THE DISTRIBUTION OF SUGAR AND RATE OF GLYCOL-

YSIS IN THE BLOOD OF SOME MAMMALS

HY MICHAEL SOMOGYI
(From the Luboratory oj the Jewish Hospital of St. Louis, St. hmis)

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(Received publication, October 16, 1933)
for

The question of the distribution of sugar between corpuscles

and plasma in human blood is fairly clarified, several workers, who
determined true sugar, having arrived at virtually identical re-
sults. Information relating to other species, however, still is
largely confined to data derived from apparent sugar values.
A few examples are given in Table I to illustrate how misleading
such figures are. In the case of sheep blood, for instance, the
apparent sugar values (total reduction in tungstate filtrates) indi-
cate that the distribution ratio, corpuscle sugar to plasma sugar,
equals 0.53, whereas the actual ratio obtained from true sugar
values (fermentable reducing substance in tungstate filtrates), is
0.09. Or, in pig blood the concentration of sugar in the corpuscles
is seemingly 41 per cent of that in the plasma as calculated from
apparent sugar values, while the true sugar values reveal that the
corpuscles contain no sugar at all. Thus, in connection with an
examination of possible relationships between the distribution of
sugar and the rate of glycolysis, it appeared necessary to determine
in terms of true sugar values the distribution in the bloods of the
several species involved.
Distribution of Sugar
In the greater part of these experiments, conducted in 1927 and
1928, the analytical method previously described (1) was followed
in its entirety. In more recent analyses the copper method (2)
was employed for the precipitation of proteins (zinc is unsuitable
for plasma and serum); the filtrates yielded true sugar values
which, in comparative determinations, agreed with the results
obtained in tungstate filtrates by the fermentation technique.
The corpuscle sugar was computed from three experimentally de-
665
666 Distribution and Glycolysis of Sugar
termined quantities: corpuscle sugar per 100 cc. = 100 (whole
blood sugar - plasma sugar X plasma volume)/cell volume.
Direct determination of the sugar content of corpuscles yields

TABLE I
Distribution Ratios Calculated from Apparent and from True Sugar
Values

Apparent supar True sqw (apparent Distribution ratio,

sugar mmus reducing corpu?.ole *ugm:
Species (total reductmn) IFXl-BlgL3CS) serum sugar

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c!orpndea( serum co~Pu.?dea~ serum Apparent True
_ ~-

“%” mztw “~27

Guinea pig. ......... 91 133 40 124 0.68 1 0.33
ox. ................. 45 91 15 8.5
Sheep. .............. 46 a7 7 80
Pig .................. 40 97 0 90

TABLE II
Partition of Sugar between C701rpuscles and Serum in Blood of Various
Mammals
- -
3istri-
True sugar : mtion
content ratio,
Species - Species COP
1 pusole
COr-
wcles skrum B
*
sugar:
wrum
.-
‘f;n”l” m
ztm ““,mPtrn
‘gcknPt@
ox. . 15 8.50.18 cat. . _. 79 331 0.24
Calf. . . 61 136 0.45 Rabbit. 42 144 0.29
“ .,.......... 101 0.36 ‘I
36 . 40 147 0.27
Sheep. . . 13 80 0.16 Guinea pig 40 124 0.33
I‘ ‘I “
. . 7 80 0.09 . .. 67 187 0.36
Pig . 0 90 0 Monkey (Maca
Dog. . 35 150 0.23 cus rhesus). 121 149 0.82
I‘ 0.42 “ ‘I
48 115 122 144 0.85
cat.:::::::::::: ‘I “
73 263 0.28 114 151 0.76
-

acceptable results only in cases where the rate of glycolysis is very

low and consequently the loss of sugar during the separation of the
cells is negligible. The blood samples analyzed were in every
instance fresh; 1 to 1.5 mg. of potassium oxalate per cc. was added
 M. Somogyi
as anticoagulant except in experiments in which glycolysis also
was studied; in these cases the blood was defibrinated.
Table II contains representative examples showing the distribu-
tion of true sugar in the blood of various species, results which
differ considerably from the multitude of data encountered in the
literature. The latter show either much higher sugar concentra-
tions in the corpuscles than we are reporting, or else state that the
blood corpuscles in all or most mammals are entirely sugar-free.
The inconsistencies are brought about chiefly by differences in

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analytical procedure. Total reduction, the value generally de-
termined, is known to include an appreciable fraction of non-sugar
reducing substances, particularly in the protein-free filtrates of cor-
puscles. The magnitude of this fraction is greatly affected by
two factors, namely by the method of deproteinisation and by the
composition of the oxidizing reagent employed. This explains
the figures which are too high. The complete absence of sugar in
corpuscles, on the other hand, is unvariably reported by investiga-
tors who considered it necessary to wash the corpuscles prior to
analysis (Otto, Bang, Lytkens, Glassmann), not suspecting that
such treatment promptly removes the total sugar content of the
corpuscles.
If we disregard the results of such faulty technique, there are
nevertheless cases in which the corpuscles seem to be actually
free of sugar. In a sample of pig blood, for example (Table II),
the corpuscles contained no fermentable reducing substance. In
a beef blood and in two samples of sheep blood the sugar in the
corpuscles was so low as to suggest that these corpuscles too, were
sugar-free and that the slight sugar content found was probably
the result of experimental inaccuracy. Such inaccuracy may slip
in at the determination of the cell and plasma volume and become
a factor of some importance in the calculation of the corpuscle
sugar. We have ascertained that after the customary centrifu-
gation for the determination of the cell volume, the amount of
plasma still adhering to the cells will in some instances suffice t,o
account for the slight sugar content attributed to the corpuscles.
If the separated corpuscles are suspended for a few moments in
isotonic salt solution, then centrifuged, the rinsed off plasma
proteins can be determined in the salt solution, The volume of
this intercellular plasma can thus be calculated, deducted from the
668 lktribution :tnd Glycolysis of Sugar
cell volume, and added to the plasma volume. In cases like the
beef and sheep blood samples under consideration, such correction
reduces the small difference between the two quantities, whole
blood sugar and plasma sugar times plasma volume, to zero or near
zero, indicating that the entire blood sugar is contained in the
plasma and that the corpuscles are wholly devoid of sugar. This
correction was not generally applied in the present study partly
because of the tediousness of the procedure, partly because the
error involved is insignificant when the sugar content of the cor-

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puscles is not very low.
Comparison of the distribution of blood sugar in the several
species examined shows great variations, the quotient, corpuscle
TABLE III
Relation of Rate of Glycolysis to Partition of Sugar and to Rate of Phosphate
Cleavage
T
Phosphate Phosphate
cleavage Ratio cleavage
Species R:Y
distri-
series of Species d&i-
series of
bution En%krdt bution En%!?=dt
Liubimova Liubimova

““,i??rt~ “zt~
Rabbit 28 0.30 Rabbit Calf. 4.5 0.43
Guinea pig 23 0.36 Guineapig Ox. . . .._ 4.5 0.18
Dog...... 22 0.42 Pig. _. 3.5 0 Pig, dog
Cat 21 0.24 Sheep, . 2.5 0.16
Monkey.. 17 0.83 cow
I
Man. 16-E 0.80 Man

sugar/plasma sugar, ranging from 0 to 0.45, values much lower

than are established for human blood. Monkey blood is the sole
exception, with a distribution ratio of 0.76 to 0.85, the same as in
human blood.
Ratio of Glycolysis
A few years ago Loewi espoused the theory (3) that the initial
step in the utilization of glucose in living cells in general is adsorp-
tion of the sugar by the cell membranes, the extent of this ad-
sorption being the determining factor in the rate of the chemical
changes involved. The only proof to support this sweeping con-
cept was based upon experiments in blood glycolysis, rather slender
 M. Somogyi 669

evidence having been offered to the effect that a shift in the dis-
tribution of sugar between corpuscles and plasma in favor of the
corpuscles enhances glycolysis. The subst.antial differences in the
concentrations of corpuscle sugar in the bloods of various animals,
as found in our experiments, prompted us to investigat,e the rela-
tionship between the concentration of corpuscle sugar and the rate
of glycolysis. The results of this investigat.ion, recorded in Table
III, show the absence of any relationship between the two quanti-
ties. In the first column of figures in Table III are presented the

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rates of glycolysis in various species, arranged according to de-
scending magnitudes, the corresponding dist,ribution ratios being
aligned in the second column. It is evident that t,he two ‘series
show no parallelism. Thus the rate of glycolysis in rabbit blood is
50 per cent higher than in human blood, although the relative
sugar concentration in rabbit corpuscles is but one-third of that
in human corpuscles. Or, comparing dog and calf blood it may
be seen that, notwithstanding the identity of the sugar content
of the corpuscles, the rate of glycolysis in dog blood is 5 times as
high as in calf blood.
While at variance with Loewi’s t.heory, our findings are in
general accord with observations made by Engelhardt and Liubi-
mova (4) which came to our attention aft,er this study had been
closed. These aut,hors discovered that t,he rate of glycolysis
runs generally parallel with the ability of the erythrocytes to
liberate inorganic phosphate, i.e. that in bloods in which the rat,e
and extent of the phosphate cleavage is greater, the rate of gly-
colysis also is higher. According to the rate of phosphate cleav-
age, Engelhardt and Liubimova arranged the animals invest,igated
in the following series: cow < goat, horse < dog, pig < man,
guinea pig < rabbit,. When the members of this series are placed
in the last column of Table III, arranged in the order of decreasing
rate of phosphate cleavage, the parallelism between this and the
rate of glycolysis becomes obvious.

SUMMARY

The distribution of sugar bet:ween the corpuscles and plasma in

the blood of various mammals was determined in terms of true
sugar values.
In all the species studied, excepting the monkey (JZncuc~.s
670 Distribution and Glycolysis of Sugar

rhesus), the ratio of the sugar content of corpuscles to that of

plasma is substantially lower than in human blood. In some
cases the corpuscles are entirely free of sugar. Human blood
and monkey blood show the same ratio of distribution.
The rate of glycolysis is not related to the partition of the sugar
between corpuscles and plasma, but runs parallel with the rate of
phosphate hydrolysis in t’he corpuscles (Engelhardt and Liubi-
mova).

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BIBLIOGRAPHY

1. Somogyi, M., J. Biol. Chem., 78, 117 (1928).

2. Somogyi, M., J. Biol. Chem., 90, 725 (1931).
3. Loewi, O., K&n. Woch., 6, 2169 (1927).
4. Engelhsrdt, W. A., and Liubimova, M., Biochem. Z., 227, 6 (1930).
 THE DISTRIBUTION OF SUGAR AND
RATE OF GLYCOLYSIS IN THE BLOOD
OF SOME MAMMALS
Michael Somogyi
J. Biol. Chem. 1933, 103:665-670.

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