certain
system
wears
with

conditions.
time,
and

(also
although

called
use-destruction
efficient
lubrication

wearing

away

of exposing,

lubricant
would
joints,

theory

be exaggerated
change
articular

cartilage

and

by factors
geometry,

polishing

such
alter

joints

are

and

as trauma,
synovial

to varying
the

degrees,

underlying

disease,
fluid, and

in-

weepThis
is
Bound-

no exception.

or attrition)
is inevitable,
greatly
minimizes
it. The

of articular

eroding,

synovial

The

lubrication
reciprocating

dicate
that synovial
joints
have a special
kind of fluid-film
lubrication
termed
ing
lubrication
(oozing
of synovial
fluid from
the articular
cartilage)
supplemented
by floating
lubrication,
that is, the classic,
fluid-film
type.
ary

have
lubrica-

of friction

surfaces

being
seem

sliding

through

of friction

is generally

The incongruity
synovial
fluid.

moves

develops
that is sufficient
to keep the
This film takes
up the effects
of friction.
synovial
folds aid in spreading
synovial

the coefficient

lubrication.
The
(articular
surfaces

is the

throughout
the whole
ment
of wedge-shaped

that

lubrication

even
most

with
com-

occasionally

bone.

Wear-and-

and biochemical
interfere
with

proccartilage

metabolism.
Absorption

from

Diffusion

phatic

Cavity

either
of the

in

capillaries

the other.
capillaries.
entrance

Joint

direction
synovial

True
solutions
When
colloidal
into

particles.
Small
are of a certain

the

synovial

colloidal
critical

by diffusion
into
capillaries.
Larger
The
rate
of
articular
pressure

that

takes
place
readily
between
the blood
membrane
on the one hand and the joint

are injected
solutions
or
membrane

into the joint

fine Suspensions
is inversely

particles
enter
both
size, probably
about

cavity
diffuse
are injected,

proportional

to

types
of capillaries.
that of the globulin

and lymcavity
on
rapidly
into
the rate
of

the

size

When
the
molecule,

of the

particles
removal

vessels
stops,
except
for small
quantities
that
enter
lymphatic
particles
remain
in the synovial
tissue.
diffusion
and absorption
of solutions
is increased
if the intrais increased,
either
by movement
or by injection
under
high pres-

sure.
THE

JOURNAL

OF

BONE

AND

JOINT

SURGERY

E.

Blood

The

arteries

most

that

supply

in the

prominent

venous

anastomoses

shunt

blood

unknown.

the

past

cellular

lying

also

the

capillary

little

joints

are

the

muscles

and

skin

spread

but

joint

nevertheless

is supplied

sensory

fibers

Section
joint
and

pain

by

that

move

in positions
the

more

or less

membrane.

that
of

blood

of

that

is

enable

specific

them

functions

flow

in

line

Arterio-

would

their

control

that

these

These
local

the

synovial

are

in joints.

there

may

responses,

numerous

can

bone

he

over-

reach

region

vertebral

of a

column

fibers
that
run
is also available

paili will
in articular
capsules

painful.

pain

with
for

capsule
Some
in

denervate
nerves
form

and

Studies

endings

not

ligaments,

of human

joints

is highly
sensitive,
of the nerve
fibers

the

adventitia

in

is of long

duration.

structure

may

addition

to

back

turn

may

of these

Joint

pain

may

be

the

pain

is severe

be. When

reflex

muscle

slowing
nerves

It often
leads
to reflex
contractions
lead to reflex atrophy
of antagonistic

spasm,

referred,

reflex

just

and

visceral

of the pulse
and
are postganglionic

as pain

sudden
and

fall of blood
sympathetic

from

in onset.,
circulatory

pressure.
fibers for the

of the smooth
muscle
in the articular
blood
vessels.
Some sensory
(mostly
pain)
in the adventitia
of the blood
vessels,
including

synovial

the

nerves

major

of the

joint

in

very

the fibrous
insensitive.

form

in the

Spasm

including
temporary
of the fibers in joint

Many

supply
endings

Each

joints

also receive
sympathetic
this perivascular
route

that
is relatively

especially

pain

supply

pain may arise in subchondral

cancellous
bone or marrow.
are especially
pertinent
to joint
pain : it. is usually
diffuse

muscles.

or visceral
be, in

that

in which

of a joint.

The

indicate

into

poorly

if the

region

branches.

most

membrane

and

deep

nerves

manner

nerve
for relief of arthritic
pain.
Many
of the fibers

are

anesthesia

Hence,
some joint
The following
points

muscles

constant

of which

vessels.

of

the

The

of branches
from
specific
nerves
and in the
Each major
articular
nerve
supplies
a wide-

major

the

endings

localized,

of

joint.

joints.

arteries

spasms

the

number
branches.
two

or stretching

accompany

to

branches

parts
of the limbs
the joint.
Possibly,

under

whereas

fibers form
those
in the

membrane.

The
endings

larger
and

are

ings

certain

fibers

in

joint

paciniform

sensitive
receptors

certain

range

receptors

to the
fire off,

of

stretch
rapidly

movement,

appear
adapt

The
movement

to

45-A,

NO.

5,

JULY

form

proprioceptive

the

joint
capsule
by movement.
and then
at

induced
at first,

After
the joint
then still others.
receptors

through

the

endings,

chiefly

Ruffini

and ligaments.
These
endWhen
a joint
is moved,
a rate that
depends
on the

has moved
Thus,
for
are

whole

more

a certain
distance,
other
a given
position
and for a
sensitive
than
others.
Some

range

of movement.

Many

recep-

continue
to discharge
after
movement
has stopped.
endings
in joints
are accurat.e
indicators
of position
and of
chief peripheral
contributors
to the kinesthetic
sense.
This

as muscle
from sensory
1963

in

certain

be active

slowly ; they
proprioceptive
and are the

also known
contribution

nerves

corpuscles,

speed
and extent
of movement.
receptors
are stimulated,
and

VOL.

arise

synovial

However,

about

relatively

from

endings.

opened

ally,

bone

of the

in joints

supplied

of a single
articular
may fail to relieve

twisting

sense,

adjacent

areas

networks.

is known

by at least

and the proximal

blood
vessels
to

tors

the

areolar

present

joints
is varied
both
in the
presence
or absence
of specific

any

and

Supply
Synovial

or

joint

and

are

Relatively

Nerve

the

1065

OF JOINTS

Most
of those
that
supply
the bone enter
the bone at or near the
attachment
and form a prominent
network
around
the joint.
arteries
that
supply
the joint
ultimately
form a capillary
network

capsular
The

PHYSIOLOGY

Supply

common.

to

I). GARDNER:

sense
or
endings

muscle-joint-tendon
iii muscles.

The

sense,
proprioceptive

receives
little,
if
endings
and

1066

A.A.O.S.

INSTRUCTIONAL

their
central
connections
movement
and posture.
Denervation
unless

of

there

muscles,

vell

Development

appear
usually

a widespread

be

does

concerned

not

lead

denervation

so that

as joints,

to

LECTURE

in the
to

movemeint.

regulation

a neuropathic

of a limb,

excessive

reflex

arthropathy

affecting
and

of

skin,

damage

bones,

and

to a joint

result..

of Joints

Most

studies

synovial

of the

mesenchymal

cells

Chondrification
future

areas

zones

becomes

menisci,

By

of the

termed

buds

joints

after

proliferate

have

limb

and

been

buds

form

concerned

appear

an axial

with

in the

mass

the

embryo,

termed

the

a blastema.

results
in the formation
of cartilage
models
of the
remains
between
the chondrifying
skeletal
elements
The mesenchyme
immediately
adjacent
to the interand most
intra-articular
structures,
such as ligaments,

interzones.

and

synovial

end

joints

of

Shortly

vascularized

the

hlastema

development

limbs.

in this blastema
blast.ema
that

The

hones.

forms

the

of

joints

adult

a joint

is also

as

also

COURSE

membrane,

of the
form

in

differentiate

embryonic
and

the

arrangement.

to

joints

occurs

during

the

having

a form

There
forms

is no recapitulation
in the
appear
as an intermediate

period

from

period,

develop
in situ and undergo
During
the latter
part

The
and

of about

it.

developing
change

arrangement

four

and

joints
from

similar

a half

to

to seven

sense that structures

phase.
Articular

those

weeks

resemble
the

adult

fertilization.

characteristic
structures
such

of the major
joints
form a joint
cavity.

cellular
in

after

no migration
having
phylogenetic
of the embryonic
period,
or early

minute
spaces
appear
in the interzones
mesenchyme.
These
spaces
coalesce
and
tion is probably
enzymatic.

closely

a generalized

of adult
lower
as ligaments

significance.
in the fetal

period,

and in the adjacent

The process
of cavita-

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