COMPARATIVE
OF

HISTOLOGY OF

THE

KIDNEY

DOMESTIC ANIMALS

By
RAMCHANDRA PRASAD YADAVA

A THESIS
Submitted to the

School

of Advanced Gradua te

Michigan State U n i v e r s i t y
Science

in p a r t i a l

of A g r i c u l t u r e

ful fi llment

f or

DOCTOR

the

of the

d e gre e

OF

PHILOSOPHY

1955

and Applied

requirements

of

D e p a r t m e n t of Anatomy

Studies

of

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ABSTRACT

This
plete

study was

and u p - t o - d a t e

u n d e r t a k e n to p rovi de

d e s c r i p t i o n of the

the kidney of the d o m e s t i c
nine

domestic

c h os e n as

animals

ent.

The

outer layer
and a few

by loos e

collagenous
in all

a lesser

elastic
and

(156|j.), dog

animals

where

and the

except the

fibers,
cat.

com­

structures

of

ma d e

on sixty-

the pig being

corpuscles

sm aller

respectively.

only one l a y e r

renal

than the

in ne r l a y e r
with

in the ox (173p),
cortical

ones,

was

pres­
col­

was f o r m e d

smooth m u s c l e f i b e r s
sheep and goat,
were

and

v ery n u m e r o u s

layer.

corpuscles

(106p) w e r e

an i m a l s

c o n s i s t e d of dense

In the

a d is t i n c t m u s c u l a r

and c a t

in domes tic

the m u s c l e f i b e r s

1 7 8 i j l , 1 3 7 (j l , 122(x, and 96p,

which w e r e
renal

fibers

extent in the ox,

(124p),

cat,

reticular

The j u xt a me du l l a r y

were

species,

of the tunica f i b r o s a

and a p p e a r e d to f o r m

lary

were

of the kidney c a ps u l e

except in the

lagenous

to

Studies

or l e s s

a type.

two-layered

present

microscopic

of seven di f fe r en t

The tunica f i b r o s a
was

animals.

a more

in the h o r s e

larger

than the

(191p),

cortical

pig
ones,

respectively.

The ju xt a m e d u l -

sheep

and goat

(147|a),

which w e r e

The j u x t a g l o m e r u l i w e r e l a r g e r

lBlp,

(15 7(jl

1S3pu, and

than the c o r t i c a l

)

158(ji,

ones

in the pig,
true.

horse,

In the

betwee n the
cortical

dog,

sheep
size

and cat;

and goat no

arteriole

along with the

As

usual
ce lls

ce lls

reverse

d iffe r e n c e

glomeruli

was

was

noticed

and th os e

(48(jl);

equal

was

in the

of the

were

more

(45p),

greater

sheep

in all

animals

of the

the

goat and

than in the pig.

diameter

of the p r o x i ­

(56p),

and s m a l l e r

some

afferent

In the

numerous

in the h o r s e

(45p);

contained

m e d i a of the

myoepitheloid c e l l s .

c o m p a r e d with the pig

convoluted tubule

and ca t

apparatus

smooth m u s c l e

the j u x t a g l o m e r u l a r

goat

ox the

ones.

spindle-shaped

mal

in the

appreciable

of the j u x t a m e d u l l a r y

The j u x t a g l o m e r u l a r

dog,

whereas

ox

in the

(50p),

dog

and

(39p)

(41 p).
The b r u s h b o r d e r

individual

cell

was found to f o r m

in the p r o x i m a l

tubule

of the pig kidney,

other

animals

they w e r e

uniformly

arranged

of the

cells.

The b a s a l

striations

were

F a t globules

were

a cluster from

along the

more

o b s e r v e d in the p r o x i m a l

each

whereas

in

luminal b o r d e r

d i s t i n c t in the dog.

tubule

of the

cat

and

dog.
The

m ac ula e

in the h o r s e ,

in all

an i m a l s

were

in which they w e r e found to be

The p a p i l l a r y
epithelium

densa e

in all

duct was

animals

lined by both

except the

dog,

single-layered

except

stratified.
s imple

and t r a n s i t i o n a l

in which the

papillary

duct

was

lined by

illary

simple

columnar

epithelium

only.

observed

in all

duct on the p a p i l l a w e r e
The

in terstitial

spaces

of the kidney

a few collagenous f i b e r s ,

fibroblasts,

all

dog and cat,

animals

except

in the

Openings
animals

contained

histiocytes,
w he r e

of the

studied.

reticular fibers,

and m a s t

mast

pap­

cells

cells

were

in

ab -

s ent.
The
in the
cl o s e

interstitial

These

in all

cell

pig,

groups
The

cell

spa ce s

groups

in the

groups

and l e s s
near

cells

the

The

macula

arteries

w er e

polygonal

ox than th os e

cell
space

or

are

between the
an i m a l s

contrary

and

in the

than in

goat,

ox and h o r s e
and dog.

than those lo c a t e d

were

The

elsewhere.

eosinophilic

larger

cell

cytoplasm

and m o r e

eosino­

of the pig.

The following findings
time,

arteries,

absent.

unit of Goormaghtigh was found to be

d e n s a in all

in the

in which they w e r e

with faintly
cells

were p re s e n t

especially

the c o r t i c a l

sheep,

larger

The

region,

cat,

numero us

in the

stained nuclei.

in the

triangular

n um e r o u s

B e c h e r ’s ce lls

ca p s u l e ,

except the

were m ore

were mostly

and darkly
philic

a n i m al s

or

cortical

neighborhood of Bo wm a n ' s

arterioles

the

intertubular

to the

aff e r e n t

and

without any
are

efferent
species

thought to be

work

iv

of

s ituated in the

arterioles,

and the

var ia tion.
reported for

recent investigators:

the f i r s t

1.
pig,

Smooth m u s c l e

in the

c a p s u l e of the h o r s e ,

goat,

and dog.
2.

goat,

Intertubular or

pig,

species

and dog,

ox,

Becher's

A. s t r a t i f i e d m a c u l a

4.

Ma s t

ce lls

in the

except the dog
Larger

sheep,
6.

and equal
7.

8.

9.

dens a in the

interstitial

cortical

horse.

s p a ce s

of the kidney of all

than j u x t a m e d u l l a r y

renal

cortical

than j u x t a m e d u l l a r y

glomeruli

slightly l a r g e r

tubule

ones

ducts

in the

in c l u s t e r s

corpuscles

in

sheep

at the

in the

ox,

and goat.

apices

of the c e l l s

of

in the pig.

T r a n s i t i o n a l epithelium

the p a p i l l a r y

ox, sheep,

and cat.

Brush b o rde rs

the p r o x i m a l

in the h o r s e ,

and goat.

Larger
or

cells

but ab s e n t in the cat.

3.

5.
the

ce lls

of all

Papillary

animals

duct openings

extending
except the

varying d i s t a n c e s

into

dog.

on the papilla of all

animals

in­

v es t i g at e d .
10.
inv e s t i g a t i o n ,

The

cell

unit of Goormaghtigh in all

the

spe ci es

in this

FRONTISPIECE

Photomicrograph of afferent
puscle

(3), unusual

tigh (5), Be ch er 's

(1) and efferent a r t e r i o l e s

double maculae densae
cells

and the distal tubule

(8).

(6), proximal
H. and E.

renal

cor­

(4), cell unit of Goormagh-

tubule with basal
420x.

(Z),

striations

Six-year-old cow.

(7),

*

f d f - ’ '.

&

\
}

kj

*

f
FRO NTISPIECE

8

£
£

FRONTISPIECE

Photomicrograph of afferent (1) and efferent a r t er io les
puscle

(3), unusual double maculae densae

tigh (5), Be ch er's

cells

H. and E.

420x.

striations

Six-year-old cow.

d

Y

renal c o r ­

(4), cell unit of Goormagh-

(6), proximal tubule with basal

and the distal tubule (8).

(2),

(7),

FRO NTISPIECE

Dedicated to my P r o f e s s o r

Dr.

M.

L.

Calhoun, D.V.M.,

P r o f e s s o r and Head,
omy,

College

of

research,

rich

human

never-ending

P h.D.,

D e p a r t m e n t of A na t­
Veterinary

Michigan State Uni ver s it y,
to

M.S.,

scientific

whose

devotion

achievements,

understanding
source

Medicine,

have

been

of i n s pi r a t i o n.

viii

and
a

ACKNOWLEDGMENTS

The

author

c i ati on to Dr.

M.

wishes
Lois

of this

to Dr.

J.

his

sincere

Professor

patie nc e,

inve s tiga tio n

Frederick

express

Calhoun,

m e n t of Anatomy, f o r h e r
course

to

and a p p r e ­

and Head of the D e p a r t ­

guidance,

and advice

and the p r e p a r a t i o n

Smithcors,

gr a t i t u de

of the

during

the

manuscript;

D e p a r t m e n t of Anatomy,

who gave

so

*

generously
helpful

of his

valuable

suggestions;

and to Dr.

f or

her

valuable

for

her

help in taking

Mrs.

Ann L.

ful

and

in reading
Esther

M.

the m a n u s c r i p t and giving
Smith, Anatomy D ep a r t m e n t ,

in the p r e p a r a t i o n of the m a n u s c r i p t

the p h o t o m i c r o g r a p h s .
the

department

Thanks

secretary,

are

also

and

due to

and o t h e r m e m b e r s

staff of the D e p a r t m e n t of Anatomy, f o r

their

help­

cooperation.
Sincere

fessor
tive

suggestions

Br o ok s,

of the faculty

time

a p p r e c i a t i o n is

of Physiology

criticism

of S u r g e r y
The

and P h a r m a c o l o g y , f o r

of the m a n u s c r i p t ,

and Medicine,
author

s pe ct to the l a t e
ology,

wishes

Dr.

and a m e m b e r

valuable help

extended to Dr.

and

f or his

and Dr.
helpful

to e x p r e s s

Frank

Thorp,

of the

author's

s uggestions.

ix

his

Jr.,

L.

his

W.

O.

F,

efforts

sincere
Professor

Pro­

and c o n s t r u c ­

Brinker,

suggestions

guidance

Wolterink,

Professor

and a s s i s t a n c e .

g ra t i t u d e

and

of Animal

committee, for

re­

Pathhis

For
author

is

Ms

aid in

deeply

m e n t of Animal

Husbandry.

D. Baten,

t o w ar ds

the

to Dr.
for

their

ment

author
L.

is

unusual

technique.
Singh,

their

G erman

presses

his

help,

Hans

suggestions

also

their

J.

Douglass

Richare

the

and Mr s .

and

equip­

Errol

in histological

Moore,

invaluable

Beck

and Medicine,

to use

and a s s i s t a n c e

due to Mr.

Ruhland f or

valuable help

Clifford C.

and p e r m i s s i o n

Clyde

are

his

of S ur ge ry

are

gr a t i t u de

sincere

to Dr.

Medicine,

Dr.

assistance

Professor

of S ur ge r y

as s i s t a n t s h i p

enabled him

F.

Clark,

Edwa r d K.

Chintamani
in many

Professor

and to Dr.

and Medicine, f o r

the

providing financial

his

studies

in the

author

Dean of the

Sales,

and Medicine,

and thus

to p u r s u e

a p p r e c i a t i o n that the

Chester

to Dr.

D e p a r t m e n t of S u r g e r y

g r a du at e
which

to Dr.

Department

due to Dr.

Depart­

extended to Dr.

for

laboratory.

with deep and

of V e t e r i n a r y

Drury,

are

the

translations.

It is

of the

Ly m a n B r a t z l e r ,

of the m a s t i t i s

helpful

Thanks

and Dr.

of the

i n ve s t i ga t i o n,

investigation.

v e r y thankful

co oper ation,

and m a t e r i a l s

Benne f o r

also

J.

this

of M a t h e m a t i c s ,

work of this

Cathey,

Many thanks
C.

for

Special thanks

Professor

statistical

Joyce

an i m a l s

indebted to P r o f e s s o r

Will iam

The

securing

ex­

College
and Head

Albert

R.

appointment to a
support

United

to the

States.

author

VITA
Ramchandra P r a s a d
candidate f o r
Doctor

Final

examination:

November
G i l t n er

Dissertation:

the

Yadava

degree

of

of Philosophy

15,

1955;

2:00 p.m.

Hall

Comparative

Histology

of the Kidney

of Domestic

Animal s
Outline

of Studies
Ma jo r

subject:

Anatomy

Minor

subjects:

Physiology;

Sur ger y

and Medicine

B i o gr a ph ica l I te ms
Born:

Novembe r

Undergraduate

1,

1920, P a t n a ,

Studies:

Veterinary

Patna

College,

Graduate

College

Studies:

College,
E x p e ri en ce :

East

stitution.
Patna

of Science

of the teaching

College,

Patna,

Professor

during

the

Appointed

as

U n i v e r s i t y in the

nary

leave

uate

studies

and Bihar

the

by the

degree,

Michigan,

Held the positions

lege, Patna,

College

B i h a r , India.

Science f r o m

Lansing,

A member

and A s s i s t a n t

India

Obtained the
Bihar

Veteri­

1943.

Master

Veterinary
1953.

in

Science

Patna,

diploma in V e t e r i n a r y
n ary

Bihar,

state

in anatomy

xi

Michigan

1953-1954.
staff of the

Bihar,

India, f r o m

of D e m o n s t r a t o r ,
at the

State

Bihar

Bihar
1943 to

Lecturer,

Veterinary

ten y e a r s

of

Examiner

in Anatomy by the

y ear

1952.

service

Col­

Granted

g ov e r n m e n t f o r

to the

in­

extraordi­

advanced g r a d ­

in the United States,

February

5,

1953.

S ur ge r y
Member

G ra d u a t e

of Bih ar

member
Indian

Assistantship

and Medicine
Veterinary

of the

since

in the D e p a r t m e n t

the fall

Association;

of

1953.

Phi

Zeta;

Society of Sigma Xi;

Students A s s o c i a t i o n

sity .

xii

associate

President

at Michigan State

of

of the

Univer­

TABLE

OF

CONTENTS

Page
I N T R O D U C T I O N .........................................................................................................................

1

HISTORICAL BACKGROUND

4

REVIEW OF
Gross

LITERATURE

Anatomy

Microscopic
The

.............................................................................

Tubules

The nephron
r en al

distal

collecting

The

arched

The

straight

The b a s e m e n t

10

11
23
27

.....................

27

t u b u l e s .......................................

30

collecting tubules

30

The p a p i l l a r y

tubules

.. .

.......................................

convoluted tubule

10

11

c o r p u s c l e .................................................................

The loop of Henle

The

............................................................

convoluted tubule

8

10

.........................................................................

The p r o x i m a l

The

8

...........................................................................................

The U r i n i f e r o u s

The

.................................................................

............................................................................................................

Anatomy

Capsule

.................................................

collecting tubules
ducts

. .

31

...............................................

membrane

of the

32

uriniferous

.....................

33

The P a p i l l a ............

33

xiii

Pag e
The I n t e r s t i t i a l
Intertubular
Cell

S p a c e ...........................................................................................
cell

groups or

Becher's

unit of Goormaghtigh

or

cells

. . . .

..................................................................................

35
36

C i r c u l a t i o n through the k i d n e y ..................

39

Arteriovenous

...........................................

41

..............................................................................

42

Lymphatic
Nerves

anastomoses

Vessels

...........................................

42

MATERIALS AND M E T H O D S ......................
Source of Animals
Techniques

.............................................................................

............................................................................................................

F i x a t i o n and P r o c e s s i n g
Sectioning

ofthe

Tissues

............................................................................................................

Staining

and Mounting

Methods

of M e a s u r i n g ..........................

.........................................................................

RESULTS AND DI SCUSSI ON .....................
Gross

34

s ocle-

p l a s m o d i u m ........................................................................................................
Blood V e s s e l s

34

Anatomy

Blood

Vessels

Lymphatic
Ne rv e s

.....................................................................
......................................................................................

Vessels

...............................................

..............................................................................

xiv

45
45
46
46
48
48
48
50
50
52
54
54

Page
Microscopic
The

Anatomy

The U r i n i f e r o u s

Tubules

The neph ron
ren al

57

..............................................................................

60

c o r p u s c l e ................................................

60

convoluted tubule

The loop of Henle
The d i s t al

...

69

..............................

73

convoluted tubule......................................

collecting

tubule

The

arched

The

straight

77

...............................................

collecting

The p a p i l l a r y

tubule

collecting
duct

tubule

82

.

83

..................

83

.................................................................

The b a s e m e n t m e m b r a n e
tubule

55

....................................................

The p r o x i m a l

The

55

........................................................................................................

Capsule

The

.......................................................................................

of the

85

uriniferous

................................................................................................................

91

The P a p i l l a ..............................................................................

91

The I n t e r s t i t i a l

95

Connective

The i n t e r t u b u l a r
Cell

unit

cell

groups

of Goormaghtigh

SUMMARY AND CONCLUSIONS
LITERATURE

Tissue

CITED

or

...................................

or B e c h e r ' s

s oc le pla s mo di um

..................................

.....................................................................

XV

cells

.

98
101
103
Ill

LIST OF

TABLES

TABLE
I.

Page
Literature
Volume

II.

Review on N um b er ,

Size,

and

of the G l o m e r u l u s .......................................................................

Diameter

of the

with E p ithe lia l

Uriniferous

Tubule,

H e i g h t ...............................................................................

xvi

13

6l

LIST

OF P LAT ES

PLATE
I.

Page
Capsule
the

of the kidney

c a ps u l a f i b r o s a

of the

goat, showing

and c a p s u l a

adiposa

..........................

59

...........................................................................

67

III.

Kidney of the p i g ....................................................................................

72

IV.

Kidney of the

II.

V.
VI.

Kidney of the

goat

c a t ...........................................................................

Kidney of the h o r s e
Papillary

ducts

.................................................

75
81

in the dog k i d n e y .....................................

88

VII.

The p a p i l l a r y

ducts

in the ox kidney

...

90

VIII.

The p a p i l l a r y

ducts

in the cat kidney

. . . .

93

IX.

Kidney of the

pig

.....................................................................

97

.......................................................................

100

connective
X.

tissue

Kidney of the

ox

stained f o r r e t i c u l a r

xvii

INTRODUCTION

This
croscopic
dog,

i n ve s tiga tio n deal s

structures

and cat.

The

m or p h o l o g i c a l
A d etailed
the

of the kidney

pig has

sim ilarity

of its

s tru ctu res

b een given,
of the kidney

his tology in the

structure

mentals
was
of

of

of the

v er y

without

Th e r e

is

the

of the

Animals."

densa,

1952,

above.
physi­

good textbooks
normal

of

structure

the

normal

an Englis h

textbook,

"Funda­

Although this book
there

is

of the kidney

no mention
such as

and i n t e r t u b u l a r

structures

species,

1

mentioned

except

German

s pe c i e s .

of i m m e n s e

with the

animals,

structures

macula

many

pig,

with the

none which d e s c r i b e s

and published in

a d e s c r i p t i o n of the
mentioning

are

that deal

of Domestic

apparatus,

an i m al s

goat,

of the kidney of

b e e n made

of the kidney is

and F i e b i g e r s '

important

sheep,

structures

of the

of the d om e s t i c

revised,

of the

c e r ta in places

value.

Histology

juxtaglom erular

made

structure

Trautmann

translated,

some

rest

But t h e r e

of the kidney

translation

microscopic

English language

of the h um an kidney.

ox,

kidney to that of the human

of the

and pathological

study of the m i ­

a type b e c a u s e

and a c o m p a r i s o n has

The m i c r o s c o p i c
ological

of the h o r s e ,

b e e n chos e n as

d e s c r i p t i o n of the

pig has

with a c o m p a r a t i v e

cells.

of the kidney has

which c r e a t e s

confusion.

In
been

2
Te r e g
domestic

(1911)

an i m a l s

cont r ibu ted

a valuable

in E l l e n b e r g e r ' s

chende m i k r o s k o p i s c h e n Anatomie
w r i t t e n long

ago

then by v ar i o u s

and many new
inve s tiga tio ns

chapter

textbook,
der

on the kidney of

"Handbuch d er

H austiere."

structures

But

V er g l ei -

this

was

have b e e n d i s c o v e r e d

since

on the kidney of m a n and l a b o r a t o r y

a nimal s .
Kunkel
the

number

mestic

(1930) and

and

size

animals.

remained

comparative

g l o m e r u l i of the kidneys

were

valuable

c ont r ibutions ,

studies

of v ar io us
but the

of

do- ■

studies

confined to a v e r y l i m i t e d field.
and Hallman

studies

but they also
apparatus,

of the

These

Langham
extensive

Rytand (1938) mad e

(1939),

of the m i c r o s c o p i c

left

macula

some

of the

d ens a,

and Eangham
structure

important

et al.

of the bovine kidney,

structures

and i n t e r t u b u l a r

(1942) made

ce lls )

(juxtaglomerular

of the kidney un­

touched.
Bloom
the

(1954)

kidney of the
This

d e s c r i p t i o n is

gators,

and c e r t a i n l y

less

is

co mplete

based

of g r e a t

of the kidneys

Nowhere

a d e s c r i p t i o n of the n o r m a l

dog and c a t in his

Cat."

s tru ctu res

gave

’’Pathology

value

so f a r

has

there

of an u p - t o - d a t e

as
are

the

investi­

microscopic

c o nc er n ed .

been found a m o r e
study

of

of the Dog and

upon the work of v ar i o u s

of the dog and cat

in the l i t e r a t u r e

description

book,

structures

or

of the c o m p a r a t i v e

3
histological
pig,

dog,

structures

and cat.

This

something t o w a r d s

this

All m i c r o s c o p i c
microscope,

of the

and the

s tudies

domestic

author hopes

th at w o r k in the
on the

reversal

serve

of the Golgi

In spite
as

histologists,

sheep,

s ci en ce .
under a n o r d i n a r y

of t h e s e

a reference
and

students

an i m a l s

cannot be

study is

necessary

work

also

to

reveal

omissions

the

to b i o l o g i s t s ,
of anatomy.

will

s tru ctu res

densa,

endings,

such

this
struc­

u n d er ta ke

remains

el e m e n t in the m a c u l a
the n e r v e

light

expected f r o m

s ome i n v e s t i g a t o r s

F urther

cells,

goat,

u n d e r t a k e n to c o nt r ibu te

w e r e mad e

that

near future.

of the j u x t a g l o m e r u l a r

vessels.
will

a s p e c t of the

Electron microscopic

tures,

u l es

in ve st i g a t i o n was

ox,

and hence the d e s c r i p t i o n of u l t r a m i c r o s c o p i c

of the kidney
work.

of the kidney of the h o r s e ,

to be

done

the g r a n ­

and the lymph

author hopes t h a t this p a p e r

physiologists,

pathologi st s,

HISTORICAL

The
fourth
pher,

study of the m a m m a l i a n kidney goes

c e nt ur y

and founder

animalium.n

and the pelvis

the kidneys
aware

Aretaeus

in

( early

which has
of the

n e ph r os

A.retaeus

the p e r c o l a t i o n of the

the

nervous

canals,

like

to

to be t r u e

probably b e c a u s e

for

urine;
reeds,

of the b l a d d e r

on each

side;

of the kidneys

to the b l a d d e r

11the

in his
nephros"
The u r e t e r

and t hese
which a r e

equal.''

4

s u s p e c t that A r e t a e u s

g l an ds ,

of the

cavi ties

a d e s c r i p t i o n of

ducts

(ducts

s m all

have

of Bellini).

and c o m p a r e d

s u p er fic ia l

are

similarity

and like

attached to

inserted

and the p a s s a g e
is

the

to c y s t i s . 1'

of the p a p i l l a r y

''their

as

an i l l u s ­

of the n e p h r o s . ' 1

led many a n a t o m i s t s

stated

gave

system

second ce n t u r y A..D.) gave

the kidneys

with the t e s t e s ,

cavity

back

a Greek philoso­

anatomy,

called the kidney

1'the

existence

considered

shape.

Aristotle

ltduct f r o m

Aretaeus

them

(384-322 B.C.),

of c o m p a r a t i v e

of the kidney

c a l l e d the

was

when A r i s t o t l e

as f a r

d e s c r i p t i o n of the m a m m a l i a n u r o - g e n i t a l

1' H i s t o r i a

was

B.C.,

scientist,

trated

BACKGROUND

into the

of urine f r o m

sieves,

each of
s houl de rs
each

side

5
Leonardo

da Vinci

investigators

of all

ske tc h of the

outline of the

Berenger
lar

time

(1452-1519),
and an a r t i s t

very

and it was

i n v e s t i g a t i o n of the f o r m

amplissimis

Berenger

m entione d that he injected

of the human b o d y . IT
There

is

described

a crude

Vesalius'

neys.

the

in his

biological

a valuable

in the

that led him
In his

anat omia m

as

"Com-

Mu n d i n i ' 1

r e n a l blood v e s s e l s .
d e s c r i p t i o n of the m a l e

monograph

" O n the f a b r i c

t r e a t m e n t of the kidney is

divided in two by a

(1520-1574) at tacked

s ievelike

Vesalius f o r having

dog instea d of that of the human.

d e s c r i b e d f o r the f i r s t
He use d magnifying

time

lenses

vascu­

to an

inadequate.

d i s c u s s i o n of the kidney in which the pelvis

sented the kidney of the
chius

super

an i l l u s t r a t e d

system

and r e p r e s e n t e d

Eustachius

interested

interest

additionibus

(1543) gave

u ri n o - g e n i t a l

ma d e

and function of the kidney.

cum

and f e m a l e

greatest

system.

much

this

m entaria

V es a liu s

of the

anatomist,

urino-genital

(1470-1530) was

bed of the o r g a n s ,

one

the c o r t i c a l

s ubstance

and i njected v e s s e l s

is

structure.
repre­
Eusta­

of the k i d ­

to aid his

study.
Ruini
his

(1599) gave

m o n og r ap h
Ruysch

the

on the

an excellent

d e s c r i p t i o n of the kidney

anatomy of the h o r s e

(1638-1731),

p e r i o d , b e c a m e f amous

one of the m o s t
f o r his

(A.natomia del
capable

Cavallo).

anatomists

injection technique

and

in

of

studies

6
of the

renal

glomerulus.

erulus

Ruyschiana"
Bellini

the

papillary

ducts

glomerulus

ducts

and hence

of Bellini.

tubules.

and,

aid of injected

they c o m m e n c e d

as

''apples

ferred
ers

to as

tubules

as

F e rre in 's

uriniferous

Bertini
columns

or

pyramids.

are

are

hence

o b s e r v e d the

specimens,

ca ps u l e s

rays.

the

small

zone of the
These

con­

noted that

containing

a clus­

arteries

now commonly

re­

one of the found­

F errein

also

groups
r en al

medullary

of

straight

pyramids
rays

d e s c r i b e d the

were

into
once

convoluted

of the kidney.

(1772-1845) d e m o n s t r a t e d f o r

which a r e

of the kidney

Malpighi was

the boundary

medullary

tubules

are

(1659) went beyond this

Malpighi

structures

(1693-1769) d e s c r i b e d

the

ducts

which in t u r n hang on the li t t l e
These

and

anatomy.

r ad i a t i ng f r o m

cortex

swellings

Malpighian c o r p u s c l e s .

of m i c r o s c o p i c

called

inflated

on a t r e e . "

F errein

the

with the

s m a l l blood v e s s e l s

the p a p i l l a r y

These p y r a m i d s

voluted t u bu les ,

like

"Glom­

composed of those p y r a m i d a l

In addition,

of

named

collecting tubules

Malpighi

of Malpighi.

ter

pyramids

straight

that the kidney was

of B e l l i n i ' s

was

him.

of the kidney,

the

by d e m o n s t r a t i n g

c a lle d

after

renal

(1662) d i s c o v e r e d the

also known as

m asses

The

inward

extensions

of the

the f i r s t
cortical

t im e

the

structure

r en al
of

7
the kidney,
are

betwee n the

al so known as
Henle

uriniferous
T h ese

the

ce l l s

of the kidney in his

William Bowman

neph r on that i s
functional

these

r en al

columns

of the

’’Allgemei ne A n a t o m i e . "

of Henle.

(1834-1897) is

with little

Hence

a d eta ile d d e s c r i p t i o n of the loops

r e m e m b e r e d f o r his

of the kidney by injecting

Working
n eedle,

pyramids.

of Be r t i n .

c a lle d the loops

Heidenhain
ing

columns

(1841) gave

tubules

are

renal

more

than

(184Z) f i r s t

rem arkable

close

method of

indigo-carmine

a microscope
drew

into the blood.

and a d i s s e c t i n g

a picture

to p r e s e n t - d a y

stain­

of the

ideas

single

of the

r en al

unit.

The

above

work of Singer

description

(1925),

of the

Castiglioni

historical

(1947),

events

and M e t t l e r

is

b as ed

(1947).

on the

REVIEW OF

Gross

Chauveau
pyramidal
lobe s

with w el l - de f i ne d

lobes

e x te r na l lobation having f if te en to twenty

Langham and H a l l m a n (1939) found six tee n to

to the

(1873), B r e m e r
structure

Th e s e

life.

kidneys

in the m a m m a l i a n fet us

authors

adult.

S tr a u s

with di s t i nc t

during i n t r a ­
of m u l t i l o b a r

s u p e r f i c i a l lobation in the kidney of
di s t i nc t l y , in the
multipyramidal

unipyramidal.
(1944a) and Smith

multipyramidal

(1951) d e s c r i b e d

with d i s t i n c t

tinct p y r a m i d s

in the kidney of the pig,

f a c e was

smooth as

in the

case

8

the kidney of the

ex t e r n a l lobation.

(1944) and S i s s on and G r o s s m a n

quite

and A.rey (1954)

s u p e r f i c i a l lobation.

(1934) found the kidney of m a n to be

Grahame

Elias

(1953),

and s o m e t i m e s , though much l e s s

and t h at of apes

elephant as

Patten

d e s c r i b e d the p r e s e n c e

Dixon (1931) d e s c r i b e d
child,

(1944),

of the m a m m a l i a n kidneys

uterine

a young

multi-

in each kidney of the ox.

Chauveau
referred

Anatomy

(1873) d e s c r i b e d the kidney of the ox as

i n each kidney.

thirty-two

LITERATURE

(1953) d e s c r i b e d

although the

external

of u ni p y r a m i d a l kidneys.

dis­
sur­

9
Elias
goat

as

a u nilobar

shepherd
those

dog,

organ

Elias

of the pig.

united;
was

(1944) d e s c r i b e d the kidney of the h o r s e ,

in the

similar

s t a t e d that the p y r a m i d s

In a g r e a t

ca t

a

simple,

Dane

German

of the kidney

the p y r a m i d s

and

resembled

w ere

c l os e ly
pyramid

present.
(1953)

tion of the kidney of the h o r s e
The i n n e r

central

which was

projected

was

called

part

a renal

into the

crest.

pelvis

The

d escrib ed distinct pyramids

constituting

minores.

sheep,

the

sixteen p y r a m i d s .
papillae
kidney,
r en al
rid ge s

The

renal

the

crest.
in

producing

r idges

projection

cribrosa.

with t h e i r
columns

In the

papillae

w ere

also

ox t hese

of
au­

projecting
v er y

into

distinct.

c r e s t was f o r m e d by the fusion of twelve

pig.

proceeded
above

a manner
a false

This

ridge

c r e s t p r e s e n t e d a number

Sisson and G r o s s m a n

Sections

such

area

renal

in the kidney of the
curved

a nonpapillated a p p e a r a n c e .

of the kidney.

r enal

thors

In the

sec­

of the m ed ull a was f o r m e d by a concave

openings

calyces

s tated that the longitudinal

presented

small

lae,

dog,

In the

u nilo ba r kidney with a simple

Sisson and G r o s s m a n

the

to that of r o d e n t s .

sheep,

as

or

also

In f r on t a l
dorsally

impression.

sections

dis tinc t

of the

and v en t r a l l y f r o m

below the

to give the

described

r en al

to

dog
the

c r e s t cut those

ap p e a r a n c e

of conical p a p i l ­

10
Microscopic

The

Chauveau
of the h o r s e

as

(1873)

chyma of the kidney into
Tereg
as

vessels,

and n e r v e s .

of the

capsule

Mollendorff
and

Copenhaver

being

capsule

layer

The in n e r

layer

was

contained blood and lymph

smooth m u s c l e f i b e r s

Maximow

the p r e s e n c e

of the human kidney

united to the p a r e n ­

(1911) and T r a u t m a n n and F i e b i g e r

(1953) d e s c r i b e d the

and Bloom
caps ule

por­

sheep.
(1952),

and Smith

of the human kidney

and a few

of a few

in the d e e p e r

(1952)

e l as tic f i b e r s .

smooth m u s c l e

in addition to the usual

cells

as

G r ee p

in the

collagenous

and

fibers.

The

Greep
of two

layers.

of the kidney of the ox and
(1930),

of the kidney

ca p s u l e of the m a m m a l i a n kidney

co mpo s ed of collagenous f i b e r s

(1954) d e s c r i b e d

elastic

the

in t i m at e l y

tunic

sent many prolongations.

The o ut e r

of

enveloping

separable

Te r e g

d e s c r i b e d the p r e s e n c e
tions

which it

easily

devoid of blood v e s s e l s .

the

membrane

(1911) d e s c r i b e d

c o m p o s e d of two

Capsule

described

a fibrous

Anatomy

Uriniferous

(1954) d e s c r i b e d

segments,

a

secretory

Tubules

a uriniferous
p or tion,

the

tubule

nephron,

as

being

having

composed

the function

11
of e l a b o r a t i o n of u r i n e ,
to

convey u ri n e

into

the p elv is

in the h um an kidney
long,

the

secretory

and a collecting

was

portion,

of the kidney.

described

p o r t i o n being

as being

30 to

the

collecting

A. u r i n i f e r o u s
50 to

tubule,

tubule

60 m i l l i m e t e r s

40 m i l l i m e t e r s

in length.

The neph ron

H uber
kidneys
the

(1932) t e a s e d

t h a t had been

nephron the

of the

renal

subjected to acid m a c e r a t i o n .

r e n a l tubule

corpuscle,

m e d u l l a r y loop,

out individual n ephrons f r o m

the

which was

the p r o x i m a l

d is t a l

adult

Huber

rabbit

called

c o n s i d e r e d to be made

up

convoluted p ortio n with the

convoluted portion,

and the junctional

tu­

bule.

The

renal

corpuscle.

kidney w e r e found to be

The

spherical

1 0 0 (jl and

2 0 0 |jl (Huber,

1932,

1935).

observed

the

of the

renal

ney to be
the

size

age,

corpuscles

with a d i a m e t e r
Langham
corpuscle

of the

reaching

renal

corpuscle

a maximum

varying between

with the

(1954) d e s c r i b e d the

of the h um an kidney

as

being

(1939)

of an adult bovine k i d ­
They noted that
advancement of

in a fully grown individual.

(1953) and G r ee p

corpuscle

increased

of the m a m m a l i a n

and Hallman

2 1 6 [jl in a fixed and stained p r e p a r a t i o n .

Copenhaver
renal

diameter

renal

Smith and

diameter

close to

2 0 0 )j l .

of the

12
The
time

glomerulus

of the m a m m a l i a n kidney was

to t i m e by v a r i o u s

investigators.

gations

showing the n u m b e r ,

uli

given in

are

siz e,

results

of t h e i r

and volume

investi­

of the

g lom er­

Table I.

Shonyo and Mann
medullary

shape,

The

studied f r o m

zone w e r e

(1944) noted that the g l o m e r u l i

usually l a r g e r

in the c o r t i c o -

than th o s e lo c a t e d m o r e p e r i p h -

e rally.
The a f f e r e n t a r t e r i o l e
Be ns le y
the

(192-9) to be

glomerulus.

expanded
afferent
erular

Schloss

arteriole

branches

s o r t of r e s e r v o i r

w er e

after

entering

s een to a r i s e f r o m

(1946) o b s e r v e d a s i m i l a r

of the human kidney and c a lle d it

this

dilatation in the
" B e c h e r ’s g l o m ­

sinus."
Dorello

of the pig was

(1948) noted that the
expanded and gave

each of which gave
Codden
teriole

guinea pig kidney was found by

expanded into a

Prim ary

sinus.

of the

rise

o r i g i n to t h r e e

to t h r e e

in the kidney

p rim a ry branches,

m o re branches.

(1949) did not find any dilatation in the

of the h u m a n kidney

af t er

He found a r a m i f i c a t i o n of the
t e r e d the

aff e r e n t a r t e r i o l e

renal

e n t e r e d the

afferent arte rio le

ren al

corpuscle.

soon a f t e r

it

en­

corpuscle.

The g l o m e r u l a r
with the help of

it

afferent a r ­

capillary

tuft was

celloidin c o r r o s i o n

studied by Wi l m e r

injection technique.

(1941)

It was

TABLE
LITERATURE

I

REVIEW ON NUMBER,

VOLUME OF

SIZE,

AND

THE GLOMERULUS

Diam­
Nu m b e r
Autho r

Animal

of

eter

of

G lo me r u l i

the

in One

Glom -

Kidney

erulu s
(p)

Vimtrup

(1928)

. . .

cat

202,813

dog

407,1 55

albino r a t
m an
McGregor
Kunkel

(1929) . .

(1930)

. .

erular
Volume
in One
Kidney
(cu.

mm .)

4-2

man

200-237

dog

163
166

m an

(1931)

Glom­

33,826
1 ,233,360

Pig
ox
Moo re

T otal

c .c .

259
800,000 to
1 ,000,000

Rytand

(1938)

. .

cat

214,500

150

2Z7

dog

408,1 00

180

1 ,247

Pig
ox

1 ,193,000

16 6

3,992,000

244

2,859
29,860

186,600

166

447

elephant

7,51 0,000

338

151,900

ox

800,000 to

monkey

Lang ha m

et a l . . .

1 ,700,000

(1942)
G ree p

(1954)

. . . .

man

1 ,000,000

14
demonstrated

that the

af f e r e n t

prim ary

and c o r r e s p o n d i n g

d et a i l e d

observations

two-thirds

of the

any a n a s t o m o s i s
Smith
kidney gave
branches,

arteriole

secondary branches.

could not be

af f e r e n t

off two to fo u r

It was

noted that

on a p p r o x i m a t e l y

system,

Wilmer

the f i r s t

did not o b s e r v e

capillaries.

(1951) found that the

fifty c a p i l l a r y loops.

made

arteriolar

between the

which in t u r n

of the h um an kidney gave

or

aff e r e n t

m ore,

arteriole

rarely

subdivided again,

of the human

up to ten, p r i m a r y

at t i m e s

The p r i m a r y b ra nc hi ng

into

as

many

as

gave the tuft a lobulated

structure
Tr abuc co
the

glom erular

and e f f e r e n t
from

one

side

a

of a

They noted that the

single

series

According

be

a dense

looping
show
in the

No

glomerulus

simple

channels

s h o r t blind
glomerulus

were

nor

to Boyer

a

a f f e r en t

was f o r m e d

the

w er e

seen.

s een to c r e a t e

sacculations.
of the

r at ,

Hall
cat,

was

skein of blind f in ge r l i k e
glomerular

tuft was

c a p i l l a r y bed with an a f f e r en t

loops

that

bent on itself.

of loops

sac cu la tion s .

channel.

vessel

between the

(1955) d e m o n s t r a t e d that the human g l o m e r u l u s

si m pl e

essentially

(1952) did not find any evidence

tuft had any looping f o r m a t i o n s

arterioles.

Boyer
neither

and Marquez,

Multiple

and

found to
efferent

anastomoses

between

a complex network which might
(1955) o b s e r v e d

dog,

rab bi t,

d irect

channels

and m a n f r o m

the

15
a f f e r e n t to the
function

as b y - p a s s e s

s e e n to
joined

e f fe r en t

give

rise

end. of the lobule.

for

medially

and divided f r e e l y

plex but unified

erythrocytes.

system

w er e

The l a r g e

to n u m e r o u s

to f o r m

Th e s e

small,

a true

capillaries

wer e

short branches

capillary

of communi cating,

thought to

network;

a n as t om os in g

which

a com­

capillaries

within the lobule.
The

ef f e r e n t

arteriole

(1929) to p r o c e e d in a
first

of c a p i l l a r i e s

efferent v e sse ls

usually

distance

as

medullary
of the

renal

vessels,

vessel.

The

much longer

capsule

of the

of the

cortex,

in a

cor tex.

which w er e

on e m e r g e n c e f r o m

w e r e found to continue f o r
efferent

at

vessels

and f o r m e d the

the
some

of the juxtaarteriole

re c t a e

medulla.
(1946) o b s e r v e d

arteriole

demonstrated

whe r e

in the human kidney a dilatation in

it left the

that in the human kidney

arterioles

were

laries.

zone

found by Be ns le y

found to t e r m i n a t e

resolved immediately

capillary

r egion w e r e

ef f er ent

ferent

toward the

It was

of the middle

a discrete

Schloss

zone

course

around the convoluted tubules

supposed to be

g l o m e r u l i into the

the

sinuous

guinea pig was

without giving off c a p i l l a r i e s .

plexus
The

of the

s ho rt,

of the

glomeruli

t h i n- w a l l ed ,

Such c a p i l l a r i e s

g lom e r u l u s .

Edwards

(1953)

20 to 25 p e r

cent of the

ef­

s ituate d in the c o r t i c o - m e d u l l a r y

and gave

w e r e found to

rise

to a network of c a p i l ­

supply blood to the p a r t s

of

16
all n ep h r o n s
efferent

arterioles

vascularized
terioles
arising

located

in the

which w er e

were

found to c o m p r i s e

from

the g l o m e r u l i

m a n and m a m m a l s
small

reticular

was

only

w h e rea s

d e s ce nd into

75 to

The long

80 p e r

or

axial

space

found by Be ns le y

surrounding

reticu lar fibers

the

of the

vessels

extending

ef fe r en t

zone.

normal

glomerulus

th e i r

a layer

en tr a n c e ,

into the lobes

of

(1930) to con­

They o b s e r v e d
at

ar­

cent of those

and Bensley

tissue.

the long

the med u l l a

in the c o r t i c o - m e d u l l a r y

amount of connective

fibers

and d el i c a t e

found to

zone,

only the m e d u l l a r y p a r e n c h y m a .

The i n t e r c a p i l l a r y

tain a

cortico-medullary

of

with s p a r s e

of the

glom er­

ulus .
McManus

et a l . (1951) made

human g l o m e r u l u s
rial

in the

intercapillary

J on es

cells.

ous

than

the

endothelial

connective

either

the

cells.

tissue

visceral

cells,

presence

of the n o r m a l

space,
ce l l s

a p pr o xi ma tel y

human g l om er u l us

of Bowman's

of age t h e i r

to that of the

had

connective

w ere found to be l e s s

years

m ate­

human g l o m e r u l u s .

which contained

epithelial l a y e r

but by t h i r t y

study of the

of c a r b o h y d r a t e

that the n o r m a l

In childhood th e s e

found to i n c r e a s e
t h eli al

space

(1953) o b s e r v e d

an i n t e r s t i t i a l
tissue

and o b s e r v e d the

a histochemical

caps ule

number

endothelial

numer­

was

and e p i ­

or

17
Ha.ll

et al

(195 3a) found the

exceedingly few i n t e r s t i t i a l
all.

B e ne de tti

and connective

ce l l s

and Scapellato
tissue

cells

rat

glomerulus

to have

and collagenous f i b e r s

(1954) o b s e r v e d

in the

stroma

or

none

a r g yro ph i l i c

of the

normal

either
at

fibrils

ren al

glom­

erulus .
The j u x t a g l o m e r u l a r
(1939)

as

being

myofibrils,
of the

compos ed

called

Ma f i b r i l l a r

cycle

the

were

or

with the

superficial

ce lls

were

at t h e i r

cortical

w e r e found to

vacuoles.

capillaries.

af fer ent

m a c u l a d e ns a

and the

pad was found to c o n s i s t
delicate, f ib r i l l a r

or

show a

basophil

These

In the

zone

afibrillar

case

af­
of

not found.

the

junction with the

devoid of

contact with the lu me n of the

glom erular

surrounding

region between the

group

th e s e

cells,

(1940) found in the m a m m a l i a n kidney

pad in co mp le tely

McManus

In the

i n t e r m i n g l e d with minute

s ecretion granules

Ed wa r d s

terioles

rabbit,

d e s c r i b e d by Goormaghtigh

smooth m u s c l e

cells.M

o b s e r v e d to be in cl ose

arteriole

dog,

of

was

culminating in the f o r m a t i o n of acidophil

s ec retion granules

ferent

mainly

n o r m a l kidney of the

g landular

cells

apparatus

arteriole

of a n e s tlik e

and occupying

a f f e r e n t and

glomerulus.
group

a periartiolar

eff erent

a

ar­

This p e r i a r t e r i o l a r
of cells

embedded in a

network.
(1942)

of a p p a r e n t l y

defined the j u x t a g l o m e r u l a r

s pe c i a l i z e d

structures

in

apparatus

as

a

r e l a t i o n to and including

18
the
to

afferent
use

the

and

efferent a r t e r i o l e s

term

' 'juxtaglomerular

"juxtaglomerular
glomerular

apparatus

Graef
the

(1943)

arterioles

hormonal

ce lls

(1946)

of the

Des

complex”

He p r e f e r r e d

i n s t e a d of G oo r m a g h t i g h ' s

"com plex"

c o n s i s t e d of the j u x t a ­

densa.

m ed ial h y p e r t r o p h y

of a p r e g n a n t

reported

woman,

the

existence

of

and h y p e r p l a s i a of
p ossibly

due to

smooth m u s c l e

1* Po l k i s s e n 1’ ( j u x t a g l o m e r u l a r

in addition to the

a

usual

ap pa r at us

epithelioid c e l l s .

These

cells

of Goormuscle

contained m y o f i b r i l s .
McManus

the

reported

glomerulus.

effect.

area

maghtigh)

His

and m a c u l a

in the c a s e

Schloss
in the

apparatus.”

of the

(1947b) d e m o n s t r a t e d the

juxtaglomerular
Prez

granules.

cells

with the help of per io dic

(1948) used M a s s o n ' s
These

ce l l s

presence

trichrom e

appeared large,

prominent

ovoid nuclei whose

somewhat

radial pattern.

cy to pla s mi c

juxtaglom erular
G om o ri

apparatus

g ra n u l e s

the j u x t a g l o m e r u l a r

cells.

in the

was

arranged

th es e

in a

Dalton (1951) d em on ­

myoepithelial

(1951) o b s e r v e d
Wilson

reagent .

all a f i b r i l l a r .

in the kidney of the

and Oltvanyi

Schiff

pale - s t a i n i n g , with l a r g e ,

With the help of e l e c t r o n m i c r o s c o p y ,
strated large

acid

in

sta in to d e m o n s t r a t e

c h r o m a t i n network

They wer e

of g r a n u l e s

ce lls

of the

rat .

a n er ve

(1952) developed

plexus

among

a spe ci al

staining

19
t echnique f o r

demonstrating

The g r a n u l e s

were

Maximow
ular

cells

the

granules

stained deep p ur pl e ,

and Bloom

in l o w e r

of the j u x t a g l o m e r u l a r
and nuclei

(1952) noted the

vertebrates

cells.

light purpl e.

a b s en ce

of j u x t a g l o m e r ­

and in c h i l d r e n below the age

of two

year s .
The function of the j u x t a g l o m e r u l a r
versial.

Goormaghtigh

b e l i e v e d that the

(1939,

afibrillar

had endo cr i ne function.
was

1940,

ce lls

The

1945a,

ap p a r a t u s
1945b,

endocrine

tion,

ischemic

the

kidney.

afibrillar

Kaufmann (1942),
(1952)

activity of the

possessed

Oberling
in c h i l d r e n

J ones

(1947,

(1944) did not find any
whose kidneys

and Schloss

of G o o r m a g h t i g h ' s

juxtaglom erular

afibrillar

r en al

and H a r t r o f t

1951)

ap p a r a t u s

s ubstance

of the

contro­

cells

present
condi­

arterioles.

and H a r t r o f t

that the j u x t a g l o m e r u l a r

an endocrine function.

in the j u x t a g l o m e r u l a r

s up po r t

1949),

view of Goormaghtigh

and adults

(1945),

1949,

suggested that in the n o r m a l

reg u l a t e d the tonus

Dunihue

supported the

apparatus

cells

ce l l s

It was

very

1947,

of the j u x t a g l o m e r u l a r

r e l a t e d to the production of the h y p e r t e n s i v e

in the

is

apparatus.

possessed

apparatus.

(1948) w er e
hypothesis

elevation of blood p r e s s u r e
abundant

Edwards
unable

granular

(1945),

Fox and

to find any

evidence

of endocrine function of the

in

20
Graef

and P r o s k a u e r

(1945) and Be c h e r

t hat the function of the j u x t a g l o m e r u l a r
renal

blood flow.

juxtaglom erular

Hartroft
cells

sodium m e t a b o l i s m
The t h r e e
by M c G r e g o r

(1929).

epithelial,

kidney.

B e ns l e y

s m all

extent of the

surface

the

wall

found to be

vital

De

that the

r eg ul at i o n of

of the g l o m e r u l u s

o b s e r v e d the p r e s e n c e

and b a s e m e n t

to be

and did not

was

were

membrane,

a continuous

found

and its

constitute

to

be

capillaries.

layer

in the

The

mem­
human

epithelium

over the

lobes.

It was

a syncytium.

structureless,

studied

of t h r e e

(1930) found the g l o m e r u l a r

of the g l o m e r u l u s

glomerular

(1938) d e m o n s t r a t e d
property

It did not f o r m

ce lls
Renyi

ep i th el ial lining
of o t h e r

mam mals

which had the

staining.

glomerular

s uggested

whole

com­

The

supporting

endothelium was

also

continuous.

Brem er
glomerulus

m em branes

McGregor

cells

membrane
of the

(1953)

might be to cont rol

involved in the h or m on al

endothelial,

in human and

basement

and H a r t r o f t

and B e ns l e y

separate

apparatus

1950) thought

and blood p r e s s u r e .

prim ary

branes,

p osed of

were

(1949,

a protoplasmic
of storing

a continuous

w e r e found to s t o r e
(1941), using
n on continuous .

capillaries.

The

f ilm

over

dye g r a n u l e s
sheet.

the

after

Only occasional

the vital dyes.

m i c r o d i s s e c t i o n t e chniques , found the
The

structure

cells
of the

resembled

the p e r i c y t e s

endothelium

of the

21
c a p i l l a r y loops
o th e r

was

different from

lining

of the

capillaries

of

that the b a s e m e n t

membrane

was

organs.
McManus

derived from
at the

(1948a) o b s e r v e d

Bo wm a n' s

glom erular

capsule

root.

The

the b a s e m e n t m e m b r a n e .
a crenated

appearance

with minute

ex t e r n a l

Oberling
carrying
Hall

was

whose

cells

cells

Rinehart

e l a b ora ting

was

considered

uct of the

studded

layer,

villilike

a continuous

whose
r idges.

epithelial

processes.

and highly porous

basement membrane,

and an e p i ­

p o s s e s s e d inte rdigitating p r o c e s s e s .

a cytoplasmi c

at many points.

cells.

and e l e c t r o n

surface

s e c r e t i o n which was

as being p r i m a r i l y

endothelial

epithelial

(1951) found the

et a l . (1953) found the

the b a s e m e n t m e m b r a n e

which was

of i nter digi tati ng

With the help of h i s t o c h e m i c a l techniques
micrographs,

revealed

and a b a s e m e n t m e m b r a n e

of m a n y - b r a n c h e d

a finely porous

enclosed by

of the kidney

(1950) found a continuous
layer,

also

arterioles

195 0).

c o v e r e d by a sy s t e m

a system

were

membrane

et a l .,

et a l . (1953a) o b s e r v e d

theli al l a y e r

loops

et a l . (1951) and Dalton

endothelial l a y e r ,

at tached to the

Electron microscopy

endothelial

surface

capillary

(Gautier

and B a k e r

a noncontinuous

and was

of the b a s e m e n t

p ro j e c t i o n s

Pease

layer

the

epithelial

in contact

The b a s e m e n t

with

membrane

a d i f fe r en tia ted cy to pla s mi c

prod­

22

Simer
and

endothelial l a y e r s

fibrillar

basement

Hall
count

as

of t h r e e

kidney
lar

intimately

r atio

related

of the

rat

membrane

an

and found

had long p r i m a r y

and divisible

and

in t h r e e
The

at tenuated in m o s t p a r t s

of the

secreting

area

found to be

structures

epithelial

a

under

nuclear

a complex
the

electron

and

rat

to be

glomerulus

Bo w m a n ' s
call

0.19

c a ps u l e

was

Bowman's

applied d i r e c t l y
was found to

was

being

extremely

was found by Book
0.3813

(1942) found the a r e a
s qu ar e

ex­

capillaries.

g lo m e r u l u s
to be

that

the middle l a y e r

cytoplasm

glomerular

g lomer u lus

Stowell

and w ere

layers,

the

cells of the g l o m e r u ­

s econdar y b ra n c h e s

endothelial

of the

typical human

while K i r k m a n

to

to epithelial

The b a s e m e n t m e m b r a n e

of high e l e c t r o n density.

proposed

provi ded with

e l e c t r o n m i c r o s c o p i c study of

that the

basement membrane.

single

was

and between the c a p i l l a r y loops

continuous

in a

was

epithelial

(rat).
(1955a) made

The

of endothelial

specialized

Pease

tended o v e r

capillary

v isceral

membrane.

The b a s e m e n t

capillaries

to the

although e v e r y

(1954) gave the

3:1.

microscope

be

et a l . (1953) did not find complete

s qu ar e

(1936)

m illim eter,

of a single albino

m illimeter.

studied by Oleynik

ca ps u le

"Shymlansky's

(1952).

Oleynik

capsule"

as

he

b e l i e v e d that Shymlansky published his r e s u l t s sixty y e a r s before Bowman.

23
Crabtree
m an's

c a ps u l e

These

cuboidal

proximal
ules

(1941) found in m i ce

was

partly

cells

capsules

co l u m n a r
capsules

cell

composed

r e s p e c t to the

stain.

Indirect

changed f r o m

a

a

This

response

(1955b) made

ability to

p ro cesses

ce lls

cell

of the

store

gran­

type to a

of cuboidal cell

earlier

in m a l e s

endocrine f a c t o r

than

or f a c t o r s .

study of the

of Bowman's

suggestive

c e lls .

evidence indicat ed that

an e l e c t r o n m i c r o s c o p i c

and found that the p a r i e t a l
had cy to pla smi c

to an

of Bow­

of cuboidal

squamous

occurred

layer

with those

Af ter p ub e r t y the p e r c e n t a g e

suggesting

Pease

times

a vital

could be

type.

parietal

w er e found to be i dentical

began to i n c r e a s e .

in f e m a l e s ,

kidney

completely

convoluted tubule with

of t r y p a n blue,

single

or

that the

capsule

rat

some­

of a r u d i m e n t a r y b r u s h

border.

The p r o x i m a l
length of the
lim eters
average

tubule
thelium

number

convoluted tubule

diameter
of t u r n s

Huber

50p to
in the

observed

with a
a

relatively

spherical

in the

narrow

nucleus,

d i s t r i b u t e d in a m a n n e r

60fjt.

Huber

(1932,

in the

rabbit to be

Grafflin

p ro x i m a l

a r e l a t i v e l y low epithelium

possessed
ules

proximal

and the

to be f o r t y .

convoluted tubule.

tubules

rabbit's

in human kidney

p ro xi ma l

He o b s e r v e d

brush border,

to give

9.4 m i l ­

(1939) noticed the

with a wide lumen,
lumen.

1935) found the

convoluted
or

a high e p i ­

that the

and m i t o ch on dr i a l

d is t i n c t b a s a l

striations.

cells
gran­

24
G r af f l i n
iron-containing
the

p i g m e n t in the

epithelium of the p r o x i m a l

tubule

of

rat.
Ma y e r

tubular
neys
ous

(1942) noted a deposition of yellow to g olden- br own

and Ottolenghi

epithelium into the

of dogs

and c a t s .

with the p r o x i m a l

epithelium,

being

Harman
cells
per
cells

cent of
in the

lateral

tubule

and Hogan

routine
v ar io us

nephrectomy

space

t u bu l ar

and was

in

a p r o t r u s i o n of

of Bo wma n ’s caps ule

Protruded

cuboidal

in the p r o x i m a l

(1947) d e m o n s t r a t e d

epithelium

distinct from

in the k i d ­

was

the

continu­

capsular

shape.

(1949) found m ult i n u c l e a t e d giant

convoluted tubules

a u to ps ie s .
tubules

of human kidneys

epithelial
in

15.2

Sulkin (1949) o b s e r v e d binucleat ed

of the

n o r m a l kidney of the

showed a significant i n c r e a s e

rat.

Uni­

in the n u m b e r

of

binucleated cells.
Lowell
proximal
c y t o pl a s m

et a l . (1953) found

convoluted tubule

occasionally

character.

The nuclei

m o u s e kidney

as

structure

r odle ts

were

a g g r e g a t e d like the m a c u l a

The u l t r a

of the c e l l s

s u r r o u n d e d by a c o n s i d e r a b l e

c h a r g e d with w e l l - p r e s e r v e d

mitochondrial

was

each nucleus

of

s paced and

densa.

of the p r o x i m a l

r e v e a l e d by high

body of

and gra nu le s

unevenly

of the

convoluted tubules

resolution

d e m o n s t r a t e d by S j o s t r a n d and Rhodin

of the

electron microscopy

(1953).

It was found that

25
the

cell

membrane

appeared

as

a thin

to w a r d the b a s e m e n t m e m b r a n e .
be

rod-shaped.

In the

zone

of condensed

The m i t o c h o n d r i a

intermediate

cell

zone t h e r e

cytoplasm

were

found to

wer e

granules

and vacuoles .
The b r u s h
(1950)

as

being

inated f r o m
l u me n

as

border

was

composed of

the c y to pla s m

observed

by Gau tie r

and B e r n h a r d

round o r polygonal tubules

of the p r o x i m a l

tubule

which o r i g ­

and ended in the

pores.

S jos tra nd
of densely

and Rhodin (1953) found the b r u s h b o r d e r

arranged

cy l i n d r i c a l

ducts

closed

co ns i s t e d

toward the tubular l u ­

men.
Hall

et a l . (1953b) noted t h at the b r u s h b o r d e r

of individual t u b u l a r f i b e r s

having

separate

was

o ri gin s f r o m

composed

the cell

s u r f ac e .
Pease
tubule

was

(1955b) found that the b r u s h b o r d e r

comprised

of extensions

cytoplasm

of the b r u s h b o r d e r

rem ainder

of the

cell,

and

of the

had the

sim ilar

apical

same

g ra n u l e s

of the proxi ma l
cytoplasm.

density
might

as

The

that of the

even be

seen

within it.
Fat
several
in the

content

of the p r o xi ma l

investigators.
proximal

tubule

Modell
of the

convoluted tubule was

(1933)
cat.

and Foote

Foote

(1936)

and Grafflin

studied by

observed fat
(1938,

1942)

26
found fat

in the p r o x i m a l

et a l . (1950),
proximal
c a t the
of the

Silver

tubule

entire pars
proximal

the

terminal

convoluta

reverse

recta
was

several

true.

au th o r s

of the two

tubule in the

They d e s c r i b e d

them

while

In the dog,

The f at was

the t e r m i n a l

on the

confined

recta

solely

other
to the

recta.
convoluted tubule

of the histology

(1938) d e t e r m i n e d

(convoluted portion
cat

and

the

and f a t - f r e e

relative

s t r a i g h t portion)

and dog on the b a s i s

as f a t - l a d e n

s eg men t
irregular

and Grafflin

of the p r o x i m a l

of fat

s eg men ts

with a m a r k e d

tours

were

more

irregular

second

more

(1942) d e s c r i b e d the c e l l s
tubule
degree

of the

cat

and dog

of i n terdigi tati on.

c o m pl i ca t ed at the lumen.

in the

seg men t

dog than in the

were

essentially

cat.

content.

of the p r o x ­

of the f i r s t

as being highly
The

The cell
The

rectilinear

was

and physiology.

tubule .
F oote

the

that in the

upper p or t i o n of the p a r s

normally fat-laden,

and Grafflin

of the p r o x i m a l

imal

and the

on the b a s i s

segments

Dallemagne

and Grafflin o b s e r v e d

d i f fe r en tia tio n in the p r o x i m a l

(1936) and F oote

lengths

Foote

cat and dog.

(1951) d e m o n s t r a t e d fat in the

was f a t - f r e e .

p o r t i o n of the p a r s

m ad e by

of both

and P l a t t

tubule w e r e

Segmental

Foote

(1951),

of the dog.

portion of the p a r s
hand,

tubule

cell

cell

con­

shapes
contours

in both

w er e
of

species.

27
Longley
the

proximal

with the help

staining

d if fe r e n t i a t e d

Schiff

rat,

and cat,

and the

segments

thin limb

The thin

in m o u s e ,

s egment

was

and

Smith
w er e

in

instances,

loop,
the

about

the thin

thin
or

Huber

cat,

1 to

large

it might

d i f fe r en t i a t ed

and chicken.

1935) found the d i a m e t e r

rabbi t

varying f r o m

epithelium.

nuclei.

only in m a m m a l s

The d is tal

as

might be

The

distal

3.6 to

7.8 m i l l i m e t e r s .

25p.

cells

small

convoluted
The

en t i re l y.

presence

p e r ce nt ag e

tubule

Huber
in

diameter

had
thin

and,
outer

Where p r e s e n t ,
limb

some

of the

of Henle' s

dis tan ce

thin

up

s egment

of the nephron of b i r d s .

(1932,

1935) found the

rabbit kidneys

was

ones,

located in the

the loop and f or

convoluted tubule.

of the

were

absent

Smith noted the

length

of

s h o r t loops

as long

only the descending

extend around

and in a

20p to

The length of the

n um e r o u s

the g l o m e r u l i

might occupy

limb.

m ou s e,

15 m i l l i m e t e r s .

seven t i m e s

whe r e

of

Periodic

in the

technique

dog,

(1932,

s egments

techniques.

segments

lined by squamous

seg men t

segment

ascending

two

the two

(1951) o b s e r v e d that in the human kidney

of Henle

the

squirrel,

r el at i v el y

s eg m e n t v a r i e d f r o m

co r t e x ,

the

of H e n l e ’s loop in the

polygonal f o r m s

s ome

of staining

alkaline p h o sp ha t a se

The loop of H e n l e .
the

(1954) d if fe r e n t i a t e d

tubule

acid

those

and F i s h e r

varying f r o m

found to be between

30p

28

and. 4 0p .

The

epithelium
ce l l s

tubule

presenting

were

p resent,

dis tal

was

a clear

lined by a low co l u m n a r

supranuclear

devoid of a b r u s h b o r d e r

though not quite

zone

cuboidal

of cyto pla s m.

but the b a s a l

so d i s t i n c t

or

as

in the c a s e

the

distal

striations

The

wer e

of the p r o x i m a l

tubule .
Graf flin

(1939) o b s e r v e d

n e ph r on took a m o r e
glomerulus

near

the

to make few t u r n s
Trautmann
d i s t al

arteriole.

and F i e b i g e r
as

(1955b) made

had

s ho rt

scattered

m entary brush

extensive

name

of the

cytoplasmi c

passed

tubule

the

was found

collecting tubule.
the nuclei of the

cells

study of the tubular

electron microscope.
of the distal

p ro cesses

tubule

suggestive

of a

cells
It

was

usually
rudi­

border.

The m a c u l a
Zimmermann

ends

of the human

esp ec ia lly l a r g e .

r a t kidney with the help of the
apical

ca s e

The distal

(1952) d e s c r i b e d

being

an

tubule

and in ev ery

until it a r c h e d to join the

found that the

the

course

aff e r en t

tubule in the h o r s e
Pease

of the

direct

that

densa

was

demonstrated for

(1933) in the m a m m a l i a n kidney.

of m a c u l a d e ns a b e c a u s e

dis tal

convoluted tubule

tubule

and the

afferent

at the

area

arteriole

of crowding

the f i r s t

t i m e by

Zimmermann
of the

nuclei in the

of contact between the

of the

g lom e r u l u s .

gave

distal

29
Edwards

(1940) p r e f e r r e d

for

macula

the

m a m m a l i a n kidney,

birds,

and f r o g s
It

as

seen

cat,

gered
cent

so

philic

row.

of this

and frog.

or

th os e

nucleus,

towards

the

two-thirds

plaque

densa

of

in m a m m a l s ,

of

of man,

rabbit,

was

often m a r k e d l y

same

cytoplasm

the wall of the
and

row but

of the

cells

and t h e i r

and a m o r e

also

was

nuclei

than those

usually

with those

n ongr anul ar
wer e

conspicuous

and

guinea

composed of

taller

nuclei w e r e

nephron

rat,

in the kidney of the whale,

that t h e i r

in the

(1943)

el e m e n t s

and Okkels

was

whereas
luminal

in the
side

(1947a)

in the m a c u l a

(1950) o b s e r v e d

reversed

m a c u l a dens a

McManus
brane

macula

plaqueM

stag­
adja­

in contact
of the

ad­

and n e u t r o ­

somewhat
nucleolus

smaller,
than

ad ja c e n t c e l l s .

e l e m e n t in the
the

cells

chromatin

McManus
of the Golgi

’’epithelial

of the

epithelial

the kidney

slightly basophilic

of the

to

wall

cl ose t o g e t h e r

contained m o r e

study

He found the plaque

of c o l u m n a r

The

the name

el liptical

one-half

not only with othe r s
jacent

extensive

he found the

s e c tio n in

one-half

bird,

an

v ar i a b l y

in c r o s s

rows

and

After

constituted

commonly
pig,

densa.

to use

of

was
rest

ma c u l a

situated towards

densa.

The Golgi

the b a s a l

of the distal tubule it was

side

of

s ituated

the nucleus.

reported

densa.

in the

that the position

The

the

absence

cytoplasm

of a b a s e m e n t m e m ­

of the

cells

of the

macula

30
d e n s a was
by the

ce ll

separated from
membrane.

thought to f a c i l i t a t e
the

macula

tu zz i

d e ns a

that of the j u x t a g l o m e r u l a r

This

the

a b s en ce

exchange

of the b a s e m e n t

of m a t e r i a l s

and the j u x t a g l o m e r u l a r

(1952) found the

macular

m a c u l a d e n s a to be l a r g e r

t u bu l a r

than the

apparatus

between the

apparatus.

ce l l s

area

membrane

to be

cells

only
was
of

Dina and M a r -

stratified

and the

of contact with the

af f e r e n t

arterio le.
The function of the m a c u l a d e ns a is
(1946)

r e g a r d e d the m a c u l a

chem o-receptor,
(1949,

influencing

the

Malpighian bodies

s ug ge s ted an angiotrophic

The collecting

of the

prim ary

rol e for

arched

collecting

periphery

tubules

arched

collecting t ubules,

The

sensory

area,

densa

Schloss

probably

vessels.

a

Becher

r eg ul at e d the flow of

of the kidney.

Okkels

(1950)

the m a c u l a densa.

tubules

The collecting
comprised

a

yet c l e a r .

co nt rac tion of the

1950) thought that the m a c u l a

blood through the

in the

d e ns a as

not

or

are
initial

c o n s i d e r e d by all
collecting

and the p a p i l l a r y

collecting
ducts

of the

ducts

tubules.

of the
co r t e x

cortex
to f o r m

or

authors

tubule, the
ducts

straight

of Bellini.

Huber

(1932,

of the

rabbit's

collecting

to be

1935) found the
kidney united

tubes

which p a s s e d

31
through

the

kidney it
tubules
thus

c o r t e x without

was

that

received further

The collecting

regular

c o l u mn a r

granules

or

tubules

cells,

striations

or

ovoid f o r m s .

ing

epithelium

their

lesser

tubules

straight

medullary

were

s ubs ta nce

The

to f o r m

tubule

cells

diameter

diameter

of the

increased

tubules

spherical

(1952)

cells,

described

of the collecting
epithelial

strictly

were

ce lls

was

lin­

animal s

as

only by

tubules.
lining

(1954)

described
s trai gh t

as

the a r c h e d

stated that the
located in the

making

collecting

up m o s t
tubules

in s u c c e s s i v e f as hion in the
p api l l a r y

4 0 |jl in the m e d u l l a r y

the tubules

the

cuboidal.

Greep

The

another

of

distinguishable

sixteen to twenty l a r g e

tubule
as

nuclei

in the domes tic

tubules.

of the p y r a m i d s .

finally

specific

r el a t i v e l y l a r g e

and F i e b i g e r

cor­

devoid of

in the human kidney were

s t a ted to fuse with one

pyramids

protoplasm,

s t a t ed that the

These

ducts

lined by v ery

s t r a i g h t collecting

rays.

and the collecting

in g e n e r a l ,

light-colored

collecting tubules

collecting

were found,

of the human kidney w e r e

The

of the

(1954)

of p r i m a r y

through the

connecting

the

co r t e x ,

In the human

while p as s in g

with c l e a r

height f r o m

Greep

of the

and with

c o m po s ed of polygonal,

branches.

b ra n c h e s

Trautmann

of the

further

a limited number

joined in the p e r i p h e r y

formed

tex.

observed

r ecei ving

fused.

ray.

ducts.
The

The epithelium was

cuboidal

32
in the p r o x i m a l
am eter

portion,

increased.

presence

but the

Trautmann

of n um e r o u s

cells

i n c r e a s e d in height as

and F i e b i g e r

fat d r o p l e t s ,

the d i ­

(1952) d e s c r i b e d the

especially

in o lde r

dom e s t i c

anim als.

The p a p i l l a r y

ducts.

Lang ham

the

epithelium of the p a p i l l a r y

tall

columnar fo rm

those

of the

the

sm aller

cells

opening

ducts

r e t a i n e d the

of

two-layered,

changing

epithelium in

to t r a n s i t i o n a l

near

the

into the pelvis.

of Be llini.

organ from

(195 3) doubted the

After

d if ferent

one p y r a m i d ,
ex is te nc e

examining

animals,

of openings

and not

of Bellini.

of the
u ri n e

excreted

existence

and

some

skeptical

ducts.

directly

routine

serial

es p ec i a l l y the

was

of openings

a few hundred

the i n v e s t i g a t o r was

n o r m a l kidney the

t h eli um

characteristics

(1952) d e s c r i b e d the

of the human kidney of all ages,

the

o th e r

a

collecting tubules.

as

D uran-Jorda
d ucts

(1942) found t h a t while

duct of the bovine kidney a c q u i r e d

T r a u t m a n n and F i e b i g e r
the p a p i l l a r y

etal.

of the

sections

sections

of this

ca t, which had only
about accepting

the

He was of the opinion that in

d ialysed through a p a p i l l a r y
into the

renal

pelvis by the

epi­
ducts

33
The b a s e m e n t m e m b r a n e

Bensley
structureless
of the
brane

and B e n s l e y

and the

sim ilar

McManus

r e l a t e d to the

thought that

it was

Pease
ture

and

s uggested a p a s s i v e

chitinous

(1950) and P e a s e
the

same

They o b s e r v e d the connective

overlying

membrane

epithel ia

appeared

rather

entirely

mem­

of the

role f o r

epithelium.

it,

McManus

in n at u r e .

b e t w e e n that they thought the b a s e m e n t
by the

This b a s e m e n t

studied the h i s t o c h e m i c a l f e a t u r e s

of the b a s e m e n t m e m b r a n e

t u bu les .

epithelium

than Bo wma n ’s capsule.

activity of the tubular

and B a k e r

of a thin

i n t e r p o s e d betwee n the

framework.

to but th inn er

(1948b)

tubules

(1930) noted the p r e s e n c e

reticular

renal b asem ent m embrane,
closely

uriniferous

basement membrane

tubules
was

of the

(1955b) found the
all

tissue

along the u r i n i f e r o u s
cells

membrane

so few

and f a r

m u s t be m ain ta ine d

than by f i b r o b l a s t s .

homogeneous

struc­

The b a s e m e n t

even under high

resolution.

The P a p i l l a

ing the
Bloom

Langham

et a l . (1942) o b s e r v e d t r a n s i t i o n a l

surface

of the p api lla in the bovine kidney.

(1952)

e p ith el ium

s t a t e d that in the human kidney the

of the

ducts

of Bellini

epithelium

Maximow and

s i m pl e

continued on to the

cover­

columnar

surface

of the

34
papilla
that,

in the

with the

t h eli um
to

area

was

cribrosa.

exception of the
replaced

columnar

Trautmann

on the

pig

and the goat,

r enal

rat

was

relatively

long

The I n t e r s t i t i a l

Kirkman

the

o b s e r v e d that the papilla

lying

in the

of f i b r o b l a s t s

of the kidney of the

cell

Schloss
human kidney

groups

intermediate

or

Becher's

Neumann

points

(1949)

where

of the g l o m e r u l i

Becher

(1949,
in the

was

The

found at
loop

of the medulla.

ce l l s

groups

the connective

of cells

tissue

in the

was

abundant.

cell

groups

of the human kidney.

1950) found th es e
immediate

rat.

of Henle' s

r e p o r t e d the lack of i n t e r t u b u l a r

vicinity

human kidney

zone

and m a c r o ­

albino

and f i b r o b l a s t s

(1946) found the i n t e r t u b u l a r
at the

ureter.

Space

of that p or tio n of the thick ascending limb

Intertubular

in the

space

c o n c e n t r a t i o n of m a c r o p h a g e s

which was

ep i ­

cuboidal

and extended into the

(1943) found the p r e s e n c e

in the i n t e r s t i t i a l

level

stated

transitional

papilla by t w o - l a y e r e d

and Schmidt-Niel s en (1952)

of the k a ng ar o o

greatest

the

(1952)

epithelium.

Vimtrup

phages

and F i e b i g e r

cell-islets

neighborhood of the

situated

in the

af f e ren t

arterioles

35
and. o t h e r

cortical

arterial

vessels

and called

them

' “p a r a p o r t a l

cell-

i s l e t s . 11
Regarding
Schloss

(1946)

p l ete ly

isolated

function.
regulate

Cell

the function of th e s e

suggested
and

Becher

that those

(1949,

1950) thought that

unit of Goormaghtigh

the

or

human kidney

in the

angle betwee n a f f e r en t
Becher

containing

nerve

Ham
gation
the

of

(1950)

glomerulus

efferent
tainty

cell-groups,

which wer e

tissue

those

com­

had an endocrine

cell-groups

might

of the kidney.

s oc le pla s mo di um

(1949),

of

s mall

and B e c h e r

(1949,

ce l l s

of Goormaghtigh)

(cells

and e f f e ren t a r t e r i o l e s

r e g a r d e d this

cluster

of

s mall

1950) found

of the g l o m e r u ­

ce lls

as

an a r e a

receptors.

cells

their

the p r e s e n c e

of a curious

with pale nuclei between the

proper,

arterioles.

about

a cluster

(1953) d e s c r i b e d

small

cells

Malpighian bodies

(1946), Neumann

in the

lus.

Becher

s u r r o u n d e d by connective

blood flow through

Schloss

intertubular

in the

Ham

nature,

was

function,

or

aggre­

m a c u l a densa

concavity between the
of the

lit tle

af f e r en t and the

opinion that t h e r e
n o m en cla tu re .

and

was

no c e r ­

36
Blood V e s s e l s

Ma c C a l l a m
dog

and cat,

(1936) found the

excl usively f r o m

verae

Ol i v e r

(1939) found a f ew pathologically
m a m m a l i a n kidneys,

found to i n c r e a s e

with age

Fitzgerald
arteries
per

Lo o m i s

No a r t e r i o l a e

MacCallum

(1939) and

changed g l om er u l i

numbe r

in the

in the

of such g l om er u l i

was

10 p e r

presence

of two main

cent of the h o r s e s

renal

and 5

examined.

(branches

in the p e r i t u b u l a r

(1942)

and Gouygou

of i n t e r l o b u l a r

c a p i l l a r y plexus

(1949) found

arteries

that t e r m i n a t e

without the

in the human kidney to occu r

interposition

in old age

and in con­

of d i s e a s e .
Grahame

and ca mel
the kidneys

(1944a,

similar

g e n e r a l plan of the

(1950) o b s e r v e d

n eig hb ors .

arterial

tree

arterial

in the

elephant

d is t r i b u t i o n in

mammals.

supplied definite

with t h e i r

1944b) found the

to the

of o th e r

Cowdry
teries

in

and J e t t - J a c k s o n

of a g l o m e r u l u s )

arising

and d i s e a s e .

each kidney

dogs

Lu dwig ’s b r a n c h e s
directly

an i m al s .

and the

rectae

eff erentia.

(1940) d e m o n s t r a t e d the

supplying

c e nt of the

ditions

found in those

va s a

rectae

normal

were

the

arteriolae

parts

that the individual

of the kidney

interlobular

and did not

ar­

a n a s t om os e

37
T hese

arteries

w e r e found to be

one was

occluded the t i s s u e

arteries

w e r e found to be o c c a sio na lly

the

phrenic,

adrenal,

Picard
in the

be

arterioles

valvular

or

a circular

suff e r ed

acutely.

when

These

a n a s t o m o s e d with b r a n c h e s

capsular

thickening

' 1e n d - a r t e r i e s 1';

of

arteries.

and Chambost
s ituated

(1952) o b s e r v e d

at the o ri g i n of the

of the j u x t a m e d u l l a r y g l o m e r u l i .

It a p p e a r e d

to

in n a t u r e .

Ruotolo
of v a r i o u s

intercostal,

on it

et a l . (1930) and P i c a r d

dog and cat

afferent

depending

typical

(1950) d e s c r i b e d

types

in the

the p r e s e n c e

tunica in t i m a

of blocking

mechanisms

and tunica m e d i a of the

arteries

of the human kidney.
Montaldo
podic b r a n c h e s
origin

(1951) found a l o z e n g e - s h a p e d l u me n of the monoof the

in the kidneys
Christensen

arteriolae

rectae

Anastomosis
served

of man,

and low i n t e r l o b u l a r
dog,

and

(1952) d e s c r i b e d

verae

of the

aff e r e n t

and

at th e i r

rabbit.

the p r e s e n c e

and Ludwig's

arteries

arterioles

of a few i s o late d

in the dog's kidney.

e f f e r en t a r t e r i o l e s

were

also

ob­

in a few i n s t a n c e s .
Rotter

intima

arcuate

of the

regarded

as

thickenings

(1952)

considered

arteries

of the human kidney

arteriosclerotic
were

the cushionlike

changes

p r e s e n t in the f o r m

as

thickenings
normal.

These

by many i n v e s t i g a t o r s .
of a ring

inside

of the
w er e
These

the b r a n c h e s

38
of the

arteries

increased
were

and w e r e found to occu r

in s i z e

and n u m b e r

supposed to

corresponding

with the

reg ul ate blood

vascular

age

supply

territories

even in young c h ild r en ,
of the individual.

and blood p r e s s u r e

and to p ro m ote

but

They

in the

an economic

dis­

t r i b u t i o n of the blood.
Superfici al
by K a z za z

veins

of the

of

drained

stellate

into the

into the

r en al

veins

running

renal

vein.

the

deep

junction

veins

side

surface

of the kidney

tributaries

joined each

of s t e l l a t e

The veins

veins

in the cow and

and joined the i n t e r l o b u l a r

protruding

side

to

sheep

into the vein lumina at the
of m a n and dog.

directly

arranged

of the veins

and

The human

the dog but not
s t a r t e d as

so

spurlike

veins.

which contained many l a r g e

in the kidney

side

and emptying into the

et a l . (1953) d e s c r i b e d the p r e s e n c e

tissue

kidney had a

r adially

interlobular s .
sim ilar

injection

of the kidney were

and on the medial

connected them to the

system

Koester

nature,

which on the l a t e r a l

dog's

The ca t was found to have

along the

developed.

connective

It was found that the

interlobular

vein.

veins

projections

veins

N um ero us

kidney had a
well

demonstrated

and Shanklin (1951) with the help of vinylacetate

and c o r r o s i o n technique.
system

m a m m a l i a n kidneys w e r e

veins

of cushions

of

of a sinusoidal

ar c u a t e - i n t e r l o b u l a r

39
C i r c u l a t i o n th r ou gh the kidney

Bowman
(with the

exception of a

s u r r o un di n g
illary

tufts

fat,

small

of the l a r g e

Malpighian bodies.

into the

vessels)

From

c a p i l l a r y plexus

and finally l e ft the

renal

artery

quantity d i s t r i b u t e d to the c a ps u le ,

and the co at s

of the

found to p a s s
tubules,

(1842) noted t h at all the blood of the

there,

s urrounding

e n t e r e d the c a p ­
the blood was
the u r i n i f e r o u s

o r g a n through the b r a n c h e s

of the

r en al

ve in .
M o r i s o n (192,6) studied the
rabbit,
finer
was

monkey,

deer,

circulation.

sheep,

It was

supplied by the

ing into

The

supply w e r e

thought

and pig with

capillaries

artery

plexus.

ence

arteriolae

White
interm ittance

p a r e n c h y m a of the

from

to its
cortex

vessels,

the i n t e r l o b u l a r

might i t s e l f t e r m i n a t e by

The n o n g l o m e r u l a r

dog,

sources

ramify­

of n ut r i e n t

to be negligible.

solely by a r t e r i o l a e

of the

cat,

reference

of the eff e r ent g l o m e r u l a r

The m e d u l l a of the kidney was
supply

special

nutrient branches

interlobular

a capillary

c i r c u l a t i o n in man,

noted that the

and in addition o cca s i o n a l
arteries.

r enal

rectae

rectae

(1939) mad e
in n o r m a l

spuriae.

verae

and

re c e i v e blood

No evidence

of the p r e s ­

was found.

o b s er v at i on s

dogs

o b s e r v e d to

indicating

rabbi ts

as

all

the a b s en ce
of the

of

glomeruli

40
were

found open all

mi g h t o c c u r

the t i m e .

in e x t r e m e

sudden i n c r e m e n t

It was

conditions

of ci r c u l a t i n g

F o x and Jones

(1946)

was

in action,
Heggie

1947)

within the kidney of the
via

the j u x t a m e d u l l a r y

relatively
vessels

as

during

while

sta ted that the

only.

The c o r t e x

stimuli.

Barclay

via the v a s a

recta.

et a l . (1946,

1948) made

kidney of the

experimental

a n im al

rab bi t
a

vasa

ally i s c h e m i c .

conditions

decreased
studied
the

or

Under
was

such

entirely

c i r c u l a t i o n in the

results

of

Trueta

recta

suppressed.
rabbit's

et a l . (1948)

et al.

could be

an extensive
and ca m e

renal

thr ough the m ed ull a via the

could

of the p e r i p h e r a l

l e a s t in p a r t ,

t h a t in the

cortical

ci r c u l a t i o n

r e d i s t r i b u t i o n of blood flow

view of Heggie that b y - p a s s i n g

the

the

r a b b i t could be main ta ine d by a c i r c u l a t i o n

to a f f e r e n t

in

diuresis,

c i r c u l a t i o n was b y - p a s s e d .

glomeruli

circulation

or

(1946) ob­

the m e d u l l a r y

su p p o r t e d the

Trueta

hemorrhage

and Havar d

active

in a n u r i a

a v a s c u l a r by v a s o c o n s t r i c t i o n

in r e s p o n s e

severe

and M a e g r a i t h

and the c o r t i c a l
(1946,

such

that i n t e r m i t t a n c e

epinephrin.

s e r v e d that in the h u m a n kidney,
c i r c u l a t i o n was fully open,

s uggested

remain
arterial

(1946,
effected,

1947)
at

inve stigation on

to the

conclusion

ci r c u l a t i o n could be continued
while the

c o r t e x was funct ion­

the flow of urine
Schlegel

kidney,

and Moses

but wer e

indicating

in the

unable

a by-pass

ureter

(1950)

to co nfi rm

through the

41
j uxtamedullary
ing

of T r u e t a

glomeruli.

et a l . (1948) t h at the only

could c i r c u l a t e
cortex were
arterioles

M o r e and Duff (1951) c o n f i rm e d the find-

in

sufficiently l a r g e

t h os e

amounts

of the j u x t a m e d u l l a r y

and the c o r r e s p o n d i n g

vessels

vasa

through which blood

to b y - p a s s

glomeruli,

the o ute r

their

e f f e r en t

r e c t a of the med u l l a in s e ­

quence.
Mos es
m edullary
rabbit.

and Schlegel

c i r c u l a t i o n following ligation of the

renal a r t e r y

in the

They o b s e r v e d t h a t t h e r e was i s c h e m i a of the o ute r

and the j u x t a m e d u l l a r y
contained blood.
with v e s s e l s
tissues.

penetrating

renal

Arteriovenous

and v a s a

observed

recta

direct

the ca ps u le f r o m

of the ca psu le

cortex

of the m ed ull a

vascular

connections

the p e r i r e n a l

and p e r i u r e t e r a l

and p e r i h i l a r

tissues

resulted

ischemia.

anastomoses

Simkin et al.

(1948) in j e c t e d g l a s s

diameter

into the

m ortem,

and the

c a t e d by

recovery from

than the a v e r a g e
on living

glomeruli

They also

Stripping

in co m p l e t e

made

(1952) found p r e s e r v a t i o n of the j u x t a -

renal

arteries

spheres

of n o r m a l human kidneys p o s t

ex i s t e n c e of a r t e r i o v e n o u s
the

diameter

ren al

vein of

of a c a p i l l a r y .

anesthetized

90p to 440fj. in

r a b bi ts

anastomoses

spheres
Similar

and dogs,

and

was

indi­

many t i m e s
experiments
recovery

of

greater
were
spheres

42
m easuring

5 Op to

circulation,
those

indicating the

animals

in the

18Op in d i a m e t e r

Similar

existence

obtained f r o m

of a r t e r i o v e n o u s

arteriovenous

h um an kidney by B a r r i e

Christensen

was

anastomoses

et a l . (1950)

the venous

anastomoses

were

demonstrated

and Pompeiano

(1952) did not find a r t e r i o v e n o u s

in

anastomoses

(1951).

in the

canine kidney.

Lymphatic

P eirce

(1944) d e s c r i b e d the l y mp ha t i c s

l a r g e blood v e s s e l s
and p e r i r e n a l

in the kidney of the

ly mp h a t i c s

Lymph c a p i l l a r y networks
the B o w m a n ' s
the

ca ps u le

rabbit.

The

cortical

w e r e found to connect with each o the r .
were

seen in the i m m e d i a t e
Valves

vincinity of

w e r e lacking

but w e r e p r e s e n t in the l a r g e

trunks

in

of the

sinus .
Rawson (1949)

the

which accompany the

dog and

of some g l o m e r u l i .

renal parenchyma,

renal

Vessels

glomeruli,

intertubular

r e p o r t e d the

and about the

ab s ence

of lymphatic

channels

a f f e r e n t and ef f e ren t a r t e r i o l e s ,

in

and the

capillaries.

Nerv es

Bradford
sels

of the

dog

(1889)

studied the inne r va tion of the

and found both v a s o - c o n s t r i c t o r

r en al blood v e s ­

and v a s o - d i l a t o r

43
fibers,
of the

the f o r m e r being well
existence

developed.

of c o n s t r i c t o r f i b e r s

No evidence

in the vagus

was

nerve

obtained

supplying

the kidney.
Gruber
kidney

(1933) concluded a f t e r

received fib ers from
O be r ling

nerve
teriole

plexus

review

the vagus

(1944) noted the

nerve

existence

of l i t e r a t u r e

that the

as well.

of a highly developed

aro un d the p r e g l o m e r u l a r p o r t i o n of the af f e r e n t a r ­

of the human kidney.
Szabo

(1948) m ad e

concluded that p os s i b l y
among the

epithelial

an extensive

vagal m o t o r

cells

adventitial

glomerulus.

cat

were p re s e n t

H a r m a n and Davies

supply o c c u r r i n g

in the

Nerve

endings

o b s e r v e d in the

veins ,

in the m e d i a of a r t e r i o v e n o u s

of the

convoluted t u bu les ,

sinus

Throughout the

s pace of the g l o m e r u l u s
el ab o r a t e

(1948a,

r en al

components w e r e found to p e n e t r a t e

vaded throughout with a f a i r l y

glomerulus.

endings

v e i n s , and epithelium.

The p e r i v a s c u l a r

were

and r a t ,

a rich nerve

in conjunction with a r t e r i e s ,
parenchyma,

nerve

and

of the convoluted tubules.

In the kidney of the
1948b) d e m o n s t r a t e d

review of l i t e r a t u r e

the

was p e r ­

complex of n e r v e f i b e r s .

adventitia of the
anastomoses,

and i n the p e r i v a s c u l a r

arteries

in the

and

epithelium

t i s s u e of the

44
Knoche
various
the

(1950) o b s e r v e d

vessels

glomeruli,

and a t e r m i n a l
the

Mitchell
ric hly

vessels,

(1950a,

innervated.

supply f r o m

the

were

interlobar

nerves.
or

cells

or

or

cells

were

tor

nerves

1950b,

of the

as

in the

convoluted tubules,
connective

tissue.

1951) found the kidney of m a m m a l s

nerves

accompanying

the

a

scanty

arteriolae

the m ed ull a d i r e c t l y f r o m

Nerve f i b e r s

epithel ial

Ma l m e j a c

in the

and the i n t e r t u b u l a r

found to e n t e r

b ead ed f i l a m e n t s

reticulum

The m ed ul l a was found to have

arcuate

No n e r v e s

nerve

nonmedullated n e r v e f i b e r s

w e r e found to t e r m i n a t e

b r a n c h e d or f u s i f o r m

endings

on the b a s e m e n t m e m b r a n e .

nerve
rectae.

the main
as f r e e ,

between the

Intrarenal

ganglia

confirmed.

et a l . (1951) found the
r enal

vessels,

s ynapses

of the

in the ganglia of the

vasoconstric­

solar

plexus.

MATERIALS AND METHODS

Source

For
This

this

adults;

from

a

Cattle:
16 months

to

Department,
Sheep:
from

the

the

Ten

Thirteen

Eleven

(nine c a s t r a t e d

from

were

and five females) ;

Rapids,

used.

males

all

Michigan,
and four females);

the m e a t l a b o r a t o r y ,

Animal

Husbandry

State Uni ver sit y.
(five m a l e s

dissection laboratory

Pigs:

and six f em a l e s ) ;

3 to24 months;

Seven

and two f emales );

4 to

24 months;

of the D e p a r t m e n t of Anatomy

and

of the D e p a r t m e n t of Pathology.

(four

males

and t h r e e females);

3 to7 months;

the m e a t l a b o r a t o r y .
Dogs:

from

males

at Gr and

Six (four m a l e s

the autopsy l a b o r a t o r y

from

castrated

slaughterhouse

Michigan

animal s

included:

(five

8 years;

sixty-nine

meat laboratory.

Goats:
from

study the kidneys f r o m

group of a n i m a l s
H orses:

of Animals

the

Nine

(six m a l e s

departments

and t h r e e females);

of Physiology

45

and Surgery

2 to

36 months;

and Medicine.

46
C ats:

Thirteen

and eight f e m a l e s ) ;
P hy si o l o g y ,

castrated

7 weeks

to

m ales,

5 years;

P a r i s i t o l o g y , and Sur g e r y

T h es e
to be f r e e

(two

animals

from

were

three

from

uncastrated

the d e p a r t m e n t s

males,
of

and Medicine.

in good n utr iti onal

condition and a p p e a r e d

disease.

T echniques

F i x a t i o n and P r o c e s s i n g

The kidneys
from

the f r e s h l y kille d

mixture

f o r four

fixed t i s s u e s
of B a r r i e
insured
tions

were

anatomic

anatomy

split longitudinally

animals,

six days.

Tissues

soon a f t e r

Small blocks

obtaining blocks

continuity of the

cl o s e l y followed.

of the m e d u l l a

their

and w e r e fixed in F.A.A.
w er e

cut f r o m

dehydr ation and f u r t h e r p r o c e s s i n g .

(1953) f or

was

vessels

for

to

of the

removal

(Lavdows k y ' s )
the well-

The technique

f r o m fixed kidney t i s s u e s

r enal

tubules

in m i c r o s c o p i c

The method took into

so that the tubules

sec­

account the g r o s s

and t h ei r

could be followed continuously f r o m

which

accompanying

the papilla to the c o r t i c o -

m e d u l l a r y junction.
Small
thick)

were

and the

blocks

(about

cut in the

papillary

ducts

1 centimeter

vertical
in

wide

plane of the

such a m a n n e r

as

and 0.5 c e n t i m e t e r

s t r a i g h t collecting
to include

tubules

a p or tion

of

47
tHe p e l v i s ,

the

the kidney.
tu bu l e s .

p a pi lla ,

This

plane

Sometimes,

divided t r a n s v e r s e l y
to ma k e
to

them

show the

Four

blocks

each

animal.

cross
were

step as
1.

parts)--4
2.

gave

in c a s e

of l a r g e

better

s ections

the

cortex,

a longitudinal

into two halves

sm a lle r for

animals,

at the

of the c o r t i c a l

of t i s s u e s

capsule

of

th es e blocks

were

c o r t i c o m e d u l l a r y junction
Blocks

were

and m e d u l l a r y

d i f fe r en t p a r t s

were

and the

s e c t i o n of the u r i n i f e r o u s

sectioning.

obtained f r o m

T h e s e blocks
step by

the medull a,

also

structures.

of both kidneys

then dehydrated

made

of

and i n f i l t r a t e d

follows:

N or mal butyl

alcohol

and 95 p e r

cent

ethyl

alcohol

(equal

hours.
N o r m a l butyl

alcohol No.

1 - - 4 hours.

3.

Normal

butyl

alcohol

No.

Z--16 hours.

4.

N o r m a l butyl

alcohol

and

52° C p araf fin

5.

52° C p ar af f i n

No.

1--2

6.

52° C p a r a f f i n

No.

2— 2 hou rs .

(equal p a r t s )--

2 hours.

7.

56°C p ar a f f i n No.

The i n f i l t r a t e d
two p a r t s

of

tissues

h ou rs .

3 -- 1 8 h o u r s .
w e r e then

embedded in a m i x t u r e

T i s s u e m a t 1 and one p a r t of 56° C paraffin.

1 Fisher

Scientific

Company,

Pittsburgh,

Pennsyl vania.

of

48
Sectioning

All

s ections

were

cat

at

Staining

seven m i c r o n s .

and Mounting

H a r r i s ’ hematoxylin and ethyl
stains.

Wei g e r t

collagenous
stain

was

and Van G i e s o n ' s

tissues

stained

the

s ections

Measurements
proximal

collecting

w er e

ples

of those

ments
renal

of the t r a n s v e r s e
corpuscle

only th os e
the

structures

vertical

and u r i n a r y

structures
plane.

diameter

poles

in the

was
same

routine
e l as tic

(1951)

and

r et i cu l um

of r e t i c u l a r f i b e r s .

of the

duct w er e

renal

corpuscle,

of Henle' s

made f r o m

the light

were

made

loop,

at t h e i r

section of the

to be cut through the

s e c tio n of the

r enal

sam ­

Measure­

centers,

judged by the p r e s e n c e

glomer­

straight

random

microscope.

of the longitudinal

which a p p e a r e d

This

used f or

Gridley's

thin limb

s ee n under

and g l o m e r u l u s

as

mounted in gum d a m a r .

of the d i a m e t e r

and p a p i l l a r y

used

of Measuring

convoluted tubule,

tubule,

w er e

demonstration

Methods

ulus,

stains

and m u s c l e f i b e r s .

employed f o r

The

eosin w er e

and f r o m
center

in

of both v a s c u l a r

corpuscle.

49
Several
urements
places
the

trials

of the

in the

average

were

s am e

same

as

measurements

ticular

well

and hence

as

species

structure

gave

of all

same

in different
was

as

average

well

animal

slides.

good as

from

measurements

species.

as

decided to take

structure

the final

in that

with twenty
in the

it was

of a p a r t i c u l a r

m e a n of the a v e r a g e s
s am e

structure

of ten m e a s u r e m e n t s

m easurem ents,

the

ma d e

as

ten m e a s ­

taken at different
It was found that

that

of the twenty

an a v e r a g e

each animal.

of ten
The

in different an imal s

m easurem ent

of

of that p a r ­

RESULTS AND DISCUSSION

Gross

The m a m m a l i a n kidneys
mida l)
l obes

or
or

multilobar
pyramids

present

d is t i n c t

any

present

twice the width.

distinct

in one kidney.

conical

The hilus

funnel-shaped,

others

s ents

of

are

(1953) as

horse

others

do not.

elongated,

bean-shaped,

Some papillae

is

The

the

is

t h at of the pig.

almost

of the

are

narrow

r e n a l pelvis

is

ex te r na l l o ­

(1873) and Sisson

right one.

card.'1

The left

The location of

The kidney in longitudinal

50

and

right kidney p r e ­

on a playing

than the

a

of the p y r a m i d s

smooth and without

d e s c r i b e d by Chauveau
Mthe h e a r t

and

majores.

b e a n - s h a p e d while the

and n a r r o w e r

of

section of the kidney p r e s e n t s

minores.

lo n g e r
like

to the numbe r

loca ted a l m o s t in the middle

kidney is
hilus

(unipyra-

Some m u lt ilob ar kidneys

wide and flattened.

the

shape .which was

and G r o s s m a n

is

unilobar

a p p e a r a n c e with the papillae

The left kidney is
a

as

The length of the kidney is

and divided into ca ly ce s

The kidney
bation.

smooth,

The longitudinal

into the c a l y c e s

while

is

lobation.

multipyramidal

projecting

described

s u p e r f i c i a l lobation while

superficial

medial border.

are

(m ul tip y r a m i d a l ) according

The kidney of the pig
without

Anatomy

section

51
is

found to be

were

not

concave

evident.

as

The

inner

The

described
m ajor

superficial

anterior

the

left kidney,

renal

similar

an terolateral
pyramids

project

into

but the pelvis

is

are

to the
an

of the kidney.

s u r fa ce

presents

of the

pig.

appearance
and m i n o r

with the
ca ly c e s

lamb

kidneys.

kidneys

lobation.

The longitudinal

appearance

the

renal

elliptical
near

a well-marked
The

cavity located

on

the medial b o r d e r .

a deep f i s s u r e
s u rf a ce .

which is

situated

On section,

The ca ly ce s

are

majores

In

the

found to
are

present

absent.

ex t e r n a l

lated

This

lobation.

out and the papillae

minores.

a

absent.

The kidneys of the sheep and the goat a r e
and without

(1953) as

m a m m a l i a n fetal

extensive

and p y r a m i d s

medull a f o r m s

elongated and p r e s e n t s

e a sily mad e

calyces

of the

and G r o s s m a n

p a r t of the d o r s a l

are

the

part

calyces

p a r t of the v e n t r a l
the hilus

not p api lla ted

into the pelvis

ox is

r i g h t kidney is

the

on the

central

and m i n o r

lobation

in the

o r g a n is

by Sisson

The kidney of the

hilus

the

ridge which p r o j e c t s

p r o j e c t i o n is
crest.

u nilobar

renal

are

The

situation of the hilus

section

Elias

in which the

assumed
individual

that

is

into the pelvis.

like that
a unilobar
The

sec tio n in a p a i r

sheep probably had compound

lobes

major

(1944) o bs erv ed the m ult ipa pil -

of the med u l l a in longitudinal

Elias

b ean - s ha pe d,

of the kidney p r e s e n t s

c r e s t projecting

absent.

smooth,

wer e

closely

united.

of

52
The kidneys
hilus

has

the

same

of the

dog

l o ca tio n as

s e c t i o n of the kidney p r e s e n t s
crest

projecting

ridges.

bling

dog El i a s

a unilobar
The
ca ly ce s

The longitudinal

appearance

renal
are

The

with the

c r e s t presents
absent.

r en al

curved

In the G e r m a n

(1944) o b s e r v e d m ul t i pa pi l l at e d kidneys

resem ­

that of the p i g .
The kidneys

hilus

and m i n o r

smooth and b e a n - s h a p e d .

that of the pig.

into the p elv is.

The m a j o r

shepherd

are

is

of the

cat a r e

s i tu ate d in the middle

of the kidney

gives

a very

are

of the medial b o r d e r .

d i s t i n c t unilobar

elongated p a p i l l a p ro j e ct i n g
calyces

smooth and b e a n - s h a p e d .
The

The
section

a p p e a r a n c e with a

into the pelvis.

The m a j o r

single

and m i n o r

absent.

Blood V e s s e l s

The
into

a numbe r

a s c e n d the
pyramids
teries.
the

These

artery

of l a r g e

renal

over

arcuate

teries,
the

renal

which they
arteries

at

main v e ss e ls ,

the kidney at the hilum

branches,

columns

arteries

arising

enters

the i n t e r l o b a r

send ar chi ng b r a n c h e s ,

n um e r o u s

extend

arteries.

These

to a pos ition level with the b a s e

do not an as t o m o s e

regular

and splits

smaller

intervals

of the

the a r c u a t e

with one anot her .

branches,

and at a l m o s t

r ad i a l l y between the

interlobular
right angles

medullary

rays.

ar­
From
ar­
to
These

53
arborize
into the
t i c al

the

of the

of the kidney

the i n t e r l o b u l a r

numerous

side

it divides

glomerulus.

The

that

capsule.

The

glomeruli

divide

capillary

size,

into the

m ed ull a,

supplying

Efferent

vessels

m a i n l y to the blood
some branches

cortex

the

interlobular

ing

arteries

aro un d the
interlobular
the b a s e

veins,

and

veins

the

which

rectae

of the p y r a m i d s .

the

cor­

an a f ­
enters

capillary

the

efferent

glomer uli b r a n c h

to

and the kidney

the j u xta m ed ulla r y
of long

straight vessels

spuriae,

which descend

of the p y r a m i d s .

zone of the c o r t e x contribute
and also

and the outer

veins.
along

These
the

tributaries

At the

Venae

the

to f o r m

components

capsule

return

the

to f o r m

branches

send

rays.

stellate

join to f o r m

arteriole

convoluted tubules

n u m er o us

tubules.

s mall

through

tubules

a tassel

the tubular

draining the

receive

cortical

cortical

in the c e n t r a l

unite to f o r m

a f f e ren t

emerging f r o m

into the m e d u l l a r y

pass

recombine

arteriolae

some

each of which is

neighboring

supply of the

venules

the

the

send

subdivides

r ep ea t e dl y , f or m i n g

of n e a r

the

from

the

efferent v essels

arising

arise,

The loops

supplies

and

arteries

As

and

efferent v essels

a network

Small

stems

to a g l o m e r u l u s .

corpuscle,

arteriole.
form

As

arteriole

renal

loops

surface

ca ps u l e .

labyrinth,

ferent
the

near

combine

path of the
from

rectae,

veins,

to f o r m

correspond­

anastomotic

c o r tic o m e d u l l a r y
arcuate

m a r g i n of

border

which

nets

the

extend a c r o s s

which r e t u r n blood f r o m

the

54
medulla,

open into the
interlobar

arcuate

veins

the

mids,

in company with the

the kidney w h e r e

veins

veins.

are formed

they join to f o r m

cortex,
nels

but

an e l a b o r a t e

apparently

them

around the

arteries.

ing f r e e l y
uli.

with

All l y mp ha t i c s

the kidney
Valves
are

are

at the hilum
lacking

present

of lymphatic

arcuate,

Large

and

fat,

renal

the l a s t

vein.

of the

the

vessels,

capsule,

two c o m m u n i c a t ­

vessels
aortic

of the

chan­

particularly

do not enter

end in the l a t e r a l

t ru nk s

of

in the

lymphatic

network,

lymphatic

in the lymph v e s s e l s

in the l a r g e

renal

vessels

u nd erlie s

The ly mp ha tic s
into

the

and i n t e r l o b u l a r

anastomotic

the p e r i n e p h r i c

drain

to the hilum

Vessels

A second plexus

each othe r .

of a r c u a t e

between p y r a ­

channel,

in the medulla.

in an i r r e g u l a r

and a t h i r d p e n e t r a t e s

but

none

arteries,

a main

network

acco mpa ny the i n t e r l o b a r ,

surrounding
rich

is

confluence

which c o u r s e

co r r e s p o n d i n g

Lymphatic

There

Through

the g l o m e r ­

which leave

lymph nodes.

r en al p ar e n c h y m a ,

renal

sinus.

Ne rv es

The n e r v e s
of the
fibers,

symp ath eti c
which

are

of the kidney a r e

der iv ed f r o m

division of autonomic
mostly

unmyelinated,

nervous
e n te r

the

ren al plexus

sytem.

the kidney

The ner ve
with the

55
renal

vessels

N er v e

fibers

tubular
among

among

especially

The

is

al so

above

d om es tic

cells,

generally

has

description

and Jenni ngs
man

(1953)

may be

The

f i b ro u s

ca ps u le

fibers.

The

Any unusual
perplasia
there

is

surface

is

adherence

in this

the

as

the

only.

tuft.

The

of the kidney of
scope of this

Hence for

study

a detailed

(1873),

Reighard

and Sisson and G r o s s ­

Anatomy

Capsule

is

covered with a delicate

composed of collagenous

ea s ily lifted f r o m

throughout the kidney
a fatty l a y e r ,

of Bowman's

supply the kidney.

a l . (1943),

of the kidney

(tunica f i b r o s a )

capsule

terminate

to.

Mi croscopic

The o u t e r

Other s

the work of Chauveau
et

arterioles.

end adja ce nt to the

layers

anatomy

short

structures

Z ie t z s c h m a n n
referred

to

ef fer ent

of the g l o m e r u l a r

gross

b e e n made

of the o rgan,

(1933),

parietal

s paces

and

segment.

c o n s i d e r e d to

confined to the m i c r o s c o p i c

gross

in the

d e s c r i p t i o n of the

animals

afferent

in the p r o x i m a l

and in the p e r i v a s c u l a r

nerve

the

the b a s e m e n t m e m b r a n e

the j u x t a g l o m e r u l a r

vagus

is

penetrate

cells,

capsule,

the

and t e r m i n a t e

regard

is

c a ps u l a

the kidney when n or m al.

indicative

substance.
adiposa.

and elastic

Outside
From

of a f ibrous

hy­

the tunica f i b r o s a
the in ne r

s ur f a ce

56
of the

capsule

fine

kidney, f o r m i n g

strands

the

interstitial

The tunica f i b r o s a
was found to be
outer

elastic
nous

fibers.

and

c o n s i s t e d of dense

The in n e r

the in n e r

of the kidney.

The i n t e r n a l

appearance

uriniferous

tubules.

tunica f i b r o s a

exceptions

were

Of the
was

and well

dog and

and a few

composed of loose
cells

tunica f i b r o s a .

surface

of the

inner

were

The

collage­
also

reticular

the p a r e n c h y m a

layer presented

a

due to the i m p r e s s i o n left by the underlying
adiposa was

lying j u s t

of the h o r s e ,

ox,

to that of the pig in g e n e r a l .

and in ne r

layers

outside

the

and n e r v e s .

sheep,

goat,

dog,

The following

of the tunica f i b r o s a ,

developed in the h o r s e ,
in the

both l a y e r s

sheep and goat,

the

inner

equally developed

and the i n n e r l a y e r

ab­

cat.

In c o m p a r i s o n with the
the

was

fibers

The

noted:

dem arcated

sent in the

collagenous

w er e found to p e n e t r a t e

of the kidney

o ut e r

better

the

of the pig

of two di s t i nc t l a y e r s .

smooth m u s c l e

of the

The ca p s u l a

sim ilar

extend into

of the kidney

and was found to contain blood v e s s e l s

The c a ps u l e
and c a t was

A few

layers

corrugated

layer

thin l a y e r

layer

tissue

tissue.

and composed

in the in n e r

from

connective

of the capsule

reticular fibers.

observed
fibers

thick

thick l a y e r

of loose

cat had m o r e

pig

all

othe r

smooth m u s c l e

domestic

in the inner

animals
layer

except
of the

57
tu nic a f i b r o s a .

The

ox,

muscle

that

portion

of the ca ps u le

the

it f o r m e d

almost
(Plate

and goat contained

a dis tinc t
I).

muscular

so much
layer

The dog had l e s s

smooth

in the

than the

deep er

pig,

and

cat had none.
Tereg

(1911)

smooth m u s c l e
of the
other
was

sheep,

sheep

in the

T r a u t m a n n and F i e b i g e r

deeper

portions

of the

Of the c u r r e n t

only one to mention

authors

(1952) d e s c r i b e d

capsule

and ox but did not mention amounts

species.

the

and

or

of histology

smooth m u s c l e

of the kidney
refer

t e xts ,

to any
Gr ee p

in the capsule

(1954)

of the hu­

m a n kidney.
While
in the

o ut e r

all

layer,

Reticular
of the

kidney

s pe ci me ns
those

fibers

of the

tubule .

of u r i n i f e r o u s
segments,

a

sheep

some

s e e m e d to have
least

el as tic f i b e r s
the

most.

developed in the

capsule

cat.

The kidney is

of two

of the

s e e m e d to be

The

number

observed presented

U r ini fe rou s

Tubules

a compound tubular gland made
tubules.

secretory

Each
tubule,

u ri nif e r ou s
or

nephron,

up of a l a r g e

tubule is

composed

and a collecting

58

PLATE I

Capsule of the kidney of the goat, showing the capsula fibrosa and
capsula adiposa.

Note the muscle cells

capsula fibrosa.

H. and E.

1237x.

1.

Capsula adiposa

2.

Capsula fibrosa:

3.

Proximal

4.

Distal convoluted tubule

in the inner layer of the

Nine months

(a) outer layer;

convoluted tubule

old.

(b) inner layer

59

X

*
I

*»

om
PLATE I

S8

PLATE I

Capsule of the kidney of the goat,
capsula adiposa.
capsula fibrosa.

showing the capsula f ibros a and

Note the muscle cells in the inner layer of the
H. and E.

1237x.

Nine months

old.

1.

Capsula adiposa

.

2.

Capsula fibrosa:

fr’
(a) outer layer; (b) inner layer

3 . Sjtniyfjqqrial
4.

plotted tubule

Distal c<J&^^Sl-U.ted tubule

59

#•»

•*

0

U

f

r»
*
PLATE X

60

The

nephr on

The n e p h r o n is
and is
the

the

renal

and the

secretory

The
spherical

renal

the p r o x i m a l

the

a vascular

vessels

and le av e,

pole,

whe re

the dilated

form

Bowman’s capsule.
renal

The

tamedullary

adult

average
renal

pole,

u ri n i f e r o u s

r en al

opposite
tubule

which encloses

to this

is

the

a

and the g l o m ­

co r p u s c l e .

invests

in a

Each

the

renal

glomerular
urinary

glomer u lus

to

of the pig kidney was found to be p e a r -

transverse

The

tubule begins

m a r k e d by the point where

and n e a r l y

corpuscles

with the

to

the

that

of

the loop of Henle,

size

diameters

were
of the

of the

137p and
renal

advancement of age,

156p,

cortical

r es p e c t i v e l y ,

co r pu s c l e
reaching

and jux-

was

as

observed

maximum

at the

stage.
In

ilar

It c o n s i s t s

Bowman’s capsule

Malpighian or

corpuscle

shown in Table II.
to i n c r e a s e

tubule.

Bo w m a n rs capsule,

the g l o m e r u l u s .

corpuscle has

shaped.

unit of the kidney

convoluted tubule,

Each u r i n i f e r o u s

known as

together form

The

of the u r i n i f e r o u s

corpuscle.

expansion,

enter

and functional

convoluted tubule.

tuft of c a p i l l a r i e s ,
erulus

structural

p or t i o n

corpuscle,

d is tal

the

horse,
of

the

the
pig.

shape
The

of

the

transverse

renal

corpuscle

diameter

of

was

the

sim ­

cortical

61

T A B L E II
DIAMETER OF THE URINIFEROUS TUBULE,
WITH EPITHELIAL HEIGHT
(in microns;

average based on ten counts per animal)

R enal
Corpuscle
Specie s

No.

Cor­
tical
Di.1

J uxtamedullary
Di.

Glome rulus

Cor­
tical
Di.

J uxtamedullary
Di.

Horse

10

178

191

159

166

Ox

13

181

173

150

141

11

153

147

1Z3

1Z3

.

6

158

157

1Z7

1Z6

. .

7

137

156

114

130

9

1ZZ

1Z4

98

100

13

96

106

8Z

87

. . .

Sheep

Goat

Pig

Dog .

Cat . .

1
Z

Diameter .
Epithelial height.

62
TABLE

P roximal
Conv oluted
T ub ul e

Di.

E.H.2

II ( C o n t i n u e d )

Thin Seg­
ment

of

Henle 1s

Distal

Straight

Convoluted

Collecting

T ub ul e

T ub ul e

Loop
Di.

E .H.

Di.

E.H.

Di.

E.H.

Papillary
Duct

Di.

E.H.

56

15

30

6

50

9

66

14

138

55

50

13

26

5

45

8

53

12

91

33

45

11

22

4

36

7

39

10

59

18

48

11

24

5

38

8

42

12

57

22

45

14

23

5

36

8

50

14

96

21

39

11

22

5

33

7

37

9

54

13

41

14

21

4

30

7

37

5

60

14

63
renal

corpuscle

was

178p

and

that

of

the

juxtamedullary

was

191m-.
The

renal

had a d i a m e t e r
tamedullary

181p in the

cortical

also

region

sheep,

of the pig.

the

shape

of the

The d i a m e t e r

r enal

renal

t i c al

corpuscle

r egion was
In the

m ore

or

of the

puscle

was
Ham

In the cat,

true for

(1953)

cortical

was

renal

147(jl.

shape

of the

ren al

sim ilar

corpuscle

In the

goat,

in the

the d i a m e t e r

lZZ\x

of the

of the ju xta m e d u l l a r y was

larger

cor­

pig,

corpuscles

and hence w e r e the
that

the

dog,

and cat.

was found that the

above

reverse

was

and j u x t a ­
124(j.,

ren al

re­

cor­

106p.

corpuscle

than those n e a r e r

the j u x t a m e d u l l a r y

observed

and

cortical

s t a ted that the j u x t a m e d u l l a r y

development

corpuscle

of the c o r t i c a l

of the dog w e r e

generally

the h o r s e ,

and goat it

173p in the j u x ­

and the d i a m e t e r

The d i a m e t e r

96p and that

tigation it was

the

corpuscles

probably b e c a u s e
during

and

and

158(jl and in the ju x t a m e d u l l a r y region was 1 5 7 p .

human kidney was

form

pear-shaped

spherical.

r en al

s pect ively.

was

dog and cat,

less

medullary

pear-shaped

corpuscle

1 5 3 p and t h a t of the j u x t a m e d u l l a r y was

was
the

of

of the ox was

r egion of the kidney.

In the
to that

corpuscle

wer e

oldest.

of the

the ca ps u le ,
the f i r s t
In this

s t a t em en t of Ham was
In the c a s e
was

true.

of the

ox,

to

inves­
also
sheep,

The j u xt a m ed ul l a r y

64
renal

corpuscles

cortical

of the ox,

an a f f e r e n t
capillary

the

arteriole

were

capillaries
The

anastomoses

dog,

observed
and

or

structures
of the

The t h r e e
visceral

of

tuft a lobulated

ca ps u le is

of the
axial

axial
wer e

animal s

space

closely

ap­

ox,

no

between the

of the

amount of connective

w e r e found to

surround

and his tioc yt es

of the glomer u lus .

observed

membranes

of

under investigation.

Fibroblasts

sheep,

of B o w m a n ’s

investigation,

of the g l om er u l us

s m all

space
also

In this

such a n a s t o m o s e s

reticular fibers

horse,

prim ary

layer

ar­

between the c a p i l l a r i e s

and man.

to contain a

o b s e r v e d in the

glomerulus

to give the

of Bowman's

tuft of the g l o m e r u l u s .

Similar

af fe r en t

The p r i m a r y branching

tends

to d e m o n s t r a t e

intercapillary

capillary

of the

cat,

glomerulus

Collagenous

also

layer

This

tuft.

of the
made

again.

arteriole

visceral

of the

pig kidney was

the

the

(1955) o b s e r v e d

glomerulus

were

than the

supplied by

arteriole.

ab r u p t division of the

subdivides

capillary

Hall

attempts

near

The

plied to the

the

sm aller

of a tuft of c a p i l l a r i e s

and d r a i n e d by an efferent

which in t u r n

capillaries

tissue.

consists

tuft is f o r m e d by the

structure.

the

and goat w e r e

corpuscles.
The g l o m e r u l u s

teriole

sheep,

goat,

of the

capsule

in the axial
dog,

of

and cat.

g lom er u lus

covering

space

co n s i s t s

the g l o m e r u l u s ,

65
the

endothelial

layer

ment m em brane
Th e s e
found to be
The f i n e r

three

Bowman's

or

or

parietal

uous

The

layer

of t hese

c a ps u l e

envelops
space

observed

to have

space

was

in the

horse

observed

double l a y e r e d .

of the capsule

of the

capillaries

s ituated between the

and the

visceral

pig and other

a flattened

animal s

squamous

found to be l a r g e r
and cat;

could not be

The

and the

inner

of the g l o m e r ­

two l a y e r s

is

contin­

convoluted tubule.

Both the p a r i e t a l
of the

the loops

investigation.

deep invagination of the

It is

the wall

of this

were

II).

is f o r m e d by the

layer forms

with the p r o x i m a l

(Plate

base­

structures.

animal s

membranes

tubule.

and the

of the g l om er u l us

and other

three

capillaries,

above two

light m i c r o s c o p e

capsular

in the kidney

glomerular

membranes

end of the p r o x i m a l

visceral

ulus.

primary

p r e s e n t in the pig

with the o r d i n a r y

outer

the

lying between the

details

expanded

lining

of Bowman's

under

equal

ox,

in the

capsule

i nves tigation were

epithelium.

in the goat,

and a l m o s t

layers

and

The

capsular

sheep;

dog,

as

smaller

c o m p a r e d to

that of the pig.
The j u x t a g l o m e r u l a r
of the

m e d i a of the

entrance
ent

into

arterioles

the

appa r at us

af f e r e n t a r t e r i o l e s

glomerulus,

b eco m e

large,

the

consists

of the modified

of the g l o m e r u l u s .

smooth mu s c l e

and p a l e - s t a i n i n g .

cells

Near

cells

of the

Their

nuclei,

the

affer­
i n s tea d

() 6

PLATE II

Kidney of the goat.
glomerular cells.

Note a
H

single-layered

and E.

6l2x.

macula densa and juxta­

Five months old.

1.

Juxtaglomerular apparatus

2.

Macula densa

3.

Glomerulus

4.

Parietal layer of Bowman’s c ap s ul e

5.

Visceral layer of Bowman's

6.

Endothelial cell of the glomerulus

7.

Basement membrane

capsule

of the glomerulus

67

¥
*

*?.

<

6

*•
%

*r

©

plate

II

66

PLATE II

Kidney of the goat.
glomerular cells.

Note a
H

and E.

single-layered
6l2x.

macula densa and juxta­

Five months old.

1.

Juxtaglomerular apparatus

2.

Macula densaj^

3.

Glomerulus

4.

Parietal layer of Bc^Sman's capsule

5.

Visceral layer of Bowman's capsule

6.

Endothelial cell^Btif the glome r u ^ s

7.

Basement membrane of the glc^^erulus

a

3

0

*
.
.
m

i

*5 * *

9
LI

19

••«*. .-> "^

U"

\

a

<fSi

*• kv*
'

'

r
I

#

a

d
P L A T E II

68

of being
taining

elongated,

myobibrils,

of t h e s e

cells

epithelioid
the

are

is

of the

usually

are

wall

and t h e i r

resemble

and tend to f o r m

c e lls .

the

c e lls .
These

an a s y m m e t r i c a l

of con­
shape

These
ce lls

myo-

thicken

cap on one

side

glomerulus.

found to be

round and

s t a i ns .

macula
of the

somewhat deeply

They w e r e found to be

dens a .

Along with th e s e

spindle-shaped
The

ox,

sheep,

the

goat

more

in s t e a d

Moreover,

epithelial

ca lle d j u x t a g l o m e r u l a r

The nuclei of the j u x t a g l o m e r u l a r

eosin

cyto pla s m,

contain g ra n u l e s .

such that they

cells

arteriolar

rounded,

structure

numerous

the

to Maximow

glom erular
vations

were

conclusions

the
were

not made
can be

were

of the
also

(Plate

stated that g r a nu le s

drawn

r egar ding

In the c a s e
appa rat us

w er e

this

of

were

ab s en t in the

In this

cells

According
are

of the
point.

absent

investigation,

in a two - month-old cat.

on young a n i m al s

some

of the h o r s e ,

of the dog and man.

of two y e a r s .

demonstrated

media

II).

(1952) the j u x t a g l o m e r u l a r

age

with the

observed.

a p pa r at us

to that of the pig.

of the kidneys

and Bloom

cells

cells

of the pig

(1951)

cells

in c h i l d r e n below

closely a s s o c i a t e d

of the j u x t a g l o m e r u l a r

than th os e

Goormaghtigh
juxtaglomerular

cells

pig kidney were

stained with hematoxylin and

of the j u x t a g l o m e r u l a r
similar

in the

modified cells

smooth m u s c l e

and cat w e r e

and dog,

ce l l s

other

juxta­
O bser­

s p e c i e s , so no

69
The
tubule

has

lined by

proximal
the

widest

simple

and the

or

and l a r g e

of c l e a r

tubule.

The b a s a l

striations

spaces

b etw ee n the

pars

This

recta,

structure

of the

p r o xi ma l

proximal

whose
cell

45p.

average

was

height was

somewhat

tubule.

the descending
same

called the
ray,

of the loop

tubule is

thick limb
as

course

called
of Henle.

that of the p a r s

tubule.

tubule

It was

the

the m e d u l l a r y

s t r a i g h t p or t i o n

The p r o x i m a l
of

enters

descending limb

the

the

which it o r i g ­

tubule is

as the

r e c t a is

or

of the p r o x i m a l

and continues

of the p a r s

is

a looped and tortuous

convoluta then

and c o ns t i t ut es

convoluta of the

diameter

ce l l s

proximal

This

striations,

the m i t o c h on dr i a

of the

of Henle.

It is

cell lining the p ro xi ma l

convoluted p o r t i o n
The p a r s

and is
abun­

The b r u s h b o r d e r

corpuscle from

straight,

The

ei t he r

basal

renal

convoluta.

pars

are

having

nuclei.

of the

becomes

the

end of the

m i t o c h o n d r i a of the

vicinity

spherical

convoluted

nephron,

cells

of a b r u s h b o r d e r ,

convoluted tubule p u r s u e s

immediate

inates.

placed

cell b ou nd a r i e s .

of the f r e e

in the

of any p o r t i o n of the

basally

covering

The p r o x i m a l

The p r o x i m a l

truncated pyramidal

by the p r e s e n c e

absence

cuticular

diameter

cuboidal

dant p r o t o p l a s m
characterized

convoluted t u b u l e .

of the pig was found to have

lined by

simple t r u n c a t e d

found to be

coarsely

granular.

14p.

The

These

an a v er a ge

pyramidal

cytoplasm

granules

cells

of the

w er e

more

70
or

less

tinct.

evenly
T h es e

they w e r e
cells
the

distributed,

grannies

not c l e a r .

masked
The

cells.

a p r o m i n e n t nucleolus
cont our

very

of the

irregular.

separate

The
the

horse,
cat,
ce l l s
and

nuclei w e r e
eosin

s ome

for

one another

Th es e

( Plate

structures

of the p r o x i m a l

ox,

equal in the

lining

the

smaller

in the

pig.

The l u m e n

goat,

ox,

ca t.

The

and

ox,

i n v e s t i g a t i o n was

greater

sheep,

almost

clusters

wer e found to be

to those

tubule was
and

as

of the

sm aller

of

in the

in the dog and

The height of the
equal in the cat;

c o m p a r e d to that

tubule was l a r g e r
dog;

rest

of the pig.

greater

(Table II).

equal in the

regular

was found to be

in the horse;

of the lumen in the
more

c h r o m a t i n g ra nu l e s.

tubule

similar

and dog,

of the p r o x i m a l

sheep;

contour

was

contained

p r o m i n e n t and f o r m e d a

sheep;

to that of the pig

tubule

and deeply

of the p r o x i m a l tubule

The d i a m e t e r

in c o m p a r i s o n

even towards

III).

u nder i n ve s t i ga t i o n were

and goat;

not d i s ­

but in some

Each nucleus

stained

was

or

circular,

s ta ins .

deeply

each cell.

middle

w ere

and t h e r e f o r e

usually b a s a l ,

lumen of the p r o x i m a l

general

animals

cell b o u n d a r i e s ,

were

The b r u s h b o r d e r

cluster

separated from

and

and

striations

situated in the

These

sta ine d with h em at o xy l i n

the

nuclei

they w e r e found to be

apex of the

The

and hence b a s a l

of the

in the h o r s e ,

and

sm aller

in the

r e s t of the

animal s

of this

than that of the pig.

Some of the

71

PLATE III

Kidney of the pig.

Note the brash border forming a cluster on each

cell of the proximal tubule.

H. and E.

1.

Proximal tubule

2.

Brush border

3.

Distal tubule

4.

Interstitial

2460x.

Seven months old.

connective tissue cells

PLATE i n

71

Z

PLATE III

Kidney of the pig.

Note the brush border forming a cluster on each

cell of the proximal tubule.

H. and E. 2460x.

Seven months old.

1.

Proximal tubule

Z.

Brush border

3.

bSl
Distal tubwle

4.

Interstitial connective tissue cells

t

72

P L A T E XU

73
nuclei

of the

goat.

The b r u s h b o r d e r

of the

ce l l s

animals
served

cells

in the

than in the

pig.

pig.

No

Basal

Fat

tubules

iron-containing
rat.

was

case

were

oval in the

were

along

of the

They did not f o r m

striations

cat

w er e

arranged

tubule in the

globules

of the

tubule

uniformly

investigation.

G rafflin

the

proximal

of the p r o x i m a l

under

proximal

of the

more

the

apices

rest

of the

clusters

as

(Plate

was

ob­

di s t i nc t in the dog

o b s e r v e d in the cells

and dog

ox and

of the

IV).

(1942) noted a deposition of yellow to golden-brown
p i g m e n t in the

such p i gm en t was

epithelium of the p ro x i m a l
o b s e r v e d in any of the

tubule

animals

of

under

i n ves tigation.

The loop of Henle.
ing
is

and an ascending
divided into

arm

The loop of Henle

arm

connected by a

a thick and a thin

segment.

co n s i s t s

s h a r p bend o r

of the

distal

arising
tively
being
of the

the thick ascending

from
short

very

convoluted tubule.
the

oute r

dis tan ce
s hort.

cortical

into

renal

recta

The loop of Henle of those
r enal

corpuscles

passes

recta
tubules

only a r e l a ­

the thin s egment

of this

other hand, the loop of Henle

of the

corpuscle

It

of the p r o x i ­

is f o r m e d by the p a r s

the medulla,

On the

juxtamedullary

arm

curve.

The thick descending

of the loop of Henle is f o r m e d by the p a r s

m a l tubule;

of a d e s c e n d ­

descends

deeply

into the

loop
tubules
medulla

74

PLATE IV

Kidney of the cat.
proximal tubule.

Note the fat globules in the pars
H. and E.

700x.

recta of the

Four months old.

1.

Fat globule

2.

Pars

recta of the proximal convoluted tubule

3.

Pars

convoluta of the proximal

4.

Brush border

5.

Basal

of the proximal

convoluted tubule
convoluted tubule

striations of the proximal convoluted tubule

P L A T E IV

74

I

PLATE IV

I

Kidney of the cat.
proximal tubule.

S

1.

Note the fat globules in the pars

H. and E.

700x.

Four months

recta of the

old

£

Fat globule

2.

Pars

recta of the proximal convoluted tubule

3.

Pars

convoluta of the proximal convoluted tubule

4.

Brush border

5.

Basal

of the proximal

convoluted tubule

striations of the proximal convoluted tubule

75

P L A T E IV

76
often

as f a r

medulla
the

is

as

the

mad e

descending

tubule;

bles

of the

am eter

than

the

tubule.

protoplasm

is

less

than

proximal

to thin

s e g m e n t of H en l e' s
The d i a m e t e r

having

an a v e r a g e

eosinophilic
more

or

less

of Hen le' s
to the
basal

clear

loop.

Z3\x.

striations

animals

stains

tubule

smaller

and is

resem ­
in d i ­

made

up

into the lumen.
less

The

eosinophilic
the

thick

seg m e n t of Henle' s

lined by flattened

The

with a s p h e r i c a l

loop in the pig

cells

were

tubule.

nucleus

.A b r u s h b o r d e r

m ar k ed l y l e s s
The cytoplasm

bulging

of the nucleus

epithelium

gave
was

was

into the lumen

a wavy contour

absent

and the

i ndis ti nc t.

under

following few

tubules

of

that of the

The t r a n s i t i o n f r o m

of the p r o x i m a l

lumen of the tubule .

is

lining

abrupt.

It was

This bulging

The g e n e r a l
all

loop is

height of 5p.

than those

ascending

segment

d is tal

tubule.

of the thin

kidney was found to be

loop r e s e m b l e s

and t h e r e f o r e

d is tal

loop within the

epithelial

with nuclei which bulge

granular
or

or

of the

The

of the

The thin

the p r o x i m a l

epithelium

part

segment.

epithelial lining

distal

either

greater

s e g m e n t of Henle' s

of a f l a t t e n e d

either

The

up of the thin

thick

proximal
that

papilla.

structure

of the

i n ve s tiga tio n was

exceptions

w ere

noted:

thin limb
similar

of Henle's

to that

loop of

of the pig.

The

77

The

d iam eter

the h o r s e ,

ox,

compared
greater
the

ox,

sheep,

The p a r s

somewhat
cells

segments,
the

the

was

sheep,

The

smaller

dog,

wider

greater

and cat,

cell height was
in the

sheep

than that

and n a r r o w e r

ascending

convoluted

o ver

in
as

slightly

and cat,

of the pig.

of the

in the dog,

(pars

an extensive

crowded

convolutions
d is t a l

tubule

pig,

in

the cat

in the

distal

recta

convoluted tubule

and the p a r s

of Henle' s

loop of Henle l e av es
convoluta),
its

nuclei

of the

makes

s ph er i c al

much l e s s

re­

tangential

con­

In this

convoluted tubule become

than in the proxi ma l

are

The

to the g l o m e r ­

The distal tubule,

the ma c u l a densa.

cuboidal with

tubule

distal

loop.

the med u l l a r y

and r e t u r n s

origin.

is

convoluta.

region with the afferent a r t e r i o l e .

distal

are

distal

of the g lom e r u l u s ,

and constitute

in the

The

ascending limb

which the nephron has

r egion of cont act the
d ens ely

pars

thick

t u r nin g to the neighborhood

cells

loop was

of equal height to those

convoluted t u bu l e.

s e g m e n t of the

ulus f r o m

the

in the

(Table II).

and goat,

recta forms

becomes

tact

of H en le' s

equal.

co m p o s e d of two

r ay ,

and

smaller

loop of Henle

The d i s t a l

thick

and

having

of the

horse,

being

horse,

animals

The l u m e n
the

and. goat;

thin limb

to that of the pig

in the

other

of the

or

There

tubule.

a r e fewer

The

ovoid nuclei.

b ro ad than those

cells

in

The

that line

78
Ihe p r o x i m a l

tubule

in the

tubule.

distal

proximal
border

tubule.

is

to be
The

tubule.

the

nuclei

each o th e r

than those

two p r o m i n e n t

was

The

ab s e n t

under
it,

the

The

spherical

striations

in the

similar
distal
sheep;

a s im ila r

c o m p a r i s o n the
ox and goat;

nuclei

as

the

stained

not c l e a r .
w er e

than that of
granular

and

of the p r o x i ­

the distal

sm aller

were

closer

coarse

to

presented

ch r o m a t i n

The b r u s h b o r d e r

tubule
pig.

tubule was l a r g e r

sm aller

8p in height.

not very distinct.

to that of the

and

was found

The nucleus

in the

of all

animals

Compared with

in the h o r s e ,

in the

epithelial height was
and

spaced

and b a s al ly placed although

and darkly
were

was

than those

These

a few d if f e r e n c e s ,

of the

irregular

cytoplasm

w er e f i n e r

nucleoli

equal

in the

contour l e s s

pig kidney

epithelium

of the p r o x i m a l tubule.

in v e s t i g a t i o n was
diameter

so acidophilic

tubule in the

also noticed.

and the b a s a l

cl os e ly

p r e s e n t although a b r u s h

lined by cuboidal

were

and goat;

equal

distal

cell b o u n d a r i e s

Except f o r

are

more

tubule.

g ra n u l e s

were

appear

not n e a r l y

striations

and its

The nuclei

a few oval

granules.

is

convoluted tubule.

eosinophilic;

one o r

was

was l a r g e r

the p r o x i m a l

mal

of the

The tubule

lumen

less

Cytoplasm

in the d i s t a l

diam eter

36p.

the nuclei

Faint basal

lacking

The

and hence

dog and cat.

greater

sheep,

ox,
By

in the h o r s e ,

dog,

and cat.

79
The l u m e n
and goat;

of

and

The
of the

the

smaller

macula

distal
the

slightly

taller
was

faintly
were

coarse

was

to that of the

ci al f e a t u r e s .
although the
irregularly
of the

as

pig

crowded

were

almost nongranular

macula

of the distal

tubule.

The

The cells

of the

with the j u x t a g l o m e r u l a r

the

same

The

d e nsa

animal s

cells.
The

stained nucleolus

under

and

investigation was

which p r e s e n t e d

densa in the h o r s e kidney was
w ere

not distinct,

as

tubule.

of the h o r s e

The cells

simple to

was

spe­

str a t i f i e d
very

epithelium

than double that of the

and did not take the

change f r o m

stratified

sim i­

some

the nuclei wer e

The height of the

a p p e a r e d to be m o r e

cells

of the

The ce lls

contained very few fine g r a n u l e s .

( Pl ate V).

other

the b a s e .

cells

except in the h o r s e ,

densa

wards

section.

s i n g l e - l a y e r e d and a p p e a r e d to be

associated

d e ns a in all

stratifications

lining

seen in c r o s s

and contained a dark

The m a c u l a

macula

sheep,

g ra nu l e s.

The m a c u l a
lar

ox,

constituted about half of the wall

stained around the nucleus.

ce l l s

spherical

in the h o r s e ,

dog and cat.

r e s t of the

cl osely

of the

chromatin

greater

d e n s a in the pig

than the

cytoplasm

nucleus

in the

was

m acula densa were

m acula densa
The

tubule

convoluted tubule

fo r m i n g

cy t o pl a s m

distal

abrupt.

of the m ac ula densa
stain,

especially t o ­

stratified

epithelium

80

PLATE V

Kidney of the horse.
400x.

Note the stratified macula densa.

Adult.
1.

Macula densa

Z.

Straight collecting tubule

H. and E

81

*£> «

*3

<91< ;£jV

M *S2s'

■’

PLATE

V

80

PLATE V

Kidney of the* horse.

Note the stratified macula densa.

1
400x.

Adult.
1.

Macula densa

Z,

Straight collecting tubule

I

H. and E

81

ft

v
-7.T

ft

PLATE

V

82
M cM a n u s

(1 9 43 ) and O kkels

(1950) d e m o n s t r a t e d the r e v e r s e d

p o s i t i o n of the G o lg i e l e m e n t in the m a c u la d e n s a of the c a t and
m an.

G o lg i e l e m e n t s

w e r e not stu d ied in th is w o r k .

The c o l l e c t i n g tu b ule

T h e c o l l e c t i n g tu b ule c o n s i s t s
ing tu b u le , the

of the a r c h e d o r in itia l c o l l e c t ­

s t r a ig h t c o ll e c t i n g tu b u le, and the p a p illa r y duct or

duct of B e l l i n i .
T h e d is t a l

c o n v o lu te d tubule i s jo in e d d i r e c t l y to an a r c h e d

c o l l e c t i n g tu b u le , into w h ic h it d i s c h a r g e s .
short ste m s

that b r a n c h f r o m

the

T h e s e a r c h e d tu b u les a r e

s i d e s of the s t r a ig h t c o ll e c t i n g

tu b u le s l o c a t e d in the m e d u lla r y r a y .

The a r c h e d tu b u les a r e

situ ­

a te d in the c o r t i c a l la b y r in th .
T h e s t r a i g h t c o l l e c t i n g tu b u le s l i e
m o s t of the

s u b s t a n c e of the p y r a m id s .

s id e by s id e and m ak e up
The s tr a ig h t tu b u les join

w ith one a n o th e r in s u c c e s s i v e f a s h io n in the p y r a m id s to f o r m
f i n a l l y th e l a r g e p a p illa r y d u c ts .
tu b u le s j o in and it r e a c h e s
c o l l e c t i n g tu b u le s
v i e w e d in c r o s s
b u les

a m a x im u m in the p a p illa r y d u c ts.

a r e n e a r ly
sectio n .

The d ia m e t e r i n c r e a s e s a s the
The

a lw a y s v e r y r e g u la r in ou tlin e when

The e p ith e lia l c e l l s lin in g the a r c h e d tu ­

and th e p r o x im a l p o r tio n of the s tr a ig h t tubule a r e

strictly

83
cuboidal.

The

cells

increase

in height as

the tubules

increase

in

d i am et e r .
The p a p i l l a r y
foram ina

s i tu ate d

ducts

on the

The a r c h e d

tubule.

It was

cuboidal

and w i d er

characteristic
spherical

or

nucleolus

and

oval,

the

arched

cat

resembled

cell

deeply
the

collecting

and ox,

of the nephron.

and b a s a l l y

some

Except f o r

horse

a r c h e d tubule was faintly

granules

placed.

height was

diameter

tubules

sm aller

greater

than the

epithelium with quite
more

The cytoplasm of

stained and devoid of

The nuclei

were

Each nucleus

either

contained a

stained c h r o m a t i n g ra n u l e s .
and cell height,

of the h o r s e ,

that of the pigand

diameter

The lu me n of the a r c h e d tubule was

than that of the d is tal tubule.

lining the

through the

The a r c h e d collecting tubule

lined by a simple

regular
cells

tubule.

a greater

cell b o u n d a r i e s .

ca ly ce s

of the papilla.

o b s e r v e d to have

distinct

the

surface

collecting

in the pig kidney was
distal

empty into the m i n o r

ox,

The d i a m e t e r

in the

sheep,

in the h o r s e

other

sheep,
was

goat,

goat,

greater

dog,

and s m a l l e r

structures
dog,

and

in the

and cat.

in the

of

rest

The
of the

animal s .

The

straight

collecting

s t r a i g h t collecting tubule

of the

tu b u l e .

The

g ene r al

pig kidney was

s t r u c t u r e of the

s i m i l a r to that of

84
the

arched

tubule
lar

collect ing

was

50p and

contour

straight

which b e c a m e
The

the

and was

collecting

cytoplasm

The

under

following

The
the h o r s e
(Table

II).

in the

ox,

greater
and dog.
goat,

The

horse,
cell

fat

droplets

in the

cuboidal

and f r e e f r o m

collecting

The

epithelium

the p api l l a r y

duct.

g ra n u l e s .

tubule

and each

ch r o m a t i n

s t r a i g h t collecting

sim ilar

straight

tubule

to that of the pig.

in the

in all
The

and F i e b i g e r

tubule was

goat,

dog,

in the hors e;

sm aller

more

in the

in the

distinct

in

and cat
and

smaller
was

sheep, goat,

in the h o r s e ,

ox,

sheep and cat.

(1952) noted the p r e s e n c e

collecting

greater

The lumen of the tubule

and

wer e

dis tinc t

sheep,

greater

and cat;

straight

collecting

and cat.

bou nd ar ie s

and l e s s

Trautmann

dog,

ox,

tubule.

regu­

distinct.

of the

sm aller

goat,

collecting

noted:

of the

and

collecting

and a few c o a r s e

were

cell height was

sheep,

and dog;

were

straight

The lumen had a

towards

stained,

than the a r c h e d

inve stiga tio n was

and ox;

The

and t a l l e r

faintly

structure

diameter

in the

wider

cell b ou nd a r i e s

d if fe r e n c e s

14[jl.

arched

stained nucleolus

The g e n e r a l
animals

was

of the

was lined by a simple

clear,

larger

contained a deeply

diameter

wider than the

tubule

was

The

cell height

somewhat

The n uc l e i w e r e

granules.

tubule.

tubules

esp ec ia lly

of n um e r o u s

that of

85

the

o ld e r

d om es tic

the

straight

animals.

collecting

No

tubules

The p api l l a ry d u c t .
duct

of the pig kidney

bule.

The

greater

diameter

than those

duct was

transitional
sitional

near

of the papilla.

epithelial

Zip,

papillary
large,

duct w e r e

dark-staining

and

the l u m e n was

regular.

were

general

noted:

II).

stained

situated

nucleolus

i n ve stiga tio n

of the

straight

collecting

The

duct on the papilla.
duct continued as

simple

p api l l a r y
tu­

duct wer e
papi lla ry

epithelium which beca me
This

the

to t r a n s i t i o n a l

duct was found to be

faintly

The

this

columnar

in

investigation.

height of the p api lla ry

of the

(Table

each n ucl eu s .

The

observed

abrupt.

tran­

epithelium

type of epi­
The d i a m e t e r

height of the p a p i l l a r y duct were found to be 9bp

respectively

spherical,

w er e

under

structure

that of the

The change f r o m

and the

animals

g ene r al

of the p a p i l l a r y

in the p a p i l l a r y

and

tall

opening

t h eli um

dropl ets

s t r a i g h t collecting tubule.

simple

the

epithelium

The

epithelial

of the

lined by a

of the

resembled

and

such fat

and

The tall

columnar

ce lls

and nongranular.

a l m o s t in the

chromatin

center

granules

The nuclei
of the

were

of the papillary

r e s e m b l e d that of the pig.

cell.

were
A

o bs e r v e d in

cell boundaries, w e r e distinct and the

structure

of the

duct in all

contour

of

animals

in

The following

exceptions

86

The
and

diameter

smaller

in the

of the

rest

epithel ial height was
in the

sheep,

papillary

dog,

duct was

papillary

of the

opening

the p a pi lla ,

almost

This
the

course.

papillary

(Plate

and

The

and

smaller

sheep the

epithelium f o r

a greater

VI).

The

columnar

simple

epi­

columnar

epithelium of

epithelium

was

extension of the t r a n ­

duct of the c a t kidney was

(1939) co ns i de re d

found

the t r a n s i t i o n a l

duct in the bovine kidney as

r ev e a l e d that the t r a n s i t i o n a l

duct in bovine kidney is

a normal

papillary

duct of the bovine kidney.
that t r a n s i t i o n a l

The

of the pig.

and Hallman

bovine kidney f o r

goat,

of the t r a n s i t i o n a l

et a l . (1942) found that only tall

the

ox,

and goat,

lined by a simple

Lang ha m

determined

ox,

and the t r a n s i t i o n a l

in the p a p i l l a ry

in ve stiga tio n has

in the h o r s e

investigation.

At the junction of the

duct

section

of the p a p i l l a r y

greater

on the papilla than that of the pig.

equal to that

La ng ha m
t h eli um

in the h o r s e ,

a beaklike p r o c e s s

epithelium

under

In the h o r s e ,

of the p a p i l l a r y

found i n longitudinal
sitional

an imal s

dog kidney was

throughout its

ep ith el ium

to be

greater

duct was

lined by a t r a n s i t i o n a l

its

t h eli um

of the

and cat.

distance from
duct

papillary

From

epithelium lines

a d is t a n c e of about

this
the

pathological.
epithelium

one

columnar

epi­

(Plate
cells

VII).
lined the

study it has
papi lla ry

1500p f r o m

the

of

been

duct of
opening

of

87

PLATE VI

Papillary ducts in the dog kidney.

H. and E.

320x.

One year

1.

Simple columnar

epithelium lining papillary ducts

2.

Beaklike process

at opening of the papillary duct

3.

Oblique section of the papillary duct opening

4.

Pelvis

old.

^

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87

PLATE VI

Papillary ducts in the dog kidney.

H. and E.

320x.

One year old,

1.

Simple columnar

epithelium lining papillary ducts

2.

Beaklike process at opening of the papillary duct

3.

Oblique section of the papillary duct opening

4.

Pelvis

t

88

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89

PLATE VII

The papillary ducts in the ox kidney.

H. and E.

320x.

months old.
1.

Transitional

epithelium

2.

Opening of papillary duct on the papilla

Nineteen

90

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89

t
PLATE VII

The papillary ducts in the ox kidney.
rn onths old.

H. and E.

320x,

^
1

Transitional

epithelium

2.

Op ening of papillary duct on the papilla

r

Nineteen

90

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w 2 i ti >t!i> -VvLv*5
o

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t ' i v ’35 .

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V

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PLATE

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91
the p a p i l l a r y

duct at which point i t

changes

to

simple

columnar

epi­

thelium .
D uran-Jorda
papillary

ducts

t h at u r i n e
ings

was

(1953) could not o b s e r v e

in the

(Pl ate

duct in the

renal

openings

of the

c a m e to the conclusion
epithelium.

ca t kidney w ere

The open­

o b s e r v e d in this

VIII).

The b a s e m e n t m e m b r a n e

The

and hence

d i al y sed through the p a p i l l a ry

of the p a p i l l a r y

investigation

c a t kidney

the

tubules

of the u ri n i f e r o u s

tubule

and the

ducts

excretory

are

enveloped ex­

t e r n a l l y by a w el l - developed and dis tinc t b a s e m e n t m e m b r a n e .
The p r e s e n c e

of the b a s e m e n t m e m b r a n e

tubule was

observed

structures

could not be

No d i f fe ren ce
tubules

was

of o t h e r

of the u r i n i f e r o u s

in the pig kidney although details

of its f i n e r

studied with the o r d i n a r y light m i c r o s c o p e .

noted in the b a s e m e n t m e m b r a n e

an i m a l s

of this

of the u r i ni f e r ou s

investigation.

The P a p i l l a

The papi lla i s
which is
surface
ings

l o ca ted

the

at the

of the papi lla is

of the p a p i l l a r y

apex of the

r enal p yr a m i d ,

the b a s e

of

c o r t i c o m e d u l l a r y junction of the kidney.
p e r f o r a t e d by f o r a m i n a which a r e

ducts.

This p e r f o r a t e d

area

of the

the

The
open­

papilla is

92

PLATE VIII

The papillary ducts in the cat kidney.

H. and E.

220x.

Seven weeks

old.
1.

Opening of papillary ducts

2.

Papilla

3.

Junction of transitional with simple columnar

epithelium

93

P L A T E Y in

92

S

E
PLATE VIII

The papillary ducts in the cat kidney.

H. a n j E.

220x.

Seven weeks

old.
1.

Opening of papillary ducts

2.

Papilla

3.

Junction of transitional with simple columnar

S

I

epithelium

93

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94
called

the

area

rmnor

reflects

epithel ium.
the

papillary
In the

found to
tion,

cribrosa.
over

This

surface

papillary

ducts
pig,

reflect

the

The t r a n s i t i o n a l

at t h e i r
the

over

the high t r a n s i t i o n a l

low and continued as
p api lla .

Af t er

came higher

a

o ver

entire

this

sides

the p api lla ry

epithelium

calyx m ino r

of

In the angle

of

was

reflec­

of the calyx m i no r became
d istance

along

the

low t r a n s i t i o n a l

sides

of the papilla

very

of the

epithelium b e ­

and apex of the papilla.

epithelium maintained

surface

angle of r e f l e c t i o n .

epithelium of the

some

calyx

on the papilla.

epithelium

and c o v e r e d the

with the

of the papilla.

s h o r t d istance

transitional

papilla never

openings

such f o r

papillary
the

epithelium blends

sides

of the

of the papilla and f o r m s

transitional
the

epithelium

The

a l mo s t uniform height

except on the

The height of the t r a n s i t i o n a l

sides

near

the

epithelium of the

a c q u i r e d the ori gin al height which lined the calyx

minor .
The papillae
similar

to that of the pig

the papi lla was
l o w er

of the

in the

higher

sheep,

othe r

animal s

investigation were

except that the t r a n s i t i o n a l

in the h o r s e ;

goat,

under

dog,

almost

and cat.

equal

epithelium

in the ox;

and

of

95
The I n t e r s t i t i a l

The i n t e r s t i t i a l
mal

connective

m a m m a l i a n kidney

forms

a fine

Connective

tissue

and c o n s i s t s

network between the

is

are

Some loose

tubules.

A. few

found only in the walls

connective t i s s u e

strands

blood v e s s e l s .

is

in terstitial

space

connective

tissue

scanty in the n o r ­
reticular

The t i s s u e

tissue

is

more

Collagenous

of the l a r g e r

which
abun­

and e l a s ­

blood v e s s e l s .

p r e s e n t between the p api lla ry ducts.

of connective t i s s u e
Fibroblasts

very

of delicate

dant in the p a p i l l a r y p or tio n of the medulla.
tic f i b e r s

Tissue

enter

the glomer u lus

and his tioc yt es

of the m a m m a l i a n kidney.

are

along the

also found in the

P r o l i f e r a t i o n of this

throughout the kidney may play an i mpor tant

part

in d i s e a s e .
The i n t e r s t i t i a l
by

reticular

the p api lla
numerous
the

fibers
(Pl ate

near

medull a.

large

IX).

histiocytes,

Re t i c u l a r f i b e r s

the blood v e s s e l s

blood v e s s e l s

tubules

of the pig kidney was

which extended f r o m

Collagenous

few v e r y fine
ous

space

fibers

es p ec i a l l y

collagenous

and the B o w m a n ’s
and m a s t

of the pig kidney.

cells

were

to the

w er e

also

occupied
apex of

w e r e found to be m o r e

also found to

spaces

of

s ur ro un d the

c o r t i c o m e d u l l a r y junction.

capsule

wer e

capsule

and in the i n t e r s t i t i a l

at the

fibers

the

mainly

o bs er v ed to invest
of the kidney.

A

the u r i n i f e r -

Fibroblasts,

noted in the i n t e r s t i t i a l

space

96

PLATE IX

Kidney of the pig stained for

reticular connective tissue.

distribution of reticular fibers
chyma and the glomerulus.
Seven months old.

in the interstitial

spaces

Note the
of the par en ­

Silver Stain (Gridley modification).

460x.

97

PLATE

DC

96

PLATE IX

Kidney of the pig stained for

reticular

distribution of reticular fibers
chyma and the glomerulus.
Seven months old.

connective tissue.

in the interstitial

spaces

Note the
of the par en­

Silver Stain (Gridley modification).

460x.

PLATE

IX

98
The

in terstitial

i n v e s t i g a t i o n was
were

The

similar

a b s e n t in the

intertubular

dog

cell

The p r e s e n c e
was

reported

195 0).

arterial

or

groups

where

These

not

the

Neumann

the

connective tis s u e

yet been

tubules.

are

and Becher

is

af f e r e n t a r t e r i o l e s ,
cell

groups

are

spaces

of the c o r t i c a l

They w er e

also found in the close

arteries

usually w er e found to be l a r g e r .

present

a regular

shape.

philic

and n o n g r a n u l a r .

The

oval.

arteries.

encapsulated.

Most of the cells

few which w e r e found to be

animals.

The

nucleus

vicinity

Those

did not

were polygonal

smaller

sur­

Some of the

The cells

cytoplasm was

was

were

region between

rounding

w e r e found to be

called

cell groups

and around the c o r t i c a l

groups

also

cortic al

of i n t er t u bu l a r

r e p o r t e d in the domestic
intertubular

in the

and other

The p r e s e n c e

these

(1949,

abundant,

ca ps u le

smaller

cells

situated in the human

of Bo w m a n ' s
the

under

in the human kidney

(1949),

groups

in t e r t u b u l a r

in the i n t e r s t i t i a l

u ri n i f e r o u s

animals

(Plate X)

groups

cell

In the kidney of the pig,
observed

cell

cell i s l e t s ’1 by B e c h e r ,
has

other

except that the m a s t

B e c h e r Ts cells

(1946),

neighborhood of the

' 'paraportal
cell

groups

intertubular

vessels.

of the

to that of the pig

of i n t e r t u b u l a r

at the points

immediate

tissue

and cat.

by Schloss

These

kidney

connective

except

faintly

and d a r k e r

a

eosino­
than

99

PLATE X

Kidney of the ox.

H. and E.

634x.

Six years old.

1.

Intertubular cell group or Becher's

Z.

Interlobar a r t e r y

3.

Proximal convoluted tubule

4.

Distal

5.

Thin limb of Henlers loop

convoluted tubule

cells

100

a m

.
&

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$« &

PLATE X

<*

99

2

£

PLATE X

Kidney of the ox.

and E.

634x.

Six years old,

1.

Intertubular cell group or Becher 's

2.

Interlobar a r t e r y

3.

Proximal

4.

Distal convoluted tubule

5.

Thin limb ^>f Henle's loop

convoluted t u b u l ^

cells

100

* *

•;>*>.

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i* *

k

<a T > -

*r

i

*

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, . a ( r ■•-:£

“. * *
k
;

; I -

J

'

•

.

o

^

>s?

_

^ £&
%
**vA« X&w-'Jf
^3

«* Si®** W G *
* > « A ‘m V

jET

* T

*

* ; w

•

7“
f
ft
*•'

<?&
.V *i*w s?.»
fe««M MR-■
r
SSP^Sl^
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PLATE X

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101
t h a t of the
erally
were

surrounding

spherical,
located

The nucleus

tubules.

although

either

a few oval

in the

presented

center

d a r k nucleoli

periphery

nucleus

the

nuclear

at ed

membrane.

at the c e n t e r

was found to be
Th e s e

cells

were

m ore

that of the pig
in the

Cell

sheep,

seen

of

small

in m i c r o s c o p i c

cell

The

space

cell

groups

with pale

proper,

s ections.

unit of Goormaghtigh,

along the

was

numerous

situ­

and b et t e r
the

cytoplasm than
developed

cat.

(Frontispiece)

nuclei between the macula

in the concavity between the

This

and

In the ox,

of the m a m m a l i a n kidneys

or

or five

between these nucleoli

w er e l e s s

socleplasmodium

cells

They

ch ro m a t i n granules.

and absent in the

eff e r e n t a r t e r i o l e s

Four

stained nucleolus

than those of the pig.

These

and dog,

and the g l o m e r u l u s

and the

deeply

gen­

of the cell.

approximation to each other

and contained m o r e

unit of Goormaghtigh or

densa

the

X).

were

noticed.

one c o r n e r

w e r e found to be m or e

eosinophilic

cells

also

appearance.

and f r e e f r o m

ox and h o r s e

goat,

nuclei were

o r towards

of the nucleus.

(Plate

A cluster

ent

A large

cell groups

of these

were found to be p r e s e n t

in close

unstained

developed in the

nuclei

a characteristic

c re sc e n t - s h a p e d
of the

The

cluster

of small

cells

Goormaghtigh* s cells.

affer­

can be
is

called

102

In the
c i ated

these

with the j u x t a g l o m e r u l a r

d en s a .
were

pig kidney

Goor maghtigh

found to be

ce lls

similar

cells
cells

of other

were found to be closely
and the cells
animals

to that of the pig.

of this

asso­

of the macula
investigation

SUMMARY AND CONCLUSIONS

1.
neys

of

Comparative

sixty-nine

d om e s t i c

pig being

c h o s e n as

s e c tio ne d

at

stains.

microscopic

Additional

animals

a type.

s ev en m i c r o n s ,
s ta ins

studies

w e r e made

of seven different

Tissues

wer e

on the k id­
spe ci es,

the

routinely fixed in F.A.A.,

and stained with hematoxylin and eosin

were

used to d e m o n s t r a t e

special

struc­

tures.
2.
domestic
layers,

The tu nic a f i b r o s a
an i m al s

an o ut e r

in this

of the

capsule

investigation was found to consist of two

and an i n n e r ,

with the

the kidney c a ps u l e had only one l a y e r ,
The o u t e r

layer

and a few e l a s t i c
collagenous
in all

extent in the

fibers,

and the

except the cat.
ox,

a distinct m uscular

pig,

exception of the cat, where
the inner la y e r being absent.

cons i s ted of dense collagenous

inner l a y e r

was f o r m e d by loose

with smooth muscle f i b e r s

In the

the m u s c l e f i b e r s

m en tio n the p r e s e n c e
goat,

tunica f i b r o s a

and r e t i c u l a r f i b e r s ,

animals

to f o r m

of the

of the kidney of the

layer.

sheep and goat,

of smooth m u s c l e

and dog.

103

and to a l e s s e r

w er e v er y numer ous ,
No r e p o r t s
cells

present

and appeared

have been found which

in the capsule of h o r s e ,

1 04
3.
pig

The j u x t a m e d u l l a r y

(1 56(a), dog

ones,

the

were

178(a,

in the

(124p),

cortical
137(a,

ox,

sheep,

were

173p,

147(a,

18lp,

153[a,

and

r a n ge d betwee n
the h o r s e ,

ox,

IZZp,

158p,

and

in the

and goat;

smaller

(137|u).

t h at c o r t i c a l

r en al

ones

as

co r t i c a l

dog,

and cat

reverse
renal

was

corpuscles

ones

renal

dog,

were

cor pus cle s
in

and cat,

co r p u s c l e s

(153(a),

tr ue

as

were

and goat (158(a),

(96(a) than that of the pig
any

w er e l a r g e r
ox,

renal

), sheep

have been found for

s e e n in the

sheep,

The c o r t i c a l

jjl

(191 jjl ),

s pe c i e s , being l a r g e r

in the

(122(a) and cat

corpuscles

The

J uxtamedullar y

smaller

(178(a), ox (181
dog

pig,

and the c o r t i c a l

191 |a in different

of the pig.

reports

No

medullary

in the

r es p e c t i v e l y ,

and

than the

the ju xta med ulla r y

respectively.

106p and

horse

in the h o r s e ,

r espectively.

where

in the h o r s e

larger

jjl

and 96[a,

157ja,

corpuscles

(10 6 ) wer e

corpuscles

and goat,

c o m p a r e d to th os e
larger

and ca t

renal

and

renal

sheep,

species

which stated

than those of the j u x t a ­
and goat in this

in ve s tiga­

tion.
4.
tical
the

ones

The j u x t a m e d u l l a r y
in the pig

cortical

159p> 98p,
(150(a) w e r e

(130(a), h o r s e

glomeruli
and

82p,

larger

and j u x t a m e d u l l a r y

g lomer uli

dog,

and cat wer e

In the ox the c o rtic al

than the j uxta medulla r y
glomeruli

than the c o r ­

(l66p), dog (100(a), and ca t

in the pig, h o r s e ,

respectively.

wer e l a r g e r

in the

ones

sheep w e r e

(141(a).
equal

(87(a),

11 4p,

glomeruli
The

(123(a),

cor tical
and m

1 05
the

goat they w e r e

127m- and

126p,

r es p ec tiv ely .

The juxtamedullary

( 1 6 6 p ) and ox (141(j.) w er e l a r g e r than those

glomeruli

of the h o r s e

of the pig

( 1 3 0 p ) , while in the sheep ( 1 2 3 p ) , goat ( 1 2 6 jj.), dog (lOOp),

(87( j.) they w e r e s m a l l e r .

and c a t

( 1 5 9 p-), ox (150(j.),

horse

t han t h o s e

of the pig

they

smaller.

were
5.

u nd er

the

goat,

equal

The c a p s u l a r

in the

7.

and
dog,

sheep;
as

of the

c e lls ,

and ca t was

compared

juxtaglomerular
8.

amounts

in the h o r s e

appa rat us

more

convoluted tubule was

and goat

(48p);

equal

in the

larger

in

and a l mo s t

in this

smooth muscle cells

apparatus
pig.

myoepi-

of the h o r s e ,

ox,

In the goat and dog,

numerous

and b e t t e r

(45p),

greater

sheep

animals

along with the usual

In c o m p a r i s o n with the pig

proximal

and cat;

in all

spind le- s h ap ed

to that of the
w er e

and r e t i c ­

to that of the pig.

The j u x t a g l o m e r u l a r

cells

of collagenous

and h istiocyt es.

aff e r e n t a r t e r i o l e

sim ilar

of the

of all animals

of Bowman’s capsule was

smaller

The j u x t a g l o m e r u l a r

media

thelioid

s mall

space

i n v e s t i g a t i o n contained some
of the

space of the glomerulus

along with f i b r o b l a s t s

ox,

g lomer u li

(114[j.), while in the dog ( 9 8 p ) and cat (82(x)

in ve s t i g a t i o n contained

6.

co r t i c a l

(123(0.), and goat (127(o) were l a r g e r

sheep

The i n t e r c a p i l l a r y

ular fibers

The

(45p);

the

and

smaller

the

developed.

diameter

in the h o r s e

sheep,

of the

(56p),
in the

ox (50p),
dog

(39p)

1 06

and ca t
was

(41p).

greater

in the

The height of the

in the h o r s e

ox (13p),

border

was

proximal

sheep

tubule

Fat

cells.

equal in the

a cluster

of the pig kidney,
were

were

observed

14p)

(14p);

and s m a l l e r

and dog

(lip).

The b r u s h

f rom

each individual

whe r ea s

striations

(pig,

cat

in other

uniformly a r r a n g e d

The b a s a l

globules

lining the tubule

(1 1ja.), goat ( l i p ) ,

found to f o r m

i n v e s t i g a t i o n they
of the

(15p);

cells

cell

animals

in the

of this

along the luminal b o r d e r

were m o r e

in the p ro xi ma l

distinct in the dog.

tubule of the cat and

dog.
9.

The d i a m e t e r

r anged between

2 lp

and 30p in different

the pig in the h o r s e ,
sheep,
ox,

dog,

sheep,

and cat.
and goat;

10.

ox,

and goat;

species,

equal to the pig in the. sheep;
pared

to that

horse

(9p);

equal in the

sheep

(7p),

dog

cat.

of the pig

in the h o r s e ,

smaller

wider

being l a r g e r

(8p), the

epithelial

ox (8p) and goat

and
ox,

cat

(7p).

sheep,

than the pig in the

and n a r r o w e r
tubule

in the dog.

ranged between

in the h o r s e ,

smaller

than

than the pig in the h o r s e ,

of the distal

and

segment of the loop of Henle

spe ci es , being g r e a t e r

to it in the cat;

The d i a m e t e r

(7p),

and

The lumen was
equal

and 5 Op in dif fer ent

greater

of the thin

ox,

and goat;

in the dog and cat.
height was

(8p);

and

greater

smaller

30p

Com­
in the

in the

The lumen of the distal tubule was

and goat;

and s m a l l e r in the

dog and

1 07

11.
was

The

macula

single-layered

stratified

and

dens a

except

in all

an imal s

in the h o r s e ,

contained m o r e faintly

found r e g a r d i n g

the

in this

in which it was found to be

stained cells.

stratified nature

investigation

No r e p o r t s

were

of the m ac ula densa of the

horse.
12.

C o m p a r e d with the pig

collecting
sm aller
The

tubule was

in the

of the
ox,

sheep

pig

were

more

the
to
ox,

the pig

in the

sheep

papilla

The d i a m e t e r
(horse),

91 p;

the

cat,

6Op;

(21p),

the

ox (33p),

in the h o r s e

(37p), and cat

(14p);

dog (9p),

sheep,

goat,

and s m a l l e r

and cat

and goat

and

(37p).
in the

in the h o r s e ,

The cell boundaries
goat,

and dog;

and

and cat.
duct was

in all

other

lined by a faintly
transitional

animals

species
59p;

epithelial
(22p);

ox,

s traight

(5p) than that

greater

and dog.

of this

of the p api l l a r y duct

s heep,

of the

(66p) and ox (53p);

dog

epithelium which b e ca m e

duct on the

138p

goat (42p),

goat (12p),

The p a p i l l a r y

columnar

dog.

in the h o r s e

d i s t i n c t than the pig in the h o r s e ,

d i s t i n c t in the

of the

equal

the d i a m e t e r

The lumen of the tubule was

smaller

13.
tall

(39p),

(lOp),

(14p).

and cat;

less

sheep

cell heig ht was

ox (12p),

greater

(50p),

goat,

m e a s ur in g
57p.

height was

and

smaller

As

simple

toward the

opening

investigation except

ranged f r om

54p (dog)

as follows:

pig,

96p;

co m p a r e d with that of

greater
in the

stained

in the h o r s e

sheep

(I8p),

dog

(55p),
(13p),

108

and c a t

(14p).

In the

horse,

ep ithel ium

extended f o r

from

openings

their

duct of the
liu m
the

its

papi lla w e r e
Langham

lining

the p a p i l l a r y

transitional

specimens

of the

openings

p elv is .

sitional

openings

in this

epithelium

d om es tic

cells

in this

were

covering

animal s

No

on

in ve stiga­

epithelium

In the p r e s e n t

(1953) doubted the

in the

study

existence

cat kidney and thought
epithelium into the

ducts of the

cat kidney were

the papilla was found to be t r a n ­

in this

spaces

fibroblasts,

in ve s tiga tio n

absent.

columnar

ducts

investigation.

The i n t e r s t i t i a l

and blood v e s s e l s ;

of this

epithe­

15 00p into the p api l l a ry duct

of the p api lla ry

f i b e r s ; a few collagenous f i b e r s

animals

simple

Duran-Jorda
ducts

columnar

animals

of the bovine kidney.

extended about

The papillary

of the p api l l a r y

p r e s e n t in all

of the p ap i l l a ry

The

in all
15.

ducts

examined.

found to be p r e s e n t
14.

The openings

simple

dialysed through the p api l l a ry

Th es e

the tr ansitional

in the p api lla ry ducts

lined only by a

course.

epithelium

was

distance

et a l . (1942) found only

in the

that u r i n e

and sheep,

on the papilla than in the pig.

found to be

tion.

goat,

a greater

dog kidney was

throughout

ox,

study.

wer e found to contain

reticular

around the tubules, Bowman s capsule,
h is tioc yt es ,

and m a s t

cells

in all

except in the dog and cat where m a s t

reports

wer e found r egarding

the p r e s e n c e

of

m a s t cells in the kidney of any spe ci es including man and l a bo r ato r y animals

1 09

16.

The

found to be
especially

intertubular

present

in the

tical

arteries,

were

larger

than cell

son with the
in the

a b s e n t in the
philic

groups

these

cat.

cell

less

In the

regarding

the p r e s e n c e

near

the

wer e

ox the ce lls
This

of these

is

nuclei.

sheep,

goat,

wer e l a r g e r

groups

the c o r ­

smaller

groups

In c o m p a r i ­
numerous

and dog;

and m or e

thought to be the f i r s t

cell

region,

mostly polygonal with

stained

in the

were

arteries

groups w er e found to be m or e

numerous

than that of the pig.

capsule,

Some of the

The cells
and dar kl y

cells

in the c o r t i c a l

groups

elsewhere.

cytoplasm

ox and h o r s e ;

Becher's

spaces

The cell

encapsulated.

pig,

or

neighborhood of Bowman's

and a r t e r i o l e s .

eosinophilic

groups

in the i n t e r s t i t i a l

close

w e r e found to be
f aintly

cell

and

eosino­

report

in the kidney of domestic

a n i m al s .
17.
under
ular

The

study to be p r e s e n t in close

cells

and the

f o r m e d by the
No

species

cerni ng
animals.
their

cell unit of Goormaghtigh was found in all

ce l l s

af f e r e n t

differences

the p r e s e n c e
Ham

nature,

(1953)

function,

of the
and

of th es e

m a c u l a densa in the

observed.
cells

stated that
or

a s s o c i a t i o n with the j u x t a g l o m e r ­

efferent a r t e r i o l e s

wer e

animals

triangular

and the

No r e p o r t s

macula densa.

were found con­

in the kidney of l a r g e

t h e r e was

n omen c l a t u r e

as

space

domestic

yet no c e rta inty

in the human kidney.

about

110
From
general
is

the

the

results

plan of the
same

except

of this

structure
a few

inve stigation it is

concluded that the

of the kidney of all domestic

spe ci es

v ar ia tion s .

animals

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