COMPARISON OF THE SIZE OF CELLS AND SOMEHISTOLOGICAL FORMATIONS BETWEEN
WHALES AND MAN
HIROSHI HOSOKAW A* AND TOSHIKO SEKINO**
INTRODUCTION, MATERIAL AND METHODS

Cells and histological formations constructing the huge bulk of thewhale's body are not so large as one might suppose. They are of thesimilar size to those of other mammals including man. Probably thesize of cells is determined by some biological factors which are, althoughunknown yet, rather common in the animal kingdom. The present
writers, following the histological studies of whales, happened to be
attracted to this notable similarity of cell size among whales and man
and tried to examine this similarity precisely and statistically.
Several kinds of histological preparations of whales and man wereused for this purpose. Whales used comprise the right whale (Balaena
gracialis Bonnaterre)***, blue whale (Balaenoptera musculus L.), fin
whale (Balaenoptera physalus L.), sei whale (Balaenoptera borealis Lesson),
one kind of the beaked whale (Berardius bairdii Stejneger) and one kindof the dolphin (Lagenorhynchus obliquidens Gill), all of which were adult.
Each of the specimens taken from these materials was embedded incelloidin, sectioned fifteen µ in thickness and stained with hematoxylin
and eosin. Measurements of nerve cells and fibres were, however, madeon the preparations stained by the Pal-carmin method.
In order to explain the method of statis-tical calculations, table 1 isto be referred to, in which the results of observations and calculations
are shown. The first column gives the items of investigation, which
includes several kinds of cells and histological units. For the measurements such cells were chosen at random that appeared to be cut throughthe center approximately. At the same time nearly round cells as possible were selected and two diameters of each cell, being at right angles
to each other, were measured by the micrometer. Then the arithmetic
average of these two values was got to represent the size of this cell.
The same holds good for such structures as the lung alveoles, Langerhans islets, renal corpuscles, etc. As to the muscle and nerve fibres,* Department of Anatomy, Faculty of Medicine, University of Tokyo.
** Brain Research Institute, University of Tokyo.
*** With the special permission of the Japanese Government, this right whale wascaught in May, 1956, for the scientific investigations.
in length.
269

It was a female and 38 feet 4 inches

270

H. HOSOKAWA AND T. SEKINO

the thickness of each fibre was measured to show its size.
Following the second and third columns which tell the species of whales
and the axis of measurement respectively, the fourth column (N) gives
the sample size or the number of cells examined. Column 5 (x) shows
the sample mean or the arithmetic average of the measurements, whichN
is represented by I:,x;/N. The unit of these figures is always " mici-1
ron." While the sixth column (S) indicates the total variance givenNby I:,(xi -x)2, two following columns (s 2 and u 2 ) state the sample and standi=l
ard variances respectively. These two variances are given by S/N andS/(N-1).

Column 9 gives the confidence interval for ' m ' or the mean of themother population. The significance level a for the figures is 0.05. It
means that the mean of the mother population lies within this intervalwith the probability of 95%. It is computed by the form
Pr.{x+uVF/N~m~x-uVF/N} =1-a.

(a=0.05)'F' is to be got by checking the table of F-distribution (n1 =1, n 2 =N-1).
Column 10 shows the critical regions for the samples, indicating thatthe measurements of all the sample must be, with the probability of
95%, within this region. It is computed by the form
Pr.{x+uv(N+l)F/N~Xi~X-UV(N+l)F/N} =1-a.

(a=0.05)

The last column gives the result of the statistical comparisons. Supposing that two samples are to be compared, of which the sizes andstandard variances are M, N and u2 , v2 respectively, the first step of
the statistical test is to get 'F0 ' given by u 2 /v 2 (u 2 >v2 ) as well as the'F' value from the F-distribution table, the degrees of freedom (n 11 n 2 )being M-1 and N-1. If ' F 0 ' is larger than ' F' (F0 > F), it means thatthe comparison of these two samples is statistically impossible (X).
For the null-hypothesis that these two belong to one and the same
mother population can be abandoned with the danger of 5%. If ' F 0 'is less than 'F' (F0 <F), the second step of the test is to be taken.
After calculating the common standard variance w 2 which is given by(n1u 2 +n2v 2 )/(n1+n2 ) or (S,,+Sy)/(M+N-2), 'F0 ' should be got by thefollowing form.

x,S,,, S11 andii represent the total variances and sample means of
the two samples to be tested. Then the ' F' value is to be got by

I
I

l

({)

C!J

C!J

.!SOil

"

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0

"+-!

~

ro

"Cl

....

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Fat cells

U

0

....

.~
...,

ea -B .-u~~{ lengthwidth
averagelengthwidth
averageI

a. Right w.
b. Manb. Mana. Right w.

a. Sei whaleb. ManPurkiuje
fibresc. Mana. Sei whaleb. Manlengthwidth
averageb. Blue w.

( lengthwidth
averagelengthwidth
averagea. Right w.

(/)

0.

·~
~

"'C!J

"t:l

(3)

(2)

Heart
musclefibres~>....C!J

~
u

~

Item

(1)
(5)

50
50
50
50
50
50
50
50
50

74.7
57.7
31.6
85.6
69.0
59.7

13.3
10.0
11.7
14.1
11. 7
12.9
92.6
56.3

30
30
30
30
30
30
50
50

13773.04605.33848.8
355.5

37.1
26.2

717.7
609.9

195.5
144.0
99.1
181.3
157.3
100.0
182.2
199.1
138.7

S(µ2)

(6)

20
10

10.7
10.6

18.6
13.9
16.3
18.3
14.9
16.5
18.0
14.4
16.3

N

Sample
mean
x(µ)

({)

100
100

(/)

"'

s ·~

0. 1:l

C!J

(4)

275.3
91.2

2.5
1.9
1.1
2.9
2.3
2.0

192.4
35.5

7.2
6.1

3.9
2.9
1.9
3.6
3.1
2.0
3.6
4.0
2.8

8 2(µ2)

(7)

281.1
94.0

2.6
2.0
1.1
3.0
2.4
2.0

202.6
39.5

7.3
6.2

4.0
2.9
2.0
3.7
3.2
2.0
3.7
4.i
2.8

u2(µ2)(8)

97.3-87.9
59.1-53.5

13.9-12. 7
10.5-9.5
12.1-11.3
14.7-13.5
12.3-ll.l
13.4-12.4

30.6-21.8

43.8~30.4

11.2-10.2
11.1-10.l

19.2-18.0
14.4-13.4
16.7-15.9
18.8-17.8
15.4-14.4
16.9-16.l
18.5-17.5
14.9-13.9
16.8-15.8(a=O. 05) (µ)

Confidence
interval ofmother mean

(9)126.2-59.0
76.3-36.3

16.7-9.9
12.8-7.2
13.8-9.6
17.5-10.7
15.1-8.3
15.9-9.9

67.7-6.5
40.7-11.7

16.0-5.4
15.6-5.6

22.7-14.5
17.3-10.5
19.2-13.4
21.9-14.7
18.5-11.3
19.4-13.6
21.6-14.4
18.0-10.8
19.7-12.9(a=0.05) (11)

Critical region
for sample(10)

TABLE 1. SIZE OF CELLS AND CELL AGGREGATES OF WHALES AND MAN.

(11)

a-b

a-b

a-b

a-b

b-c

a-c

a-b

x

+

x

-

-

(a=0.05)

Statisticalcomparison

t'l
t""'

l'V

"'1
......

(fJ

t""

n

>:i:j

0

t'l

N

(fJ

t'l

::c:

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0

z

0

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n

0
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...
....

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Cl)
u

~
c

·D.
UJ

~

u

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0
0
...

~

c

Cl)

.µ

q:::

..0

>

rn

~

·oj

...

(/)

*'1l

::s

~

...

lengthwidth
averagee. ManI( a. Berardiusb. Lagenorhynx
I\ c. Man

lengthwidth
averagelengthwidth
average!
!
!

{ lengthwidth
averagelengthwidth
average

!

(/)

<t:

·~

d. Right w.

c. Blue w.

b. Fin w.

a. Sei w.

D.
UJ

Cl)

·o

0'"

\ c'. ManI

. r a'. Berardius
S 8 . b'. Lagenorhynx(/l.j..>~...; ....,
0
::s 0

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(/)
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Item

Cl)

(3)

(2)

(1)

rn

(CONTINUED)TABLE 1.

(4)

UJ

19.3
15.9
17.6
19.1
15.3
17.2
20.8
15.7
18.3
23.1
16.7
19.9
18.0
15.1
16.8
17.1
13.5
10.6
15.1
11.8
10.0

N

50
50
50
50
50
50
50
50
50
50
50
50
50
50
50
50
50
50
50
50
50

Sampel
mean

(5)

x(µ)

aro ·-"'

~ ~

Cl)

6.0
5.3
2.8
7.0
6.4
3.1
4.1
2.2
1.9
9.2
6.7
4.3
6.2
5.2
4.2
2.9
1.2
1.6
3.0
1.2
1.8

145.4
60.0
78.7
152.0
59.8
92.0

g2(µ2)

(7)

297.8
264.0
140.0
351.1
320.9
154.6
202.7
112.0
93.3
459.6
333.3
214.2
308.4
258.7
211.5

S(µZ)

(6)

2.95
1.2
1.6
3.1
1.2
2.0

6.0
5.4
2.9
7.2
6.5
3.2
4.1
2.3
1.9
9.4
6.8
4.4
6.3
5.3
4.3

uZ(µZ)(8)(9)

17.6-16.6
13.8-13.2
11.0-10.2
15.4-14.8
12.1-11.5
10.4-9.6

20.0-18.6
16.6-15.2
18.1-17.1
19.9-18.3
16.0-14.6
17.7-16.7
21.4-20.2
16.1-15.3
18.7-17.9
24.0-22.2
17.4-16.0
20.5-19.3
18.7-17.3
15.8-14.4
17.4-16.2(a=0.05) (µ)

Confidence
interval ofmother mean

(10)20.5-13.7
15. 7-11.3
13.5-7.7
17.2-13.0
13.9-9.7
12.9-7.1

24.3-14.3
20.9-10.9
21.2-14.0
24.8-13.4
20.3-10.3
20.8-13.6
25.1-16.5
18.6-12.8
21.2-15.4
29.5-16.7
21.7-11.7
24.2-15.6
23.0-13.0
20.1-10.1
21.1-12.5(a=0.05) (µ)

Critical region
for sample

(11)

{n

a
b-c
a'-h'}
b'-c'
a'-..c'a-a'}b-b'c-...c'

+

+

x

+

x

+

x

le
d-e

x

-

c-J d

+

x

e

c

+

+

d
e

+

c

b- d

a-

b

(a=0.05)

Statisticalcomparison

N>
N>

0

z

t".I
:;:::
......

(f);3

zi:::;

;t>
;t>

~

;t>

~

0(f)
0

::i::

t:i:

-'1

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'1l

~

~

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...,,

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Cl)

~

t)

~

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i:i.

......

z

~~

8 ~

.~

f
*oo ..;.:.Sa> l::::: l3 u

d . ·- (cj ~~ lZ ..c !)() l

Item

(1)
00

(f)

i:i.

II

a.Bluew.
b. Right w.
c. Mand. Right w.***b. Mana. Right w.

-

;:;
>
<G

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~

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lengthwidth
averagelengthwidth
average

<G

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(3)

(2)
gi

(CONTINUED)

a. Sei w.
b. Manc. Mousea. Sei w.
b. Manc. Mousea. Sei w.
,.-..,~b. Manc. Mousea. Sei w.
SJ b. Man~c. Mouse

TABLE I.

50
50
50
50

50
50
50
50
50
50

50
50
50
50
50
50
50
50
50

50
50
50

35.9
35.3
35.0
32.0

336.7
217.1
276.6
195.0
144.3
170. 7

34.4
31.l
15.4
80.0
40.5
17.7
68.9
36.7
20.0

60.7
29.4
25.3

z(µ)

N

U'J

§- 'CiiN(5)

Sample
mean

'1l

(4)
~

914.0
841.6
679.0
855.5

200857131060
97344
73853
35913
35785

705.9
674.6
150.1
8051.62605.4383.0
2659.51485.2503.02377.0
466.6
460.2

S(µ2)

(6)

18.3
16.8
13.5
17.1

4017.12621.2
1946.91473.1718.3
715. 7

13.5
3.0
161.0
52.1
7.7
53.2
29.7
10.l

14.1

47.5
9.4
9.2

s2(µ2)

(7)

18.7
17.2
13.8
17.5

4099.12674.7
1986.61507.2
732.9
730.3

14.4
13.8
3.1
164.3
53.2
7.8
54.3
30.3
10.3

48.5
9.5
9.4

u2(µ2)(8)(9)

37.1-34.7
36.5-34.l
36.0-34.0
33.2-30.8

354.6-318.8
231.8-202.4
289.3-263.9
206.0-184.0
152.0-136.6
178.4-163.0

36.2-32.6
32.2-30.1
15.9-14.9
83.6-86.4
42.6-38.4
18.5-16.9
71.0-66.8
38.3-35.1
20.9-20.0

62.7-58.7
30.3-38.5
25.6-25.0

Confidence• t
1 0f
mtherva
mo er mean
(a=0.05) (µ)(10)

44.5-27.3
43.7-27.7
42.2-27.9
40.6-23.4

464.5-208.9
322.1-112.l
367.3-185.9
273.5-116.5
199.3-89.3
225. 7-115. 7

47.3-21.6
38.6-23.6
19.0-11.9
105.7-54.3
55.3-25.7
23.4-12.0
83.8-54.0
478.-25.6
26.7-13.3

75.0-46.4
35.7-23.1
27.3-23.3

Critical region
for sample(a=0.05) (µ)

(11)

_..

b

a}
b -d
c

~:::::~

a-b}

a-b

x

x

x

+

+

x

f x

l
f
b1
\
b -c

a-

b-c

a

a-b
b-c
b
ab-c

Statisticalcomparison(a=0.05)

tI1

~
c.>

CJ>

!:lr<

(")

'Tj

0

trj

N

UJ

>-l
:::c:

'T1

O
OZ

(il

::>::l

>

~

"i::I

(")

"'

<G

C><

~

"'

lengthwidth
average

(4)

if)

80698
50869
55820
43687
16731
26026
23660
13013
12210
39462
28730
31561

163.8
132.6
148.0
165.1
135.2
151.0

50
50
50
50
50
50

S(µ.2)

(6)

175.5
111.8
144.0
141.7
109.2
125.0

(5)
Sample
mean
x(µ)

50
50
50
50
50
50

N

"' "'

8 ·-

A ~

Cl)

473.2
260.3
244.2
789.2
574.6
631.2

482.9
265.6
249.2
805.3
586.3
644.0

170.0-157.5
137 .2-128.0
152.5-143.5
173.2-157.0
142.1-128.3
158.2-143.8

187.0-164.0
120.7-102.9
153.6-134.4
150.2-133.2
114.6-103.8
131.5-118.5

1614.0 1646.9
1017.4 1038.1
1116.0 1139.0
873.7 891.6
334.6 341.4
520.5 531·1

8 (µ2)u2(µ2)(8)(9)
Confidence
interval ofmother mean
(a=0.05) (µ)(7)* Stained by the Pal-carmin method.
** (1) at the level of rostral collicu!Us, (2) at the level of caudal colliculus.
*** Embedded in paraffin.

su

Cl)

~

b. Man"'l':l"' ::s"'C><

u

-

~

lengthwidth
averagea. Right w.

..-l

"'

lengthwidth
averagelengthwidth
averageb. Mana. Right w.

if)

Cl)

l':l ·-

Cl)

Ui

Cl)

b!I

~

"'.... "'

l':l

·~

·u"'

Item

!
!
!
!

(3)

Cl)

(2)

(CONCLUDED)(1)

TABLE 1.

208.1-119.5
165.4-99.8
180.1-115.9
222.9-107.3
184.5-85.9
202.4-99.6

257.6-93.4
175.3-48.3
212.5-75.5
202.4-81.0
147.8-70.6
171.4-78.6(10)
Critical region
for sample(a=0.05) (µ)

a-b

a-b

x

x

(11)
Statisticalcomparison(a=0.05)

Nl

0

z

~

~

(fJ

...,i

zt:I

;i..

;i..

::El

;i..

~

::i::
0
0
if)

::i::

""'

-:i

275

COMPARISON OF THE SIZE OF CELLS

checking the F-distribution table, n 1 and n 2 being 1 and M+N-2 respectively. If 'F0 ' is larger than 'F' (F0 >F), it means that there isstatistically a rational difference between these two samples ( + ), while
in case F 0 <F, it is shown that there is no rational difference at all
between these two (-).

RESULTSAll the results of observations and calculations are shown in table 1.
Each of the items will be explained briefly. Accompanying figures andgraphs are to help the explanation.

30

25
20

10

5

~
+"'

..r::

Oil

~
<!)

Pa ..E

"°

i::
«!

~

'-------~

~

·v
Ul

~
+"'

i::
«!

~

..r::

i::

~

~

~

~

i::
::J § «!
<!)

rn

rn

.8 0'..r::::Ju

"O
....
«!

i:: i:::>, i::
Oll,.r:: «!
<!)

rn

::J

rn

::J

'..r::;a.... OU
i:: i::i::....«! «!~E
Cl).... «!

Hepatic Heart m. Adreno·
cells
fibres corticalcells
Fig. 1.

Parietal cells

Ventr. r.

~
+"'

i::

..r::

i::

~ Ul ~
~~

Pa

~

....
Oil
«!
·v ri:i:: P'.i Pa ~ &:lj'-'~Pa ~ Ul"° . . ,'----_,--/"------ ,.------" ~_,.---------'" ' - - - - - - - - - - - - - . _ , - - - /~Dors. r.

~Spinal root fibres

·v

«!

Oil

«!

~,--/

Purkinje Fat cells
fibresAverage means of the cell sizes.

Hepatic cells (Figs. 1, 2.)
Not only the microscopical appearances but also the size of the hepatic
cells is quite similar to each other among the right whale, blue whale
and man. The sample means of the longitudinal and transversal
diameters are 18.6µ and 13.9µ for the right whale, 18.3µ and 14.9µ forthe blue whale, and 18.0µ and 14.4 µ for man. The averages of these

276

H. HOSOKAWA AND T. SEKINO

25

15
.£l

'E

....

....0

~ JO

....OJ

Q)

..c

s
z::l

"S::l

z

/O-

II-

12-

13-

14-

15-

4-

Size of cells (3.0=4µ)Fig. 2.

5-

6-

7-

8-

9-

JO-

//-

12- 13-Size of muscle fibres (3 .0=4µ)Size distribution ofhepatic cells.

Fig. 3.

Right whale
Blue whale
Human

Size distribution of heart
muscle fibres.
Sei whale·--·Human

15

.£l

.£l

....0

....0

al
u

al
u

....
Q)
..c

....
Q)
..c
8

s
z::l

z::l

B-

9-

JO-

Size of cells (3.0=4µ)Size of cells (1. 0= 13µ)Fig. 4. Size distribution ofadrenal cortical cells.

Fig. 5. Size distribution of fat
cells (in the gastric submucosa).

Right whale·--· Hum<1nRight whale·--· Human

COMPARISON OF THE SIZE OF CELLS

277

two diameters are also of high similarity, being 16.3µ, 16.5µ and 16.3µrespectively. Furthermore, it is illustrated in figure 2 that these cellsof whales and man show very similar mode of distribution of the size.
The statistical comparison revealed that there were no rational differencesamong these three populations of cells.
Heart muscle fibres (Figs. 1, 3)
As shown in figure 3, muscle fibres constructing the heart of the sei
whale and man are almost the same size in their thickness. The sample means are. 10. 7µ and 10.6µ respectively and the distribution curvefor each takes a similar shape and position. The statistical calculations
showed that no rational differences in thickness were recognizable between these two groups of muscle fibres.
Purkinje fibres (Fig. 1)
Contrary to the high similarity of the thickness of ordinary heart
muscle fibres between the sei whale and man, Purkinje fibres present
remarkable difference in size between
these two species. The former, of
which the average thickness is 37.1µ,I5exceeds the latter by more than 10µ.
That the whale's heart is provided
with very thick Purkinje fibres is in..'!lteresting, when it is reminded of that ~ 10
the hearts of ungulates have also 'B....
remarkably thick Purkinje fibres.
,.0
Cortical cells of the adrenal gland ::l8z 5
(Figs. 1, 4)
Adreno-cortical cells in the deep
layer abutting on the medulla were
measured.
Although the sample
10- 11- 12- 13- 14- 1s- 16- 11- 18- 19means of these cells are not so difSize of cells (3. 0=4µ)ferent from each other between theFig. 6. Size distribution of parietal
right whale and man, their mode ofcells of the gastric mucosa.
distribution is noticably different.
· - · Right whale(Fig. 4) Necessarily a rational dif·- - -· Blue whale·-----· Fin whale
ference was encountered by the· ------ · Sei whale
statistical comparison.
·-·HumanFat cells (Figs. 1, 5)
Fat cells in the gastric submucosa of the right whale and man showeda remarkable discrepancy in their sizes, the former being nearly twiceas large as the latter. Their modes of distribution are also discrepant
to such an extent that the statistical comparison turned out to be impossible.
(j)H. HOSOKAWA AND T. SEKINO

278

Parietal cells of the gastric gland (Figs. 1, 6)
Parietal cells in the gastric mucosa were compared among the sei,fin, blue, right whale and man. Generally speaking these cells showeda considerable uniformity in size, the sample means ranging from 16.8µin man to 19.9µ in the right whale. Figure 6 illustrates diagrammatically the mode of distribution for each species. According to thestatistical calculations, however, the comparison of these whales andman gives rather un-uniform results. For in some cases there is norational difference (f. i. between fin and sei whale or man), while insome other cases there is rational differences (.f. i. between sei and

20

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Size of nerve fibres (3. 0=10µ)Fig. 7.

Size distribution of spinal root fibres (large medullated fibres).

·-· ventral root l Human·------·

dorsal root·-· ventral root·------· dorsal r.

·=--· ventral r.
·-----· dorsal r.

frhynchus} Lageno-l Berardiusfblue, right whale or man, between right and fin whale or man), andstill in other cases the comparison is impossible (f. i. between blue andfin, right whale or man).

COMPARISON OF THE SIZE OF CELLS

279

Spinal root fibres (Figs. 1, 7)
Thick-medullated fibres of the cervical nerves were chosen at random
and measured in their thickness for the Berardius, Lagenorhynchus andman. The sample mean of the ventral root fibres as well as of thedorsal ones is largest in the Berardius and smallest in man, that of theLagenorhynchus taking the mediate value in both groups of fibres. If
the ventral root fibres are compared with the dorsal fibres, the former
always exceed the latter in thickness. Hence the distribution curvefor the ventral fibres takes a position left to that for the dorsal fibres
for each of the species examined. (Fig. 7).
The results of the statistical comparisons among these six groups ofnerve fibres are rather irregular. In some cases it was revealed thatthe comparison was impossible (f. i. between the ventral root fibres ofBerardius and those of Lagenorhynchus or man, between the dorsal

.,.

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29- 31~Size of cells (2.0=5.1 µ)
Fig 8.

Size distribution of nerve cells of the hypoglossal nucleus.
• + + + · Sei whale·-·Human.-·-····Mouseroot fibres of Lagenorhynchus and those of Berardius or man). In othercases, however, the comparison turned out to be possible, althoughrational differences were recognized in most cases.

H. HOSOKAWA AND T. SEKINO

280

Nerve cells (Figs. 8-13)
Nerve cells in the hypoglossal and trigeminal mesencephalic nuclei aswell as the Purkinje cells in the cerebellum were measured and examined in the sei whale, man and mouse. The average of the longitudinal
and transversal diameters of each cell was used for the statistical calculations. As shown in figures 8, 9 and 10, in every item of the invesitigation it was made clear that the brain of the sei whale was furnishedwith the largest cells and the mouse's were smallest, while the humanmaterials were of the intermediate size. Statistical relations in size
among these groups of nerve cells are not so intimate. So far as thesize is concerned, most of them seem to belong to special kind of
population.

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Size distribution of the Purkinje cells.
· +++· Sei whale·-·Human-- -- · Mouse

When those three kinds of cells are compared to each other in eachof the species examined, the cells of the hypoglossal nucleus are largest
and the Purkinje cells are smallest for both the sei whale and mouse.

281

COMPARISON OF THE SIZE OF CELLS

25

20
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73~

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Size of cells (2.0=5.lµ)Size distribution of nerve cells of the trigeminal mesencephalic nucleus.
·+++· Sei whale
Human
Mouse

30
25
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19- 21- 23-

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Size of cells (2.0=5. lµ)Size distribution of nerve cells of the sei whale.
Nucl. nervi hypoglossi
Purkinje cells
Nucl. mesencephalicus V

H. HOSOKAWA AND T. SEKINO

282

30

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Size of cells (2.0=5. 1µ)Fig. 12.

Size of cells (2.0=5. lµ)Size distribution of nerve
cells of man.

Fig. 13. Size distribujon of
nerve cells of the mouse.

Nucl. nervi hypoglossi
Purkinje cells .
Nucl. mesencephalicus V

Nucl. nervi hypoglossi
Purkinje cells
Nucl. mesencephalcus V'· ',,,
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Size of acini (3. 0=10µ)Size of alveoles (1. 0=13µ)Fig. 15. Size distribution of the pancreas-acini.
Right whale
Blue whale
Human
Right whale (embedded in paraffin)

Fig. 14. Size distribution of alveoles of the lung.
Right whale
HumanCOMPARISON OF THE SIZE OF CELLS

283

In the case of man, however, the cells of the hypoglossal nucleus are

smallest, although the order in size of other two kinds of cells is thesame as in the sei whale and mouse. (Figs. 11, 12 and 13)
Alveoli of the lung (Fig. 14)
The pulmonary alveoli of the right whale are by far larger thanthose of man, the sample mean of the inner diameters being 276.6µ and170. 7µ respectively. The distribution mode is also quite different fromeach other and the comparison is statistically impossible.
Pancreatic acini (Fig. 15)
Outer diameters of the pancreatic acini of the blue, right whale andman are fairly alike, the sample means being approximately 35µ ineach case. Their distribution modes are, as shown in figure 15, alsosimilar to each other so extremely that they can be regarded statistically,so far as the size is concerned, as belonging to one and the same motherpopulation.
In the preparations of the right whale pancreas embedded in paraffin
the acini were smaller than those in the celloidin preparations. Thisdecrease in size is apparently due to the shrinkage. As the sample
mean for the paraffin preparations is 32µ compared to 35.3µ in the celloidin preparations, the decrease is calculated to be 9.3%. 1) Because ofthis decrease in size the distribution curve for the paraffin preparations
is shifted to the left considerably. The shape of the curve itself is
however almost the same as that for the celloidin preparations, thus
resulting merely in the rational difference statistically.
Langerhans islets
The Langerhans islets of the right whale pancreas are on the average
a little larger than those in the human pancreas. The sample meansof the measurements are 144µ for the former and 125µ for the latter.
Their distribution shows an irregular curve in both cases and the statisticalcomparison turned out to be impossible.
Renal corpuscles
Malpighian corpuscles of the right whale kidney are exceeded slightly
in size by the human equivalents. Furthermore the distribution for theright whale represents a steeper curve than that for the human material.
The curve for the latter is provided with a broader foot contour.
Relation in size between the nucleus and cell body
By the measurement of the adreno-cortical cells and the parietal cellsof the gastric glands their nuclei were measured too. The averagel)According to FUJITA (1947) the radii of the renal corpuscles of the rabbit were in the
average 32. 9µ, in the paraffin preparations, 36. 2µ in the celloidin preparations and 42. 3µin the frozen sections. Thus the decrease in size of the paraffin preparations comparedto the celloidin ones was 9. 1 %H. HOSOKAWA AND T. SEKINO

284

sizes given by the arithmetic mean of the length and width of nuclei
as well as of cell bodies are in table 2. The ratio between these twovalues seems to be approximately the same for each kind of cells of

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Size of cell-bodies (3. 0=4µ)Comparison of the size of cell-bodies and nuclei of the parietal cellsof the gastric gland.

whales and man. In the adreno-cortical cells the proportion of thenucleus to the cell body is nearly 50% in both the right whale and man.
The same ratio is shown to be considerably smaller and less than 40%
in the parietal cells for man and every kind of whales examined.

COMP ARJSON OF THE SIZE OF CELLS

285

The scatter diagram~ in figures 16 and 17 are to show the correlationin size between the nuclei and the cell bodies. Judging from these
diagrams there seems to be no simple relationship between these two.
The cell bodies are by far more variant in size than the nuclei within them.
TABLE 2.

RATIO IN SIZE BETWEEN THE NUCLEUS AND CELL BODY.

ItemSpecies

Cell body (11)

Nucleus (µ)

RatioAdrenocorticalcells
Parietalcells ofthegastric
gland

Right w.
Man

11. 7
12.9

5.3
6.3

2.2:1=1:0.45
2.1: 1=1:0.49

Sei w.
Fin w.
B:ue w.
Right w.
Man

17.6
17.2
18.3
19.9
16.8

5.4
6.0
6.8
6.6
5.8

3.2:1=1:0.31
2.9: 1=1 :0.35
2.7:1=1:0.37
3.0:l=l:0.33
2.9:1=1:0.35

HISTORICAL REVIEW

Cell size in general*
Since Schleiden and Schwann enunciated the cell-theory in 1838-1839,
the cell has attained and kept its position as the most fundamental andessential subject in the fields of biology. More than half a century has
elasped, however, before the problem of the size of this tiny mass ofjelly-like substance called protoplasm attracted the notice of scholars.
For it was Julius Sachs (1893) who for the first time paid special
attention to the fact that the cells forming big plants are not necessarily
larger than those of small plants. Amelung (1893), Strasburger (1893)
and Rahl (1899) followed Sachs and Driesch (1900) mentioned of '' thelaw of fixed largeness of the organ cells." It would be paraphrased
that not the cell size but the number of cells of organisms varies inproportion to the body size. Studying on the Crepidula (gastropod) andrabbit respectively, Conklin (1912) and Painter (1928) reached to thesame conclusion that the equivalent cells of animal bodies are, regardless of the individual variations in body size, almost the same.
Meanwhile, Hardesty (1902) stated, based upon the measurements ofCavazzani (1891) and Buhler (1898), that the variations in the size-of
cell-bodies of the spinal ganglia are not directly proportional to thevariations in the size of the body of the animal, though in general thelarger animal possesses larger cells. According to his own measurements of nerve cells in the spinal cord for several mammals, however,
the average mean diameter of the cell-bodies of the columna anterior in* Among others. surveys of the pertinent literature by Chambers (1908), Wilson (1925),
Wassermann (1928), Jacobi (1935), and Bucher (1954) are extensive and useful for reference.

286

H. HOSOKAWA AND T. SEKINO

the intumescentia cervicalis turned out to decrease gradually througha series of mammals of diminishing body weight. His observations
showed further that the volume of the cell-body varies more nearly inproportion to the body size of animals and the volume of the entire
neuron bears a still more constant ratio to the bulk of the animal
body.
Comparative examinations of the cell size of several kinds of mammals
led Levi (1905) to a noteworthy conclusion as follows. According tohim, there are two categories of cells, one of which comprises such cells
that show a notable uniformity in size all through the animals, while
the cells of the second group vary in proportion to the body size.
Epithelial and glandular cells belong to the first group and the large
ganglion cells represent the typical example of the second group. Levi
tried to explain the difference in relation to the histogenesis of eachcells. For the cells of the first group maintain the potentiality of cell
division during the whole life of animals, while the second ones lose
the possibility early in the development. Observations of Obersteiner(1913) on Purkinje cells of the cerebellum of the sei whale, elephantand mouse gave the same relationship between the cell size and bodysize. Ganglion cells of the invertebrates were, however, shown by
Erhard (1912) to vary capriciously and independently of the body size.
Even for mammals he denied the intimate correlation between the sizeof cells and animal bodies.
Hatai (1902) measured the nerve cells in the spinal ganglia of thewhite rat and noticed the increase of cell size following the growth ofanimal. According to Pfuhl (1932), same kind of growth of the hapatic
cells was reported by several authors for man and animals ; Harting(1845) and Toldt-Zuckerkandl (1876) for man, Kretschmar (1914) for thepig, Plenk (1911) for the rat, Heiberg (1907) for the mouse and Illing(1905) for domestic animals. Akiyama (1928) made a similar observation
for the adrenocortical cells of the white rat. According to Berezowski(1910) the epithelial cells of the intestinal villi were observed to growand lengthen too. Detailed investigation of the growth of cell size wasmade later by Rohrbacher (1927), who stated that each kind of cells
has its specific size particular to the growth stages of animals.
Another factor influencing the cell size was pointed out by Chambers(1908), who noticed that the frog developed from a small ovum had
smaller cells than the frog from a larger ovum. So it is possible, as
Wassermann (1929) said, "<lass bis zu einem gewissen Grad die difinitive
Zellgrosse auch <lurch die Grosse der Ursprungszelle, des Eies, bestimmt
sein kann."
The last and important matter to be considered in relation to the cell

COMPARISON OF THE SIZE OF CELLS

287

size is the quantitative correlation among the cell, the nucleus and thechromosomes. R. Hertwig (1903) was the first to notice the constancyof the ratio in volume between the cytoplasm and the nucleus and heestablished the theory of the nucleo-cytoplasmic relation or the karyoplasmic ratio (Lehre der Kern-Plasma-Relation). Erdmann (1908, 1909),
Koehler (1912) etc. followed Hertwig and developed his theory. Erhard(1912) too stated that "die Grosse der Kerne (der Ganglienzellen) richtet
sich nach der Grosse der Zellen, nur haben Zellen mit reichlicher Nisslsubstanz stets kleine, solche mit wenig oder gar keinem Tigroid stets
grosse Kerne."*Deviations of the nucleo-plasmic relation were studied by several
authors such as Erdmann (1911), Lanz (1926), Stieve (1926), Tretjakoff(1928), etc. Excepting pathological cases, most of these changes of thenucleo-plasmic relation were apparently connected to the fluctuations of
the hormonal condition. The functional hypertrophy is another element
which causes the shifting of this relation in favour of the cytoplasm.
According to Bucher (1948) the nucleo-plasmic relation changes as theaging progresses, resulting smaller nuclei in the cells of higher ages.
Studying upon the chromosomes of the sea-urchin larvae, Boveri (1905)
found the parallelism between the cell size and the number of chromosomes or the amount of chromatin in the nucleus. Thus the cell body,nucleus and chromosomes are shown to be in quantitative correlation to
one another, although no one knows whether one of these three takes
the initiative in determining the volume or size of the others.
Such a harmonious balance prevailing in the. intracellular structures
fascinated Heidenhain (1907-) and he called it "syntonischer Zustand"or " Syntonie " or " Kanon der Teile und des Systems." The idea of" Syntonie " has developed hand in hand with that of the " dividing
bodies " (" Teilkorper ") such as protomeres, histomeres and histosystems. For both of these ideas represent important constituents of his
famous " synthetische Morphologie " or " Synthesiologie." By meansof these unique ideas Heidenhain tried to grasp the biological principle
prevailing in the intra-, extra-, as well as the "supra "-cellular structures of the living body.
At least one part of Heidenhain's hypothetical theories was proved byfact, when Jacobj (1925) found the rhythmical variations of nuclear
volumes in the hepatic cells of rats and mice. The so-called .J:acobj's
law or " das rhythmisches Verdoppelungswachstum " of the nuclear
volume was tested and accepted by Voss (1928), Clara (1928, 1930) and* G, HERTWIG (1931) stressed upon the necessity of careful!ness in considering thenucleo-plasmic relations. For he showed clearly that the cytoplasm reacts to fixatives
with irregular shrinkages of higher grade than the nucleus.

lt. lIOSOKAWA AND T. SEKINO

288

many other scholars. Jacoby (1935) himself investigated later on manykinds of human cells and classified them into a series of c1asses, inwhich the nuclear volumes increase by doubling. Standard nuclear sizefor each class is as follows.
Class

Diameter (µ,)

Volume (cub µ,)

K 1/8
K 1/4
K 1/2
K 1
K2

3.25
4.1
5.2
6.5
8.2

18
36
72
144
288

K 16

16.4

2304

In recent years new attention has been paid to Jacobj's doubling
phenomenon of the nuclear volume. For some of the histochemical
studies showed a parallelism between the nuclear volume and the DNA
amount. (cf. Swift, 1953; Alfert, 1955 etc). New methods of precise
caryometry are also being devised. (f. i. Bucher, 1954.) At the sametime the nucleo-cytoplasmic relationship of R. Hertwig is increasing its
importance in the fields of modern biology (cf. Frankenhauser, 1952;
Hammerling, 1953 etc.), and the ratio between nucleus and nucleolus too
is becoming an interesting subject. (cf. Junqueira and Hirsch, 1956.)*
So far is the review of the literature on the problem of cell size ingeneral. Concerning the items of the present writers' investigationsome supplemental survey of the history will be added.
Hepatic cells
Several works on the quantitative study of the hepatic cells and nuclei
have been mentioned already. Data given for the human hepatic cells
are as follows.
v. Ebner (1899)
Bucher (1948)
Kopsch (1955)
Bargmann (1956)

18-26 µ, in diameter
20-25
18-35
13-30

Average means of the cell size for several animals were tabulated by
Pfuhl (1932) in the following way. Apparently the hepatic cells show
a fairly uniform size among animals.

* It .would be noteworthy that Hsu (1954) studied in vitro on the chromosomes of humanneoplasms and showed in graphs that the chromosome number distributes in the mode of
doubling increase just like the nuclear volumes in Jacobj's graphs.

289

COMPARISON OF THE SIZE OF CELLSSIZE OF HEPATIC CELLS (PFUHL, 1932)
Animal (adult)
Rat
Rabbit
Cat
Dog
Horse
Pig
Ox
Goat
Sheep

Author
Plenk
Schlater
I11ingAuerbach
I11ingNucleus (µ,)
8.0
8.3

Cell (µ,)
23.0
25.7
21.1
20.0
26.3
26.5
21.4
23.3
23.6
29.0
21.5
20.7

7-8

,,
,,

Kretschmar
I1lingBaum
I11ing,,Heart muscle fibres and Purkinje fibres
For the detailed survey of the literature Benninghoff (1930) and
Haggqvist (1931, 1956) are to be ref erred to. Measurements of the widthof the human cardiac fibres were reported as follows.
Letulle (1897)
v. Ebner (1899)
Marceau (1904)

5-25µ,
9-22
5-40 (average 20)

The size of the human heart muscle fibres in relation to the aging wasstudied very extensively by Schiefferdecker (1916). According to him,
the muscle fibres as well as the nuclei increase in size with aging.
For animals Schiebler's data (1953) will be cited.
Animal
Ox
Calf
Pig
Dog
Cat

Ordinary fibres of
the heart muscle
10 µ,
10
10
10
10

Purkinje fibres
35-45 µ,
18-32
20-26
14-20
11

Notable uniformity seems to be prevailing in the size of heart muscle
fibres, while the size of Purkinje fibres varies from one animal to another.
According to Hirai (1943), however, the heart of such animals as withhigher activity is made of compact bundles of thinner muscle fibres.
SIZE OF HEART MUSCLE FIBRES (HIRAI, 1943)
Animals with higher activity
Wild dog
Dog
Japanese
spanielHare
Water rat

157±5.3 µ,182±5.8
181±7.1
178±5.8{ 185±6.3
189±5.4

Animals with lower activity
211±9.4 µ,Rabbit { 225±10.5
Mouse { 224±9.0
234±9.5

H. HOSOKAWA AND T. SEKINO

290

Davies and Francis (1952) studied comparatively the hearts of mammals
of various kinds and said that the ordinary myocardical fibres in all theanimals examined showed a remarkable uniformity in diameter. On theother hand the fibres of the conducting system showed considerable
difference in size among animals, the Purkinje fibres always exceeding
the fibres of the atrio-ventricular bundle. Furthermore they examined
the relationship between the size of the Purkinje fibres and the duration of QRS of the electrocardiogram for each animal, and got thesuggestion that in hearts of thicker muscle fibres of the conducting
system the cardiac impulse spreads through the ventricles at a higher
rate than in those of thinner fibres.
DIAMETERS OF THE HEART MUSCLE FIBRES. (DAVIES AND FRANCIS, 1952)
Atrio-ventr.
bundle(µ.)

Sub~ndocardial

Animal
Horse
Cow
Human
Wallaby
Sheep
Dog
Cat
RabbitRat
Swan
Pigeon

35
30
11
36
30
12
9
10
9
35
11

88
40
18
40
40
18
15
14
13
44
12

Purkinje
fibres(µ.)

Ventricularmyocardium
(µ.)

QRS
(1/lOOsec.)

12

7
9
8
3.5
3
4
4
3
2
3
2.7

II
II
II
II
II
II
II
II

9
8

Pulmonary alveoles
According to the sur.vey of literature by Bargmann (1936), the numberof alveoles of the human lungs are 300-500 millions and the total areaof their internal surfaces amounts to 50-100 square meter. The average
diameters of the human alveoles reported by several authors are as
follows. (Data marked with U) are cited from Bargmann, 1936.)
Rossignol (1847)'
Frey (1859)'
Kolliker (1880)'
v. Ebner (1899)
Schulze (1906)
Ogawa (1920) 1
Wilson (1922)'
Marcus (1928)
Claus (1935)'
Kopsch (1955)
Braus (1956)
Bargmann (1956)

adult
newborn
infant200-250µ.
50-166.7
60-90
160-220-370
200
100-19075x90x125150
300-600
45-60
100-150150-350230
150-600

Z91

COMPARISON OF THE SIZE OF CELLS

Concerning the lungs of various mammals, the extensive data given
by Schulze (1906) and Marcus (1928) will be cit€d here. It would be
added further that Fiebiger (1915) studied on the lung of a dolphin andstated the size of alveoles as 260x140µ.
MEASUREMENTS OF LUNG AL VEOLES. (SCHULZE, 1906)
AnimalDiameter ofalveoles (µ)

Cat
Sloth
Man
Dolphin100
400

Respiratory area(sq. meter

Number ofalveoles

400 millionas
6.Z5
150
437

zoo

140

zo
5
30
43

(Schulze regarded the difference of the respiratory area as indication of activity of animals.)
MEASUREMENTS OF LUNG AL VEOLES. (MARCUS, 19Z8)
AnimalVolume oflungs (ccm)

Respiratory Diameter ofmass (ccm) al veoles (µ)

Dolphin
50
Mouse
0.9
Rat
7
Cat
180
Man
1880
Bat
3
Galeopithecus 10
Young deer
420
Calf
3050
Horse
17500

40
7.2
5.6
144
1500
2.4
8
336
2440
14000

200
30
50
100
150
Z5
150
120
160
140

Numberofalveoles5 millions
Z66
45
144
444
160
Z.5
200
600
5000

Respirat. Resp. area (sq. cm.)area
per gr. of(sq. m.)body weight

1
0.1
0.6
7.Z
50
0.5
0.3
14.4
76.8
500

31
54
33
Z8
7
100
10
21
13
11

Renal corpuscles
Numerous papers have hitherto been published which treated directly
or indirectly the size of the renal corpuscles of man and animals. Data
given in those works will be arranged and tabulated as follows. (Values
put in the brackets concern the corpuscles situated deep in the renal
cortex. Data of authors with the mark C'> or C"> are cited from
Mollendorff-1930- or Vimtrup-1928- respectively.)Man:

217 '"

zoo

200-300
200-300
213 (~). 196 (Sf-)
Z37
200 : 300, 149-21Z
130: 220
210
19Z : 159
Z00-300
176-212

Bowmann (184Z)''
Schweigger Seidel (1865)
Sappey (1879)''
Toldt (1E88)'
Eckhardt (1888)''

Kiilz (1899)"
Glantenay and Gosset (1901)'
v. Ebner (1902)'
Moore (1903)
Peter (1909)
Prenant (1911)'
Moberg (19Z9) 1

292

H. HOSOKA WA AND T. SEKINO
159. 54 (paraffin)
169. 82 (celloidin)
193. 86 (frozen section)
218: 171200
130-220200-300

Pig

Mouse:

180-350
175
128 : 149 (167 : 219)
161 : 210 (212 : 270)
176 : 192 (209 : 240)l Fujita (1947)J

Abe (1953)
Greep (1954)
Kopsch (1955)
Bargmann (1956)
Ki:illiker (1863) 1
Schweigger-Seidel (1865)~

f

Roost (1912)'
Schweigger-Seidel (1865)
Peter (1909)v. Mi:illendorff (1927)'
Abe (1953)60
103: 86
136 : 110, 89 : 84
88: 68Schweigger-Seidel (1865)
Abe (1953)

Guinea gig: 128
84: 101
Bat:

75

Mole:

63

Peter (1909)

Schweigger-Seidel (1865)
II

II

II

II

122
102
124 : 124 (175 : 153)
96 : 80 (144 : 128)

Miller and Carlton (1895)''
Peter (1909)
Roost (1912) 1

Sheep:

210
173 : 153
144 : 128 (192 : 176)
158: 132

Schweigger-Seidel (1865)
Peter (1909)
Roost (1912) 1
Grundmann (1922)'

Weasel:69

Schweigger-Seidel (1865)

Goat:

200 : 176 (240 : 208)
150: 122

Roost (1912) 1
Grundmann (1922)'

Rabbit:116: 9134-40
76. 94 (paraffin)
84. 16 (celloidin)
101. 04 (frozen section)
114: 89

Peter (1909)
Boycott (1911)

Cat:

Ox:

209: 172
224 : 193 (188 : 170)
200 : 176 (270 : 240)
225: 153

1\[

Fujita (1947)
Abe (1953)
Peter (1909)
Inouye (1909)
Roost (1912)'
Abe (1953)

293

COMPARISON OF THE SIZE OF CELLS
Porpoise (Phocaena communis):Peter (1909)
Inouye (1909)

130 : 103
ff:"

Horse:

270 : 240 (272 : 240)

Roost (1912)'Dog:

256 : 240 (288 : 272)
162 : 127

Roost (1912)'
Abe (1953)

White rat: 127 (62, newborn)124
113: 91

Kittelson (1917)
Arataki (1926)
Abe (1953)

Supplemental data about the renal corpuscles will be added in thefollowing,
MEASUREMENTS OF THE RENAL CORPUSCLES. (PtlTTER, 1927 AND
MOLLENDORFF, 1930)

AnimalSurface ofeach glomerulus(sq. mm.)Mouse
Rabbit
Cat
Sheep
Man
Ox
Pig
Echidna

0.087
0.101
0.144
0.249
0.293
0.335
0.425
0.183

Total no. ofglomeruli inboth kidneys

Total surface
area of glomeruli(sq. cm.)

10 thousand
285
460
1010
1700
8050
1400
180

20.5
288
662
2520
4950
27000
5980
330

Relative glomeruli
surface per gr.
of body weight
1. 08 sq. cm.
0.144
0.221
0. 07180.07080.06000.089
0.165

According to " Biological Data " edited by Spector (1956) (Table 145),
the volume of glomeruli (cub. mm.) per one gram of kidney is as follows,
showing a considerable uniformity among animals and man; man: 29,
cat: 28, dog: 40, elephant: 42, ground hog: 75, guinea pig: 42, monkey: 50, mouse: 21, opossum: 49, ox: 47, rabbit: 46, albino rat: 40,
kangaroo rat: 30, swine: 37.
Langerhans' islets
Quantitative studies of the Langerhans' islets of man and animals
were reviewed and surveyed extensively by Bargmann (1939) in v.
Mollendorff's Handbuch der mikroskopischen Anatomie des Menschen,
Bd. 6, 2 Teil, S. 209-. Among many papers treating this subject, merely
the representative ones will be nominated here: Clark (1913) and Nakamura(1924) for man; Bensley (1911) for the guinea pig; Hess and Root (1938)
for the white rat ; Glaser (1926) for the mouse ; Clara (1924) for birds.
So far as the size of the Langerhans' islets is concerned, however,294

H. HOSOKA WA AND T. SEKINO

the data of Heiberg (1909) are most extensive and some of them will
be cited here.
AVERAGE DIAMETERS OF LANGERHANS' ISLETS. (HEIBERG, 1909)
AnimalDog
Cat
Pig
Monkey
Sheep
Goat
Ox
Horse
Pig
Ox

Long axis : Short axis
64: 43µ75: 54
86: 64
122: 75
93: 54
97: 54
155 : 75
100: 64
108 : 86 (cauda)
75 : 55 (caput)
75 : 52 (lobus dexter)
158 : 99 (cauda)
108 : 82 (caput)
126 : 82 (lobus dexter)

DISTRIBUTION OF SIZE OF L ISLETS. (HEIBERG, 1909)
AnimalMan
Mouse
Guinea pig
Dog
Cat
Pig
Horse
Sheep
Ox

below 75 µ 76-125 126-175 176-225 226-275
4
10
23
23
38
4
11
24
27
33
2
8
18
35
37
1
7
64
28
1
11
48
40
1
2
34
13
50
5
5
24
47
18
14
4
51
31
12
1
39
47

276-325 above 325
1
1
1

1

DISCUSSION

Comparative observations of the cell size in whales and man revealed
that some kinds of cells such as hepatic cells, adreno-cortical cells andparietal cells of the gastric gland as well as the heart muscle fibres
show a considerable uniformity in the size. On the other hand there
are other kind of cells which show a remarkable difference in the size
between whales and man. The Purkinje fibres of the heart, fat cells,
nerve cells and fibres represent examples belonging to the second catagory. A perusal of the pertinent literature suggests that the uniformityof size found in the cells of the first group holds good to a wider extent
in the kingdom of animals, so far as the special attention is paid toCOMP ARISON OF THE SIZE OF CELLS

295

mammals. As Levi (1905) pointed out, probably the epithelial andsecretory cells may be representatives of this group.
The literature tells at the same time that the variability of cell-size
encountered in the second group prevails also among the equivalent cellsof other mammals. It is an interesting and perhaps noteworthy factthat the nervous elements, especially larger ones, are the typical examples of this group.
Surveying the members belonging to the second group, it would benoticed that they are represented by relatively large cells. Also manyof those cells are furnished with some special intracellular structures or
inclusions such as Nissl bodies for the ganglion cells and the large dropof neutral fat for the fat cells. The extraordinary richness in sarcoplasm of the Purkinje fibres may be reminded of too. The ovum containing yolk granules as well as the cells of the sebaceous gland furnishedwith coarse droplets of fatty substance doubtlessly represents anotherexample. Also the pigment cells in which the pigment granules accumulate belong to the same category.
Nissl bodies of the nerve cell are, however, not to be regarded ascorresponding to fat drops, yolk substance, pigment granules and so on.
For the latter are merely cellular inclusions, while the Niss! bodies arecertainly a sort of important cellular constituent of the nerve cell. The
examination of the nucleo-plasmic relation will be probably useful, atleast to some extent, to discern between these two categories of intracellular structures. Cellular inclusions such as fat drops in the fat
cells do not affect the correlation between the size of nucleus and theamount of the proper cytoplasm. So the nucleus of such a cell is rather
small for the considerable bulk of its cell body. In the case of Nissl
bodies, on the contrary, the correlation holds good between the nucleus
and the total volume of the cytoplasm, resulting thus in such a cell as
furnished with nucleus of a fairly largeness.
By the way the present authors have no knowledge as to whether
the volume of the neurite and dendrites is to be taken into accountwhen the nucleo-plasmic relation of the neuron is considered. If it
should be, supposing that there are two nerve cells or perikarya ofa similar size which are though provided with axon or dendrites ofdifferent length and numbers, the cell with longer and more processes
must have a larger nucleus.
Returning to the earlier discussion, let us bring up a question. Is
there any way to explain the above mentioned difference between two
groups of cells in the animal tissues? This question is necessarily related to the problem of what is the definitive factors to determine thesize of cells.

296

H. HOSOKAWA AND T. SEKINOIn the literature, various factors have been stated by many authors
as influencing the size of cells~ For instance the activity of animals,
the rate of metabolism, the innate potentiality for further cell divisions,the size of animal body, the grade of growth and aging, functional
influences such as due to hormons and training, etc. have been examined
and their correlations to the cell size were proved to some extent. Probably each of those factors is ,working in its own way and the sum ofthem, in cooperation with still other innate factors, will determine thesize of cells, although its detailed mechanism is at present far beyond
our knowledge.
So far as the morphology is concerned, the quantitative correlation
between the cell body and nucleus, which was noticed by Hertwig (1903),
developed and elaborated by Heidenhain (1912-) and Jacoby (1925-) et al,seems to be of an important meaning. Especially Jacoby's phenomenonof the " rhythmical, doubling growth of the volume of nuclei " isastonishing and must be of a great importance.
Checking Jacobj's review table (1935), the present authors cannot
avoid such a suggestion that the discrimination of two cell groups
mentioned in the beginning of this chapter is related to the differenceof respective classes of those cells. That is to say, cells of lower classes or of smaller nuclear volumes (f. i. K 1 / 8 , K 1/ 11 K /~, Kl) belong to
the first group of the present description, where the cell size showsa considerable uniformity among animals. On the other hand, cells ofhigher classes or of larger nuclear volumes (f. i. KB, K16) are comprisedin the second group, where the cell size .varies from one animal toanother. The higher variability of cell size found in the latter groupis not difficult to understand, if it is taken into consideration that thestandard nuclear volumes here are of high values. It is also easy tocomprehend that the difference between those two groups of cells is not
an absolute but relative one. For there can be intermediate classes of
medium-sized cells (f. i. K2, K4). Probably nerve cells of small andintermediate sizes represent examples of this category.
On the size of some large histosystems.
The digestion, respiration and elimination of wastes are three principal
functions which are indispensable for keeping the vegetative life. In
the protozoa like amoeba all of these functions take place through the
body surface. In higher animals, however, each of these three functions
is carried out in a special part of the body ; the intestinal canal, lungs
and kidneys. Thus the intestinal villi, pulmonary alveoles and the renal
corpuscles may be regarded as specialized equivalents of the body surfaceof amoeba.
1

COMPARISON OF THE SIZE OF CELLS

297

By the way measurements of the lung alveoles and renal corpuscles
revealed that the size of these both shows considerable differences
between whales and man. Judging from survey of the pertinent literature, a similar discrepancy of size seems to be prevailing widely among
animals. Although the variations in diameter are quite irregular fromanimal to animal, the relative respiratory as well as glomerular surface
per unit of the body weight is in nearly inverse proportion to the bodysize of animals. Supposing a cell or amoeba shaped like a ball, therelative surface per unit volume is given by 4rrr 2 / 3/ 4rrr 3 =3/r. Thus it isalso inversely proportional to the size of the body. Probably this is
one of the reasons for the limitation of cell size. For, if it enlarges
beyond a certain size, its surface cannot take in food and oxygen fastenough to maintain its bulk.
In spite of the variability among animals the average diameter of
pulmonary alveoles as well as of renal corpuscles for every mammal
falls in a limited range of some 50-300µ. So we can still speak of
a fairly uniformity in size for these histological structures. The pancreatic acini represent an example of a higher uniformity. According
to Miziarsky (1900) the same relation was observed in the secretory
alveoles of the parotis too.
To explain the size of these structures is as difficult as to explain thenature of the cell size. In his superb thinking way of "Synthesiologie ",Heidenhain called those structures with a generic name '' Histosysteme ''(adenomeres, pneumomeres, etc.), and assumed that common biologicalfactors for cell size would be responsible for determining the siz'3 ofthese "supra "-cellular units or systems too. The present writers haveneither fact nor theory to develop the discussion further.
SUMMARY

1. Several kinds of cells and histological structures were measured
and compared statistically between whales and man. Historical reviewof the literature extended the comparison to other mammals.
2. Some kinds of cells such as hepatic cells, adreno-cortical cells,parietal cells of the gastric glands, heart muscle fibres, etc. were shownto have a fairly similarity in size among animals and man.
3. Nerve cells and fibres, fat cells, Purkinje fibres of the heart andso on were shown to belong to other category, where the size varies
considerably from one animal to another.
4. The problem of cell size in general was discussed, with special
remarks on the difference between two groups of cells just mentioned.

298

H. HOSOKAWA AND T. SEKINOThe authors should like to acknowledge the advise and encouragement given by Prof. T. Ogawa, Director of anatomy department ofTokyo University.

REFERENCES
ABE, T. (1953). A biometrical study on the glomeruli in the normal kidneys of man andseveral mammals. (in Japanese) Jap. J. Urology, 44: 243-287.
AKIYAMA, S. (1928). Postnatal growth of adrenals in white rats. (in Japanese) Acta Anat.
Nipponica, 1: 401-418.
ALFERT, M. (1955). Quantitative cytochemical studies on pattern of nuclear growth. Symposium on fine structure of cells, 157-163.
AMELUNG, E. (1893). Ueber mittlere Zellengrossen. Flora, 77: 176-209.
ARATAKI, M. (1926). On the postnatal growth of the kidney, with special reference to thenumber and size of the glomeruli (albino rat). Amer. J. Anat. 36: 399-436.
BARGMANN, W. (1936). Die Lungenalveole. v. Mollendorff's Handbuch d. mikrosk. Anatd. Menschen, Bd. 5, 3. Tei!.
. ____ (1939). Langerhanssche Inseln. ibid. Bd. 6, 2. Tei!.
(1956). Histologie u. mikroskopische Anatomie des Menschen. (2. Aufi.)
BENNINGHOFF, A. (1930). Blutgefiiss und Herz. v. Mollendorff's Hundbuch, Bd. 6, 1. Tei!.
BENSLEY, R. R. (1911). Studies on the pancreas of the guinea pig. Amer. J. Anat. 12:
297-388.
BEREZOWSKJ, A. (1910). Studien iiber die Zellgrosse. 1. Mitt. Ueber das Verhaltnis
zwischen der Zellgrosse und der Gesamtgrosse des wachsenden Organismus. Arch.
Zellforsch. 5: 375-384.
BOVERI, TH. (1905). Zellenstudien. 5. Ueber die Abhangigkeit der Kerngrosse und Zellenzahl der Seeigel-Larven von der Chromosomenzahl der Ausgangszellen. Jena.
BOYCOTT, A. B. (1911). A case of unilateral aplasia of the kidney in a rabbit. J. Anat.
and Physiol. 45: 20-22.
BRAUS, H. (1956). Anatomie des Menschen. Bd. 2, (3. Aufl.)
BUCHER, 0. (1948). Histologie. Bern.
_ _ _ (1954). Caryometric studies of tissue culture. Internat. Rev. Cytol. 3: 69-111.
BUHLER, (1898). (cited from Hardesty, 1902).
CAVAZZANI, (1891). (cited from Hardesty, 1902).
CHAMBERS, K. (1908). Einfluss der Eigrosse und der Temperatur auf das Wachstum unddie Grosse des Frosches und <lessen Zellen. Arch. mikrosk. Anat. 72: 607-661.
CLARK, E. (1913). The number of islands of Langerhans in the human pancreas. Anat. Anz.
43: 81-94.
CLARA, M. (1924). Das Pancreas der Vogel. Anat. Anz. 57: 257-265.
- - - (1928). Untersuchungen der menschlichen Hodenzwischenzellen. Zugleich ein Beitrag zur Kenntnis des rhythmischen Wachstums der Zellen <lurch Verdoppelung ihres
Volumens. Z. mikrosk.-anat. Forsch. 13: 72-130.
_ _ _ (1930). Untersuchungen an der menschlichen Leber. 2. Tei!, Ueber die Kerngriissen
in den Leberzellen. ibid. 22: 145-219.
CONKLIN, E. G. (1912). Cell-size and nuclear-size. J. exp. Zool. 12: 1-98.
DAVIES, F. AND E. T. B. FRANCIS, (1952). The conduction of the impulse for cardiac contraction. J. Anat. 86: 302-309.
DRIESCH, H. (1900). Die isolierten Blastomeren des Echidnenkeimes. Arch. Entw.-mechan.
10: 361-434. (cited from Wasserman, 1929.)
EBNER, v. (1899). Kolliker's Handbuch der Gewebelehre. Bd. 3, (6. Aufl.)
ERDMANN, RH. (1908). Experimentelle Untersuchung der Massenverhaltnisse von Plasma,

COMPA RISON OF THE SIZE OF CELLS

299

Kern und Chromosomen in dem sich entwickelnden Seeigelei. Arch. Zetlforsch. 2:
76-136.
-- (1909). Kern- und Protoplasmawachstum in ihrer Beziehungen zueinander. Erg.
Anat. Entwick.-gesch. 18: 844-893.
(1911). Quantitative Analyse der Zellbestandteile bei normalem, experimentell
verandertem und pathologischem Wachstum. ibid. 20: 471-566.
ERHARD, H. (1912). Studien ii.her Nervenzellen. I. Allgemeine Grossenverhaltnisse, Kern,
Plasma und Glia. Arch. Zellforsch. 8: 442-547.
FIEBIGER, J. (1915). Ueber Eigentii.mlichkeiten im Aufbau der Delphinlunge und ihre
physiologische Bedeutung. Anat. Anz. 48: 540-565.
FRANKENHAUSER, G. (1952). Nucleo-cytoplasmic relations in amphibian development. lnternat. Rev. Cytol. 1: 165-193.
FUJITA, S. (1945-1947). Planimetric studies on the renal corpuscles. I-VIII. (in Japanese)
Juzenkai Zasshi, Bd. 40-42.
GLASER, M. (1926). Ueber die Veranderungen im Pancreas der weissen Maus nach Thyroxininjection. Arch. Entw.-mechan. 107: 98-128.
GREEP, R. 0. (1954). Histology. New York and Toronto.
HARDESTY, I. (1902). Observations on the medulla spinalis of the elephant with someco.nparative studies of the intumes::entia cervicalis and the neurons of the columna.. anterior. J. Compar. Neural. 12: 125-182.
HAMMERLING J. (1953). Nucleo-cytoplasmic relationship in the deve~opment of Acetabularia.
lnternat. Rev. Cytol. 2: 475-498.
HAGGQVIST, G. (1931). Gewebe und Systeme der Muskulatur. v. Mollendorff's Handbuch,
Bd. 2, 3. Tei!.
- - - (1956).
. ibid. Bd. 2, 4. Tei!.
HATAI, S. (1902). Number and size of the spinal ganglion cells and dorsal root fibres in
the white rat at different ages. J. Compar. Neural. 12: 107-123.
HEIBERG, K. A. (1907). Ueber eine erhohle Grosse der Zellen und deren Teile bei dem
ausgewachsenen Organismus, verglichen mit dem noch nicht ausgewachsenen. Anat.
An.z. 31: 306-311, 1907.
- - - (1909). Die Inseln in der Bauchspeicheldriise (Langerhanssche Inseln) nebst kurzer
Uebersicht iiber einige andere neuere Pancreasarbeiten. Erg. Anat. Entw.-gesch. 19:
948-1032.
HEIDENHAIN, M. (1908-1911). Plasma und Zelle. I, II. Jena.
------ (1912). Ueber Zwillings-, Drillings- und Vier!ingsbildungen der Diinndarmzotten,
ein Beitrag zur Teilkorpertheorie. Anat. Anz. 40: 102-147.
- - - - (1913). Ueber die Entstehung der quergestreiften Muskelsubstanz bei der Forelle.
Beitrage zur Teilkorpertheorie II. Arch. mikros!;;. Anat. I. Abt. 83: 427-447.
- - - (1914). Ueber die Sinnesfelder und die Geschmacksknospen der Papilla foliata desKaninchens. Beitr. zur Teilkorpertheorie III. ibid. 85: 365-479.
- - - (1919). Ueber die Noniusfelder der Muskelfaser. Beitr. IV zur synthetischen Morphologie (Teilkorpertheorie). Anat. H. I. Abt. 56: 321-402.
- - - - - (1921). Ueber die teilungsfiihigen Driiseneinheiten oder Adenomeren, sowie ii.her
die Grundbegriffe der morphologischen Systemlehre. Zugleich Beitr. V zur synthetischen Morphologie. Arch. Entw.-mechan. 49: 1-178.
· - - - (1923). Ueber die Entwicklungsgeschichte der menschlichen Niere. Beitr. VI zur
synthetischen Morphologie. Arch. mikrosk. Anat. I. Abt. 97: 581-609.
HERTWIG, G. (1931). Der Einfiuss der Fixierung auf das Kern- und Zellvolumen. Z. mikrosk.-anat. Forsch. 23: 484-504.
HERTWIG, R. (1903). Ueber Korrelation von Zell- und Kerngrosse und ihre Bedeutung fiir
die ge3chlechtliche Differenzierung und Teilung der Zelle. Biol. Zbl. 23: 49-62 and108-119.

300

H. HOSOKAWA AND T. SEKINO

HESS, W. N. AND C. W. ROOT, (1938). Study of the pancreas of white rats of different
age groups. Amer. J. Anat. 63: 489-498.
HIRAI, T. (1943). Comparative-histological study on the heart muscle fibres. (in Japanese)
Acta Anat. Nipponica, 21: 545-549.
Hsu, T. C. (1954). Mammalian chromosomes in vitro. IV. Some human neoplasms. J.
National Cancer Inst. 14: 905-932.
ILL'NG, G. (1905). Vergleicbende histologische Untersuchungen iiber die Leber der Haussiiugetiere. Anat. Anz. 26: 177-193.
INOUYE, M. (1909). Die Harnkanalchen des Rindes und des Tiimmlers. Karl Peter's
Untersuchungen iiber Bau und Entwicklung der Niere. I. Heft, II. Jena.
JACOBJ, W. (1925). Ueber das rhythmische Wachstum der Zellen durch Verdoppelung ihres
Volumens. Arch. Entw.-mechan. 106: 124-. (cited from Pfuhl, 1932.)- - - (1935). Die Zellengrosse beim Menschen. Ein Beitrag zur quantitativen Cytologie.
Z. mikrosk.-anat. Forsch. 38: 161-240.
JUNQUEIRA, L. C. U. AND G. C. HIRSCH, (1956). Cell secretion: A study of pancreas andsalivary glands. Internat. Rev. Cytol. 5: 323-364.
KITTELSON, J. A. (1917). The postnatal growth of the kidney of the albino rat, withobservations on an adult human kidney. Anat. Ree. 13: 385-408.
KOEHLER, 0. (1912). Ueber die Abhangigkeit der Kernplasmarelation von der Temperatur und vom Reifezustand der Eier. Arch. Zellforsch. 8: 272-351.
KOPSCH, FR. (1955). Rauber·Kopsch's Lehrbuch und Atlas der Anatomie des Menschen.
(19. Aujl.)
LANZ, T. v. (1926). Ueber Bau und Funktion des Nebenhodens und seine Abhiingigkeit
von der Keimdriise. Z. Anat. Entw.-gesch. 80: 177-282.
LETULLE. (1897). (cited from Benninghoff, 1930.)
LEVI, G. (1905). Vergleichende Untersuchungen iiber die Grosse der Zellen. Anat. Anz.
27, Erg.-H. 156-158. oder Verh. anat. Gesell. 19. Versammlung, 156-158.
MARCEAU, FR. (1904). (cited from Haggqvi£t, 1931.)
MARCUS, H. (1928). Lungenstudien. V. Vergleichende Untersuchungen iiber die respiratp~ische Oberfliiche und ihre Verhaltnis zum Korpergewicht. Gegenbaur's Morph. Jb.
59. 561-566.
MIZIARSKI. (1900). (cited from Wasserman, 1928.)
MOLLENDORFF, W. v. (1930). Der Exkretionsapparat. v. Mollendorjf's Handbuch, Bd. 7,
1. Tei!.
MOORE, F. C. (1903). The unsymmetrical kidney; its compensatory enlargement. J. Anat.
and Physiol. 38: 71-81.
NAKAMURA, N. (1924). Untersuchungen iiber das Pancreas bei Foeten, Neugeborenen, Kindern
und im Pubertatsalter. Virchow's Arch. 253: 286-349.
0BERSTEINER, H. (1913). Die Kleinhirnrinde vom Elephans und Balaenoptera. Arb. aus d.
Neural. Inst. a. d. Wiener Univ. 20: 145-154.
PAINTER, TH. S. (1928). Cell size and body size in rabbits. J. exp. Zool. 50: 441-453.
PETER, K. (1909). Untersuchungen iiber Bau und Entwicklung der Niere. I. Heft. Die
Nierenkanalchen des Menschen und einiger Siiugetiere. Jena.
PFUHL, W. (1932). Die Leber. v. Mollendorjf's Handbuch, Bd. 5, 2. Tei!.
PUTTER, A. (1927). Die Nierenindex. Ein Beitrag zur Kritik der Methoden physiologischanatomischer Forschung. Z. Anat. 83: 228-240.
RABL, C. (1899). (cited from Wassermann, 1929.)
ROHRBACHER, H. (1927). Untersuchungen iiber die Grosse der Zellen in ihrer Beziehung
zur Korpergrosse, Alter, Rasse und Geschlecht. Z. Zierziichtung, 9: 163-206.
(cited from Wassermann, 1929.)
SACHS, J. (1893). Physiologische Notizen. VI. Ueber einige Beziehungen der spezifischen

COMPARISON OF tHE SIZE OF CELLS

301

Grosse der Pflanzen zu ihrer Organisation. Flora, 77: 49-81.
SCHIEBLER, TH. H. (1953). Herzstudie. I. Histochemische Untersuchung von Purkinjefasern
bei Siiugern. Z. Zellforsch. 39: 152-167.
SCHIEFFERDECKER, P. (1916). Untersuchung des menschlichen Herzens in verschiedenen
Lebensaltern in bezug auf die Grossenverhiiltnisse der Fasern und Kerne. Pjluger'sArch. ges. Physiol. 165: 499-564.
SCHULZE, FR. E. (1906). Beitriige zur Anatomie der Saugetierlungen. Sitzgsber. preuss.
Akad. Wiss., Physik.·math. Kl. VI, 225-243. (cited from Marcus, 1928.)
ScHWEIGGER-SEIDEL, F. (1865). Die Nieren des Menschen und der Saugetiere in ihrem
feineren Bau. Halle.
SPECTOR, W. S. (1956). Handbook of biological data. Table 145. (p. 174.)
STIEVE, H. (1926). Die regelmassigen Veriinderungen der Muskulatur und des Bindegewebes
in der menschlichen Gebiirmutter in ihrer Abhiingigkeit von der Follikelreife und derAusbildung eines gelben Korpers, nebst Beschreibung eines menschlichen Eies im
Zustand der ersten Reifeteilung. Z. mikrosk.-wnat. Forsch. 6: 351-397.
STRASBURGER, E. (1893). Ueber die Wirkungssphare der Kerne und die Zellgrosse. Histologische Beitriige, H. 5, 95-124.
SWIFT, H. (1953). Quantitative aspects of nuclear nucleoproteins. Internat. Rev. Cytol.
2: 1-76.
TRETJAKOFF, D. (1928.) Die Riesenwuchs der Knorpelzellen. Anat. Anz. 65: 241-255.
VIMTRUP, BJ. (1928). On the number, shape, structure, and surface area of the glomeruli
in the kidney of man and mammals. Amer. J. Anat. 41: 123-151.
Voss, H. (1928). Die Kerngrossenverhiiltnisse in der Leber der weissen Maus. Z. Zellforsch.
7: 187-200. (cited from Pfuhl, 1932.)
WASSERMANN, F. (1928). Wachstum und Vermehrung der lebendigen Masses. v. Mollendorff's
Handbuch, Bd. 1, 2. Tei!.
WILSON, E. B. (1925). The cell in development and heredity. New York.