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The Structure and Development of Articular Cartilage: An In-depth Look, Lecture notes of Physiology

An in-depth exploration of the embryological development, gross structure, and microscopic features of articular cartilage. It discusses the role of Dr. Sydney M. Friedman's apparatus, the function of synovial membranes, and the cellular composition of cartilage. The text also covers the vascular supply, the role of minerals, and the metabolism of articular cartilage.

What you will learn

  • How does the vascular supply impact the development and function of articular cartilage?
  • What are the main components of the cellular composition of articular cartilage?
  • What role does Dr. Sydney M. Friedman's apparatus play in the development of articular cartilage?
  • How does the synovial membrane affect the function of articular cartilage?
  • What minerals are essential for the proper development and function of articular cartilage?

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THE
STRUCTURE
AND FUNCTION OF SYNOVIAL JOINTS,
WITH
PARTICULAR REFERENCE TO THE
MECHANISM
OF THEIR LUBRICATION
by
MICHAEL STAFFORD
PIPER
M.D.,
University
of
British
Columbia,
1968
A THESIS
SUBMITTED
IN PARTIAL FULFILMENT OF
THE
REQUIREMENTS
FOR THE
DEGREE
OF
MASTER
OF SCIENCE
in
the
Department
of
Anatomy
We
accept
this
thesis
as
confirming
to the
required
standard
THE UNIVERSITY OF BRITISH
COLUMBIA
JUNE, 1972
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THE STRUCTURE AND FUNCTION OF SYNOVIAL JOINTS, WITH PARTICULAR REFERENCE TO THE MECHANISM OF THEIR LUBRICATION

by

MICHAEL STAFFORD PIPER M.D., U n i v e r s i t y of B r i t i s h Columbia, 1968

A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE

i n the Department o f Anatomy

We a c c e p t t h i s t h e s i s as c o n f i r m i n g t o the r e q u i r e d s t a n d a r d

THE UNIVERSITY OF BRITISH COLUMBIA JUNE, 1972

In p r e s e n t i n g t h i s t h e s i s in p a r t i a l f u l f i l m e n t o f the requirements f o r an advanced degree at the U n i v e r s i t y of B r i t i s h Columbia, I agree that the L i b r a r y s h a l l make it f r e e l y a v a i l a b l e f o r reference and s t u d y. I f u r t h e r agree t h a t p e r m i s s i o n f o r e x t e n s i v e copying o f t h i s t h e s i s f o r s c h o l a r l y purposes may be granted by the Head o f my Department o r by h i s r e p r e s e n t a t i v e s. It i s understood that copying o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l gain s h a l l not be allowed without my w r i t t e n p e r m i s s i o n.

The U n i v e r s i t y o f B r i t i s h Colui Vancouver 8, Canada

Department of

( i i i ) As a r e s u l t o f t h e s e i n v e s t i g a t i o n s , i t was f o u n d t h a t t h e s y n o v i a l f l u i d s a m p l e s f r o m p a t i e n t s w i t h r h e u m a t o i d a r t h r i t i s c o n t a i n e d a s i g n i f i c a n t l y l o w e r c o n c e n t r a t i o n o f i o n i z e d s o d i u m. I t i s c o n c l u d e d t h a t t h e l o w e r c o n c e n t r a t i o n o f s o d i u m i o n s i n s y n o v i a l f l u i d o f r h e u m a t o i d a r t h r i t i s may r e s u l t i n a d i m i n u t i o n o f e l e c t r i c a l r e p u l s i v e f o r c e s a c t i n g w i t h i n s y n o v i a l j o i n t s , a n d e x p l a i n , i n p a r t , t h e c a r t i l a g e a t t r i t i o n s e e n i n t h i s d i s e a s e.

S y d n e y M. F r i e d m a n P r o f e s s o r a n d H e a d D e p a r t m e n t o f Anatomy U n i v e r s i t y o f B r i t i s h C o l u m b i a ( S u p e r v i s o r )

( i v ) TABLE OF CONTENTS THE STRUCTURE AND FUNCTION OF SYNOVIAL JOINTS, WITH PARTICULAR REFERENCE TO THEIR MECHANISM OF LUBRICATION. Page No. I n t r o d u c t i o n 1 Embryology of S y n o v i a l J o i n t s 4 Gross Anatomy o f S y n o v i a l J o i n t s 6 B l o o d and Nerve Supply o f S y n o v i a l J o i n t s 8 L i g h t and E l e c t r o n M i c r o s c o p y of S y n o v i a l J o i n t s 13 S y n o v i a l Membrane and S y n o v i a l F l u i d 18 B i o c h e m i s t r y o f A r t i c u l a r C a r t i l a g e 25 M e t a b o l i s m o f A r t i c u l a r C a r t i l a g e 28 L u b r i c a t i o n o f S y n o v i a l J o i n t s 30 C o n c l u s i o n " 37 B i b l i o g r a p h y 39

( v i )

ACKNOWLEDGEMENT

I w o u l d l i k e t o a c k n o w l e d g e t h e a s s i s t a n c e g i v e n me i n t h e p r e p a r a - t i o n o f t h i s t h e s i s. I n p a r t i c u l a r I w o u l d l i k e t o t h a n k t h e o r t h o p a e d i c s u r g e o n s a n d r a d i o l o g i s t s a t t h e V a n c o u v e r G e n e r a l H o s p i t a l f o r t h e i r a s s i s t a n c e i n o b t a i n i n g s y n o v i a l f l u i d s a m p l e s , a n d D r. G e r f r i e d G e b e r t f o r h i s t e c h n i c a l a s s i s t a n c e. I n a d d i t i o n , a p p r e c i a t i o n i s e x p r e s s e d t o D r. S y d n e y M. F r i e d m a n f o r t h e o p p o r t u n i t y t o c o n d u c t t h i s w o r k , a n d f o r h i s d i r e c t i o n a n d s u p p o r t.

THE STRUCTURE AND FUNCTION OF SYNOVIAL JOINTS, WITH PARTICULAR REFERENCE

TO THE MECHANISM OF THEIR LUBRICATION

X INTRODUCTION "The F a b r i c o f the J o i n t s i n the Human Body i s a S u b j e c t so much the more e n t e r t a i n i n g , as i t must s t r i k e everyone t h a t c o n s i d e r s i t a t t e n t i v e - l y w i t h an Idea o f f i n e m e c h a n i c a l C o m p o s i t i o n. Where-ever the M o t i o n o f one Bone upon a n o t h e r i s r e q u i s i t e , t h e r e we f i n d an e x c e l l e n t Apparatus f o r r e n d e r i n g t h a t M o t i o n s a f e and f r e e : We s e e , f o r I n s t a n c e , the e x t r e m i t y o f one Bone moulded i n t o an o r b i c u l a r C a v i t y , t o r e c e i v e the Head o f a n o t h e r , i n o r d e r t o a f f o r d i t an e x t e n s i v e P l a y. Both a r e c o v e r e d w i t h a smooth e l a s t i c C r u s t , t o p r e v e n t mutual A b r a s i o n ; c o n n e c t e d w i t h s t r o n g L i g a m e n t s , t o p r e v e n t D i s l o c a t i o n ; and e n c l o s e d i n a Bag t h a t c o n - t a i n s a p r o p e r F l u i d d e p o s i t e d t h e r e , f o r l u b r i c a t i n g the Two c o n t i g u o u s S u r f a c e s. So much i n g e n e r a l. " thus w r o t e W i l l i a m H u n t e r , i n 1743 i n a r e m a r k a b l e e s s a y ; "Of the S t r u c t u r e and D i s e a s e s o f A r t i c u l a t i n g C a r t i l a g e s ( 1 ) ". H i s o b s e r v a t i o n s , r e c o r d e d over two hundred y e a r s ago, were e x t r e m e l y a c u t e ; i n f a c t , h i s d e s c r i p t i o n o f t h e b l o o d s u p p l y o f j o i n t s , the " C i r c u l u s A r t i c u l i V a s c u l o s u s " has p e r s i s t e d t o the p r e s e n t t i m e. That the s y n o v i a l j o i n t s c o m p r i s e d a u n i q u e body t i s s u e e a s i l y a c c e s s i b l e t o i n v e s t i g a t i o n l e d e a r l y a n a t o m i s t s t o t h e i r s t u d y. Thus, w i t h the i n t r o d u c t i o n o f the m i c r o s c o p e , the u n i q u e h i s t o l o g i c a l a p p e a r - ance of the a r t i c u l a r c a r t i l a g e s was e s t a b l i s h e d , and c a r t i l a g e was known t o have a h i g h m a t r i x t o c e l l r a t i o ( 2 ). Much i n f o r m a t i o n on the n a t u r e of s y n o v i a l j o i n t s has been accumu- l a t e d i n r e c e n t y e a r s , and i t i s the purpose o f t h i s paper t o r e v i e w

c o n t r o l l e d p a i n l e s s m o t i o n o f t h e l i m b s. To t h i s end, n a t u r e h a s p r o - v i d e d m u s c l e s a n d g r a v i t y f o r m o t i o n a n d l i g a m e n t s a n d a r c h i t e c t u r a l d e s i g n f o r s t a b i l i t y. H o w e v e r , i t i s t h e m e c h a n i s m o f l u b r i c a t i o n i n s y n o v i a l j o i n t s t h a t h a s p r o v i d e d t h e g r e a t e s t i n t e r e s t as w e l l as t h e g r e a t e s t s o u r c e o f c o n t r o v e r s e y i n r e c e n t y e a r s. T h i s t h e s i s c o n c l u d e s w i t h a d i s c u s s i o n o f t h e v a r i o u s m e c h a n i s m s p r o p o s e d t o a c c o u n t f o r t h i s l u b r i c a t i o n ; a n d , as a r e s u l t o f r e s e a r c h c o n d u c t e d t h i s y e a r , a t h e o r y o f l u b r i c a t i o n e n h a n c e m e n t b y m o n o v a l e n t c a t i o n s w i t h i n s y n o v i a l f l u i d.

  • 4 -

Embryology of S y n o v i a l J o i n t s

Fore and h i n d limb buds appear i n the human embryo at f o u r weeks of

age as outgrowths of p r i m i t i v e mesenchyme. (5, 6, 7) W i t h i n a few days,

a c e n t r a l core of mesenchyme condenses t o become what i s termed the

"blastema". The blastema subsequently becomes f u r t h e r condensed i n t o

c h o n d r i f i c a t i o n c e n t e r s , one f o r each s k e l e t a l element. That p a r t of the

blastema which remains f o l l o w i n g segmentation i s c a l l e d the " i n t e r z o n e. "

These are l o c a t e d at the s i t e s o f f u t u r e j o i n t s , and are continuous w i t h

the perichondrium surrounding the c a r t i l a g i n o u s models of the bones. (8)

At about f i v e and one h a l f weeks (embryo 11-17 mm) the i n t e r z o n e has

f u r t h e r d i f f e r e n t i a t e d i n t o a three l a y e r e d s t r u c t u r e w i t h two chondro-

genic l a y e r s sandwiching a loose middle l a y e r. (9) The chondrogenic l a y e r s

are continuous w i t h the perichondrium of the neighbouring s k e l e t a l seg-

ment. The i n t e r m e d i a t e l a y e r i s continuous w i t h the adjacent e x t r a - >

b l a s t e m a l mesenchyme, which has become v a s c u l a r i z e d at t h i s stage and i s

r e f e r r e d t o as the " s y n o v i a l mesenchyme", ( s i x and one h a l f weeks - 25 mm).

At t h i s stage the outer l a y e r s of the s y n o v i a l mesenchyme have begun

t o d i f f e r e n t i a t e i n t o the f i b r o u s capsule, w h i l e deeper l a y e r s are

d i f f e r e n t i a t i n g i n t o s y n o v i a l membrane and the i n t r a c a p s u l a r s t r u c t u r e s.

Most authors agree t h a t the s y n o v i a l space begins t o develop at about

the s i x week stage of growth as minute spaces i n the s y n o v i a l mesenchyme

and the loose middle l a y e r of the i n t e r z o n e. These s m a l l spaces subse-

q u e n t l y coalese to form the j o i n t c a v i t y. (6, 8, 11, 12) However, there

i s some d i s p u t e as t o how c a v i t a t i o n a c t u a l l y comes about. Some f e e l that

c a v i t a t i o n i s an a c t i v e process accomplished by c e l l u l a r p r o l i f e r a t i o n o f.

Gross Anatomy o f S y n o v i a l J o i n t s A j o i n t or a r t i c u l a t i o n i s formed where two or more bones o f the body meet one a n o t h e r. ( 1 5 , 16, 17) The c h a r a c t e r and s t r u c t u r e of j o i n t s a r e l a r g e l y d e t e r m i n e d by t h e i r f u n c t i o n. Thus, j o i n t s may be f i b r o u s as seen i n t h e s u t u r e s between c r a n i a l bones, or the syndesmosis o f the i n f e r i o r t i b i o f i b u l a r j o i n t. S e c o n d l y , j o i n t s may be c a r t i l a g i n o u s. P r i m a r y c a r t i l a g i n o u s j o i n t s e x i s t as the e p i p h y s e a l p l a t e between t h e e p i p h y s i s and d i a p h y s i s o f growing bones. As such they a r e temporary and e v e n t u a l l y o s s i f y. Secondary c a r t i l a g i n o u s j o i n t s a r e seen where the a r t i c u l a r s u r f a c e s of the bones a r e c o v e r e d w i t h h y a l i n e c a r t i l a g e and c o n n e c t e d by a d i s c o f f i b r o c a r t i l a g e. These j o i n t s a l l o w some degree of m o t i o n , and a r e found i n the median p l a n e of the body - the s t e r n o - m a n u b r i a l and i n t e r v e r t e b r a l j o i n t s and the p u b i c s y m p h y s i s. A l l o t h e r j o i n t s i n the body a r e r e f e r r e d t o as s y n o v i a l or d i a r - t h r o d i a l j o i n t s , and have common c h a r a c t e r i s t i c s. The c o n t i g u o u s bony s u r f a c e s a r e c o v e r e d w i t h a r t i c u l a r c a r t i l a g e and a r e n o t a t t a c h e d t o one a n o t h e r. The a r t i c u l a r c a r t i l a g e i s u s u a l l y h y a l i n e , however, i n the case o f membranous bone the a r t i c u l a r c a r t i l a g e i s w h i t e f i b r o c a r t i l a g e. The b a s a l l a y e r o f t h e a r t i c u l a r c a r t i l a g e i s c a l c i f i e d and f i r m l y bound t o the u n d e r l y i n g bone, the " a r t i c u l a r l a m e l l a. " The bones o f s y n o v i a l a r t i c u l a t i o n s a r e bound t o one a n o t h e r by a f i b r o u s c a p s u l e c o n s i s t i n g of a dense c o n n e c t i v e t i s s u e c u f f. G e n e r a l l y the c a p s u l e i s a t t a c h e d t o the bones n e a r the p e r i p h e r y o f the a r t i c u l a r s u r f a c e. A v a r i a b l e number o f t h i c k e n i n g s i n the c a p s u l e a r e s e e n. These l i g a m e n t s a c t t o p r e v e n t e x c e s s i v e or abnormal movements o f the j o i n t s. L i n i n g the i n n e r s u r f a c e o f the f i b r o u s c a p s u l e and a t t a c h i n g t o the a r t i c u l a r margins i s a s y n o v i a l membrane. T h i s h i g h l y s p e c i a l i z e d s t r u c t u r e

e n c l o s e s a p o t e n t i a l s p a c e ( t h e t r u e j o i n t s p a c e ) w i t h i n w h i c h i s a s m a l l amount o f " S y n o v i a " o r " S y n o v i a l F l u i d ". The j o i n t c a v i t y may be d i v i d e d , c o m p l e t e l y o r i n c o m p l e t e l y , by a f i b r o c a r t i l a g i n o u s a r t i c u l a r d i s c. T h e s e d i s c s a r e u s u a l l y a t t a c h e d t o t h e c a p s u l e a t t h e i r p e r i p h e r y , b u t a r e n o t c o v e r e d w i t h s y n o v i a l mem- b r a n e. S y n o v i a l j o i n t s a r e f u r t h e r c l a s s i f i e d a c c o r d i n g t o t h e k i n d o f m o t i o n p e r m i t t e d i n them. T h i s i s d e t e r m i n e d b y t h e s h a p e o f t h e a r t i c u l a r s u r f a c e s , t h e m u s c l e s t h a t a c t u p o n t h e j o i n t , a n d t h e l i m i t s p l a c e d o n t h e j o i n t s ' e x c u r s i o n b y t h e c a p s u l e a n d l i g a m e n t s.

The b l o o d s u p p l y o f the m e t a p h y s i s changes between f o e t a l and a d u l t l i f e. (19j 20, 21) I n the f o e t u s , the metaphysis i s s u p p l i e d by the n u t r i e n t a r t e r y a l o n e , w h i l e i n the a d u l t , the m e t a p h y s e a l a r t e r i e s s u p p l y the p e r i p h e r a l two f i f t h s of the m e t a p h y s i s , the c e n t r a l t h r e e f i f t h s s t i l l b e i n g s u p p l i e d by the n u t r i e n t a r t e r y. (22) T h i s s i t u a t i o n e x i s t s i n the " g r o w i n g end" o f l o n g bones. I n the " n o n - g r o w i n g e n d " e v e n t u a l l y a l l b l o o d d e r i v e s from the m e t a p h y s e a l a r t e r i e s. L i k e w i s e t h e r e i s a change i n the b l o o d s u p p l y of e p i p h y s e s. I n the i n f a n t , (up t o one y e a r ) , m e t a p h y s e a l v e s s e l s p i e r c e the c a r t i l a g i n o u s e p i p h y s e a l p l a t e and thus s u p p l y the b a s a l p a r t of the e p i p h y s i s. A f t e r one y e a r of age the e p i p h y s e a l p l a t e becomes i m p e r v i o u s t o these meta- p h y s e a l v e s s e l s , and i s s u p p l i e d s o l e l y by the e p i p h y s e a l v e s s e l s. (23) Thus i t can be seen t h a t the b l o o d s u p p l y o f the e p i p h y s e a l p a r t of l o n g bones, and hence t h e s u b c h o n d r a l r e g i o n i s o f v a r i a b l e o r i g i n , depending upon the age o f the bone and the s i t e o f s t u d y. T h i s f a c t may be of r e l e v a n c e i n c o n s i d e r i n g the e t i o l o g y of a v a s c u l a r n e c r o s i s of the c a p i t a l f e m o r a l e p i p h y s i s. (19) The r o l e o f the s u b c h o n d r a l e p i p h y s e a l v e s s e l s i n the n u t r i t i o n o f a r t i c u l a r c a r t i l a g e w i l l be d i s c u s s e d l a t e r , however, the v a s c u l a r anatomy of the e p i p h y s i s now deserves m e n t i o n. Many e p i p h y s e a l n u t r i e n t a r t e r i e s a r i s e from the c i r c u l u s a r t i c u l i v a s c u l a s i s and anastomose w i t h i n the e p i p h y s e s of l o n g bones. I n the immature s k e l e t o n some branches pass d i r e c t l y t o the s u b c h o n d r a l a r e a o f the c a r t i l a g i n o u s growth. p l a t e ( e p i p h y s e a l p l a t e ) , but the m a j o r i t y o f the e p i p h y s e a l a r t e r i e s pass towards the a r t i c u l a r and n o n - a r t i c u l a r s u r f a c e s of the e p i p h y s i s. As these e p i p h y s e a l n u t r i e n t a r t e r i e s pass from t h a t p a r t o f the e p i p h y s i s a d j a c e n t t o the m e t a p h y s i s towards the a r t i c u l a r s u r f a c e , they

  • 10 - f e e d i n t o a number o f a r t e r i a l arcades w h i c h s u b s e q u e n t l y g i v e r i s e t o f u r t h e r r a d i a t i n g a r t e r i e s. (15) T h i s p a t t e r n of the a r t e r i a l s u p p l y o f t h e e p i p h y s i s i s s i m i l a r t o the p a t t e r n o f t h e v a s c u l a r s u p p l y seen i n the c a r t i l a g i n o u s e p i p h y s e a l p r e c u r s o r. (24) The a n a l o g y between a v a s - c u l a r n e c r o s i s o f the f e m o r a l head i n a d u l t s (25) and t h a t seen f o l l o w i n g r e d u c t i o n o f a c o n g e n i t a l l y d i s l o c a t e d h i p (26) i s e a s i l y s e e n. I t has been n o t e d t h a t the d i s t r i b u t i o n o f e p i p h y s e a l a r t e r i e s i s n o t r e l a t e d t o t h e t r a b e c u l a r a r c h i t e c t u r e o f the bone. (27) The p a t t e r n o f «» t h e t r a b e c u l a e i s known t o be v a r i a b l e depending upon the b i o m e c h a n i c a l f o r c e s o p e r a t i v e i n v a r i o u s c o n d i t i o n s. (28) However, the d i s t r i b u t i o n o f e p i p h y s e a l a r t e r i e s appears r e m a r k a b l y c o n s t a n t. The t e r m i n a l e p i p h y s e a l a r t e r i o l e s w h i c h emanate from the a n a s t o - m o t i c a r c a d e s i n the c e n t r a l p a r t of t h e e p i p h y s i s r a d i a t e towards the s u b c h o n d r a l bony p l a t e w h i c h s u p p o r t s the c a r t i l a g e. Here they form s i n u s o i d a l l o o p s a b u t i n g a g a i n s t the b a s a l c a r t i l a g e l a y e r s. These a r e s i m p l e s i n u s o i d s not a t a l l l i k e the d i l a t e d , v a r i c o s e s i n u s e s seen on t h e m e t a p h y s i a l s i d e o f the growth p l a t e. (23) O c c a s i o n a l l y t h e s e s u b - c h o n d r a l l o o p s can be seen t o r e a c h i n t o t h e c a l c i f i e d zone o f c a r t i l a g e , but o n l y v e r y r a r e l y i n t o the more s u p e r f i c i a l zones o f u n c a l c i f i e d c a r t i l a g e. (4) The s i n u s o i d a l network i n the immediate s u b c h o n d r a l a r e a o f the e p i p h y s i s tends t o be p a r a l l e l t o the a r t i c u l a r c a r t i l a g e. T h i s v a s c u l a r anastomosis i s s e p a r a t e d from the most b a s a l l a y e r o f the c a r t i l a g e ( t h e c a l c i f i e d zone o f c a r t i l a g e ) by a t h i n p l a t e o f bone. As has been men- t i o n e d t h i s bony p l a t e i s o c c a s i o n a l l y seen t o be p e r f o r a t e d by s m a l l v a s c u l a r c h a n n e l s w h i c h extend i n t o the c a l c i f i e d zone of the a r t i c u l a r c a r t i l a g e. T h i s f i n d i n g l e n d s c r e d e n c e t o the s u g g e s t i o n t h a t the a r t i c u l a r c a r t i l a g e d e r i v e s a t l e a s t p a r t o f i t s n u t r i t i o n from the sub-

Many o f t h e f i b e r s i n a r t i c u l a r n e r v e s a r e v a s o m o t o r a n d v a s o s e n s o r y. T h e s e s u p p l y t h e b l o o d v e s s e l s i n t h e f i b r o u s c a p s u l e a n d s y n o v i a l mem- b r a n e , a n d t h e r e b y c o n t r o l b l o o d f l o w t h r o u g h t h e j o i n t. The e l e v a t e d t e m p e r a t u r e o f t h e o v e r l y i n g s k i n i n d i s t e n d e d p a t h o l o g i c a l j o i n t s i s e v i d e n c e o f t h e common s u p p l y o f j o i n t s a n d o v e r l y i n g s k i n. ( 3 2 ) One c h a r a c t e r i s t i c o f t h e n e r v e s u p p l y t o s y n o v i a l j o i n t s i s o v e r l a p. T h a t i s , e a c h r e g i o n o f a j o i n t i n n e r v a t e d b y one n e r v e i s s u p p l i e d b y a t l e a s t one o t h e r a r t i c u l a r n e r v e. T h i s p a t t e r n o f o v e r l a p p i n g i n n e r v a t i o n may l e a d t o f a i l u r e o f p r o c e d u r e s d e s i g n e d t o d e n e r v a t e p a i n f u l a r t h r i t i c j o i n t s.

  • 13 -

L i g h t and E l e c t r o n M i c r o s c o p y o f A r t i c u l a r C a r t i l a g e

I n r o u t i n e h i s t o l o g i c a l p r e p a r a t i o n s o f a r t i c u l a r c a r t i l a g e s chon- d r o c y t e s a r e seen t o l i e i n l a c u n a e d i s t r i b u t e d w i t h i n an abundant h y a l i n e i n t r a c e l l u l a r m a t r i x. (33) The c h o n d r o c y t e s a r e s p a r c e , even i n v e r y young c a r t i l a g e. ( 3 4 , 35) E a r l y m i c r o s c o p i s t s d e f i n e d f o u r d i s t i n c t zones i n a d u l t a r t i c u l a r c a r t i l a g e , and d e s c r i b e d d i f f e r e n c e s i n the s p a t i a l arrangement as w e l l as the c e l l u l a r morphology i n t h e s e z o n e s. (34) Zone I , the s u p e r f i c i a l or t a n g e n t i a l s t r a t u m c o n s i s t s o f f l a t t e n e d o r markedly o v o i d c e l l s l y i n g a d j a c e n t t o the s u r f a c e w i t h the l o n g a x i s o f the c e l l s p a r a l l e l t o the a r t i c u l a r m a r g i n. T h i s a r e a has a l s o been r e f e r r e d t o as the " g l i d i n g zone". (36) I n Zone I I , the i n t e r m e d i a t e or t r a n s i t i o n a l s t r a t u m , the c h o n d r o c y t e s a r e plump and o v a l and d i s t r i b u t e d i n a random f a s h i o n. Zone I I I i s r e f e r r e d t o as t h e deep o r r a d i a t e s t r a t u m. The c e l l s i n t h i s zone t e n d t o be s m a l l and r o u n d. They a r e a r r a n g e d i n s h o r t , i r r e g u l a r columns p e r p e n d i c u l a r t o the a r t i c u l a r s u r f a c e. The b a s a l l a y e r o r Zone IV i s the c a l c i f i e d s t r a t u m , and l i e s a d j a c e n t t o the s u b c h o n d r a l bony p l a t e. I n t h i s a r e a the c e l l s t e n d t o be s m a l l and i r r e g u l a r w i t h p y k n o t i c n u c l e i. On h e m a t o x y l i n and e o s i n s t a i n the c a l c i f i e d s t r a t u m i s s e p a r a t e d from t h e more s u p e r f i c i a l l a y e r s by a t h i n wavy b l u e l i n e. The n a t u r e o f t h i s b a s o p h i l i c a r e a i s unknown; i t has been c a l l e d the " t i d e m a r k ". F u r t h e r mention w i l l be made o f the v a r i a t i o n s found i n the c h o n d r o c y t e s of d i f f e r - ent zones, p a r t i c u l a r l y w i t h r e g a r d t o s t u d i e s w i t h the e l e c t r o n m i c r o s c o p e. M e n t i o n s h o u l d be made now, however, o f the l i g h t m i c r o s c o p i c appearance of t h e i n t r a c e l l u l a r m a t r i x o f a r t i c u l a r c a r t i l a g e.