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QP91 .H7S^'''°°°'uLin.esuga, SUGAR IN THE BLOOD OF PIGEONS
RECAP
BY
HANNAH ELIZABETH HONEYWELL
A DISSERTATION
Submitted in Partial Fulfillment of the Requirements for the Degree
OF Doctor of Philosophy in the Faculty of Pure Science,
Columbia University, New York City
1921
Qpq
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intljeCttpoflrttigork
CoIIese of Ij^^psimns mis burgeons
Hibrarp
STUDIES IN THE SUGAR IN THE BLOOD OF PIGEONS,
BY
HANNAH ELIZABETH HONEYWELL
A DISSERTATION
Submitted in Partial Fulfillment of the Requirements for the Degree
OF Doctor of Philosophy in the Faculty of Pure Science,
Columbia University, New York City
1921
Reprinted from The American Journal of Physiology
Vol. 58, No. 1, November, 1921
W7r
STUDIES OF THE SUGAR IN THE BLOOD OF PIGEONS
HANNAH ELIZABETH HONEYWELL
From the Department of Physiology, Columbia University
Received for publication July 1, 1921
WTiile it is a recognized fact that much of our experimental data
in physiology must be obtained from animals other than man, there
has been very little hesitation on the part of many experimenters in
drawing conclusions concerning phenomena in man from data derived
from other animals. In many cases the results have warranted the
practice, in others disappointment has resulted. Actually the larger
the number of experiments performed, and the greater the number of
species from which the data have been derived, the more justification
there is for a generalized statement, and for its application to species
other than those upon which work has been done. With this idea
in mind it was thought wise to make the determinations leading to the
data published in the present paper, on an animal hitherto little used
in metaboUsm work. A bird rather than a mammal was chosen
largely because of the fact that in this form the red blood corpuscles are
nucleated. That this makes a striking difference in the physiology of
the blood is well illustrated by Warburg's (1) work on oxygen consump-
tion in the drawn blood of birds as compared with that of mammals.
The fact that the red corpuscles of birds are nucleated should prove of
especial value in studies made on them with the idea of using the re-
sults as a means of giving a better understanding of the physiology of
the cells of tissues other than blood. A considerable literature has
developed upon the exchange of material between the red cells and the
plasma. This is especially true of the chlorides and the sugars. The
data upon which this literature is based have been obtained from
studies on the blood of mammals. It seems probable that parallel
studies on bird 's blood with its nucleated cells should lead to a better
understanding of this process.
The fact that the birds, characteristically, have a higher temperature
than mammals is an additional reason why their metabolism should be
studied for comparison with that of mammals. In fact, it has been
152
BLOOD SUGAR OF PIGEONS 153
suggested by Bierry (2) that this is the reason for the very high con-
centration of sugar found in their blood as compared with that found
in the blood of mammals. That there is a correlation between body'
temperature and blood sugar concentration is shown by the work of
Hollinger (3) and others.
Since most of these properties are of general interest the bird should
be an instructive addition to our list of laboratory animals. Com-
paratively little has been done on the metabolism of birds aside from
some work on polyneuritis. It is therefore advisable to extend our
knowledge of this phase of their physiology.
From birds in general the pigeon was selected and it is thought
that it should be a practical and convenient bird with which to work
for the following reasons: a, it is of convenient size; h, it is comparatively
easy to obtain; c, its first cost is not prohibitive; d, it is easily and
economically kept; e, it is a seed eater and therefore herbivorous.
Further, in the use of the carrier pigeon there is an opportunity for the
study of fatigue. Mosso (4), in fact, has already made some use of it
for this purpose. This phase of the work we hope to extend, and con-
sequently further data on resting ])irds are necessary for comparison
with those obtained from fatigued birds and from resting and fatigued
mammals.
Method: 1. The care of the birds. The pigeons were confined in
the laboratory until they became accustomed to their surroundings.
They were subjected to frequent handling in order to reduce their fear
of the operator, and so to minimize as far as possible results which might
arise from fright. In most cases the birds wholly ceased to resist
handling when taken from the cage. During periods between experi-
ments the pigeons were kept in a large enclosure giving them oppor-
tunity to fly about freely. The food consisted of a mixture of com, .
oats, barley and some other grains, and a plentiful supply of fresh
water. Access was given to both food and water at all times not
otherwise noted.
2. Method of estimation of sugar. MacLean's (5) micro-method was
used throughout the research. The macro-method had been success-
fully used in this laboratory for other blood sugar determinations.
The micro-method possesses the obvious advantage of requiring only a
small quantity of blood (0.2 cc). It therefore may be used for animals
having a comparatively small amount of blood, and it permits of
drawing consecutive samples at relatively short intervals, without pro-
ducing serious effects from hemorrhage. The extreme limits of error
154 HANNAH ELIZABETH HONEYWELL
were found to be about 10 mgm. per 100 cc. of blood. Variations of
this amount or less may, therefore, be disregarded.
3. Special teGhniqiie. While the samples of blood were being drawn
the birds were encased in a strong cloth jacket made especially for the
purpose. This was done as a matter of convenience to the operator
and of safety to the birds. The jacket laced across the ventral surface
of the body in such a manner as to be adjustable to birds of varying
size, and to secure the wings and legs.
The blood was drawn directly from the heart into a 0.2 cc. pipette
by means of a long hypodermic needle. This needle was inserted at
the end of the breast bone, going directly through the skin and body wall
into the abdominal cavity. The tip of the needle followed the breast
bone up to the region of the heart, and so avoided puncture of the liver
and other viscera. When the heart could be felt at the tip of the
needle the point was dropped slightly and plunged into the heart tissue.
Following this procedure the needle struck the heart in such a way
that arterial blood was drawn. Post-mortem examination showed
that the needle usually entered the heart near the apex on the left
side. Hj^podermic needles, 20 gauge, and 3 inches long, were used
most successfully. Since the distance from the tip of the breast bone
to the heart is very neaily equal to the length of such a needle, it is
obvious that this method of drawing blood would not be practicable
for birds much larger than the pigeon. It is also probable, because of
the lesser quantity of blood and the smaller and more delicate heart,
that this method would not be applicable to species much smaller.
All the experiments were performed during the fall and winter of
1920-21.
Study of the concentration of sugar in the blood of indi-
vidual BIRDS. It is a common practice in physiological labora-
tories to keep the experimental animals for only a short time. While
there are probably many cases where repeated observations have
been made on individuals, they have not been published with this
point in view, and as a result it is difficult to find data showing whether
or not the concentration of the sugar in the blood is approximately
constant, or whether it is markedly variable from time to time in one
animal, and from individual to individual.
In order to determine these points for the pigeons, two were set
apart to be used exclusively for this study. They were kept in the
laboratory throughout the period of ol>servation and had access to
food and water at all times. The observations were extended from
BLOOD SUGAR OF PIGEONS
155
October 29, 1920 until the death of pigeon A on February 20, and until
March 13, 1921 on pigeon B. The first six observations were made at
intervals of one week. Following this the intervals were lengthened
as indictited in table 1.
From this table it will appear that pigeon A, in all but four of the
eleven determinations, had a sugar content of 175 mgm. or 180 mgm.
per 100 cc. of blood; pigeon B, with four exceptions, 170 mgm. or
175 mgm. per 100 cc. These variations are well within the limits of
error for the method, and so are to be considered constant.
TABLE 1
Blood sugar of normal birds
October 29...
November 5.
November 13
November 20
December 5 . .
December 11.
December 18,
January 8 . . . .
January 15 . . .
February 6 . . .
February 20 .
March 13
Weight
grams
317
325
327
330
330
335
335
337
340
337
342
Glucose
per 100 cc.
mgm.
200
175
175
180
150
180
175
180
175
185
195
Weight
grams
368
372
375
380
377
375
370
375
377
375
377
375
Glucose
per 100 cc.
mgm .
140
170
170
175
215
175
170
170
175
165
190
175
Obviously, then, there is a concentration of sugar which is charac-
teristic of the blood of a given bird. The fact, however, that the two
birds which happened to be selected yielded characteristic values so
close together, does not warrant the extension of this value to other
birds, as will appear later. But while it is evident that there is a
value which is characteristic of a given bird, it is also evident that
this value will not necessarily be obtained at all times. The occa-
sional striking variations shown in the table illustrate the fact that here
we are dealing with an organism which is responsive to modified condi-
tions, and one of its means of adjustment to its environment involves
changes in the concentration of blood sugar. This offers experimental
confirmation of the statement of Pike and Scott (6) that the concen-
tration of sugar in the blood is one of those internal conditions existing
THE AMERICAN JOURNAL OF PHYSIOLOGY, VOL. 58, NO. 1
156
HANNAH ELIZABETH HONEYWELL
in higher organisms which are generally constant, and which are
regulated by the general nervous and physico-chemical mechanisms
of the organism as a whole.
Three cases offered an opportunity for comparing tlie blood sugar
concentration given by individuals at various times after a period of
48 hours' inanition. These data are given in table 2, from which it
will be observed that each bird has a value which appears to be
characteristic for it.
TABLE 2
Blood sugar of individuals after inanition
BLOOD SUGAR PER 100 CC.
DATE
Normal
After inanition
rngvi.
mgm.
/
10-11
190
165
' 1
12-17
175
• {
10-22
185
165
1- 7
185
20 1
11-4
140
100
12-18
135 •
TABLE 3
Blood sugar of normal dogs at different times
DOG
DATE
BLOOD SUGAR PER 100 CC.
mgm.
^ {
11- 6
75
1- b
73
" {
10-31
67
11- 7
65
= {
11-20
59
1-16
63
These results are similar to those reported by Scott and Hastings
(7) for dogs. Two determinations each were made on three different
dogs, the greatest difference between consecutive determinations upon
the same dog being 4 mgm., as shown in table 3.
In a series of 22 blood sugar determinations made by Kramer and
Cofl&n (8) on a quiet dog, the amount of glucose varies from 87 mgm.
to 93 mgm. per 100 cc, the average being 89 mgm. per 100 cc.
BLOOD SUGAR OF PIGEONS
157
Jones (9) states that in making repeated observations on individual
rabbits the variations were within the experimental error, and so could
be considered as constant. She frequently found, however, a con-
siderable variation in passing from individual to individual.
In a recent paper Strouse (10) reported a series of observations on
five normal persons covering a period of 8 months. In the series the
variations for individuals range from 27 mgm. to 59 mgm. per 100 cc.
of blood with an average variation of 41 mgm. per 100 cc. Thallinger
(11) made repeated observations on a boy with furunculosis, whose
blood sugar was abnormally high, varying from 145 mgm. to 155 mgm.
per 100 cc. on a liberal diet which was low in carbohydrates.
Thus it will be seen that the pigeons agree with the rabbits and dogs
in possessing a characteristic sugar value. Strouse 's figures would
TABLE 4
Effect of excitement on blood sugar
BIRD
DATE
BLOOD
SUGAR PER
100 CC.
REMARKS
mgm.
4
/
I
10- 9
1- 7
185
265
Excited by presence of several strangers
8
{
10-22
10-22
185
300
Excited by loud talking
10
{
11-19
11-20
175
250
Excited by escape from cage
indicate that the range may be greater in man, although it is not clear
whether this greater range is due to a more ready response to changes
in the environment than occurs in the other animals studied, or.
whether the sugar-controlling mechanism is not so perfectly developed,
or whether possibly the conditions of hving were not so carefully
standardized
Effect of handlixg; emotional glycemia. As noted in the
foregoing, noise, loud talking and the presence of strangers produce a
rise in the blood sugar. Rough, sudden or uncertain handling also
disturbs the pigeon and increases the blood sugar. In one instance
the bird escaped from the cage just before a sample was drawn and
some confusion attended its recapture. The amount of sugar in the
sample of blood was high, as shown in no. 10, table 4.
158 HANNAH ELIZABETH HONEYWELL
From this table it will be seen that the bird offers no exception to the
principle long ago pointed out by Boehm and Hoffman (12), Pavy
(13) and others, and more recently by Cannon (14), Shaffer (15) and
Scott (16), that to obtain blood sugar figures of value, samples of blood
must be obtained without pain or other emotional disturbance of the
subject. The fact, however, that birds which were known to be ex-
cited have such high figures as appear in table 4, indicates that the
lower figure of about 185 mgm. per 100 cc. may be assumed to be normal.
This was the value obtained by Scott and Honeywell (17) and though
Fleming (18) found a much lower value for ducks, in fact a figure quite
comparable with that characteristic of mammals, the normal value for
the pigeon, at least, appears to be much higher and to agree well
with the values published for other birds (cf. Scott and Honeywell).
Effect of inanition. For reasons which will be discussed later,
the birds were subjected to a 48-hour period of inanition in determining
the alimentary glycemia curve to be described in the following section.
In all cases the concentration of sugar in the blood was determined soon
after the arrival of the birds at the laboratory, and again at the close
of the 48-hour fast and just before feeding the glucose. While the
effect of this inanition was not the primary purpose of the experiment,
this procedure offered opportunity for its study, provided that the
initial values can be taken as normal values for birds on full feed. The
propriety of this is in some doubt, as the birds had not yet become fully
accustomed to their new environment and had not been subjected to
standard conditions. This would probably result in rather wide
variation from values characteristic for the individual with a general
tendency to yield high values. It is felt, however, that the figures as
they stand merit some attention.
As noted by Rogers (19), the general effect of inanition on the normal
pigeon is to increase its natural restlessness. It becomes irritable and
fights on the slightest provocation, such as a sudden noise. This
might lead one to expect higher blood sugar values in birds subjected
to inanition, and may explain those values which are even higher than
normal that were occasionally found.
From table 5, which contains the data for the blood sugar of pigeons
after inanition, it appears that in 36 experiments 19 pigeons show a
lower blood sugar after inanition than before. The results vary from
3.1 per cent to 70 per cent below the initial value. Fifteen pigeons ex-
hibited an increase in blood sugar ranging from 5 per cent to 100 per
cent. Two pigeons showed no change whatever. The entire series
BLOOD SUGAR OF PIGEONS
159
TABLE 5
Effect of inanition
WEIGHT
BLOOD SUGAR PER 100 CC.
DATE
PERIOD
OF
INANITION
BIRDS
Before
inani-
After
inani-
Per
cent of
Before
inani-
.^fter
inani-
Per cent
of
tion
tion
loss
tion
tion
difference
hours
grams
grams
mgm.
mgm.
10
November 17 to 19
48
280
250
10.7
200
175
- 8.00
11
November 17 to 19
48
350
320
8.5
175
120
-31.0
12
November 17 to 19
48
300
240
20.0
155
150
- 3.1
13
December 15 to 17
48
300
280
6.6
160
100
-37.6
14
December 15 to 17
48
310
275
11.3
105
90
-14.2
5
December 15 to 17
48
340
325
4.4
190
175
- 7.8
9
January 5 to 7
48
280
272
2.8
140
185
32.0
17
January 5 to 7
48
325
312
4.0
185
265
43.0
18
January 5 to 7
48
300
290
3.3
175
310
77.0
19
January 5 to 7
48
350
330
5.6
150
290
93.0
20
November 2 to 4
48
280
248
11.4
140
100
-28.0
21
November 2 to 4
48
310
280
9.65
120
105
-12.5
22
December 8 to 10
48
330
305
7.6
200
170
-15.0
23
November 10 .to 12
48
340
310
8.8
155
155
0.0
14
November 10 to 12
48
330
315
4.5
105
210
100.0
25
November 10 to 12
48
230
215
6.5
105
170
62.0
26
November 10 to 12
48
290
250
13.8
255
155
-39.0
27
December 2 to 5
48
345
340
1.4
120
150
25.0
28
December 2 to 5
48
330
305
7.6
150
140
- 6.6
29
December 8 to 10
48
365
340
6.8
120
175
46.0
8
December 16 to 18
48
3;3'0
300
9.1
185
55
-70.0
9
December 16 to 18
48
300
262
12.6
140
135
- 3.5
4
December 16 to 18
48
300
290
3.3
185
100
-46.9
33
December 20 to 22
48
340
330
2.9
160
175
9.4
12
December 20 to 22
48
330
317
3.9
155
155
0.0
11
December 20 to 22
48
300
285
5.0
175
150
-14.2
36
December 19 to 21
48
300
280
6.7
185
110
-40.6
37
December 19 to 21
48
300
285
5.0
190
160
-15.8
38
December 19 to 21
48
380
340
10.5
185
115
-37.9
9
December 19 to 21
48
350
340
2.8
140
180
28.6
2
September 11 to 14
48
220
195
11.3
290
300
3.4
3
September 11 to 14
48
216
200
7.4
190
200
5.2
8
October 20 to 22
48
356
300
15.7
185
200
8.1
9
October 20 to 22
48
360
280
22.2
187
165
-11.7
51
October 20 to 22
48
330
272
17.6
195
205
5.1
52
November 2 to 4
48
340
320
5.9
140
160
14.3
Aver
age
166
165
- 0.6
*^o^ •
160 HANNAH ELIZABETH HONEYWELL
gave an iaverage decrease in blood sugar of 0.6 per cent. It may,
therefore, be that 48 hours inanition has practically no effect on the
blood sugar of the pigeon. As pointed out above, the evident irri-
tabihty of the birds subjected to inanition with its possible effect upon
the concentration of sugar in the blood should be borne in mind.
Effect of ingestion of glucose: 1. Special technique and dis-
cussion. As noted in the previous section, the sugar was determined
upon the arrival of the birds in the laboratory. Also as described
above, after the birds had become accustomed to the laboratory, they
were subjected to a fast of 48 hours and the sugar in the blood again
determined. The results of these two . determinations were given in
table 5. In addition to the reasons usually assigned for a preliminary
period of inanition, this somewhat prolonged period seemed to be
necessary to empty the crop and so to insure a rapid passage of the
sugar to the region of the alimentary tract where absorption might be
expected to take place. It will be readily appreciated that this is even
more essential in the case of such birds as the pigeon, which are pro-
vided with crop and gizzard, neither of which is presumably a region
of absorption, than it is with the mammals, and possibly than it would
be with other birds. The alimentary canal was empty in all birds
examined, so this period of inanition may be considered as sufficient
to fulfill its purpose.
After the second sugar determination, the appropriate amount of
glucose was administered. To facilitate the feeding, the glucose was
made into tablets and a weighed amount, 1, 2 or 3 grams, according to
the series, was given to each bird. In feeding the sugar, the beak was
opened and the tablets were dropped well back into the haouth. If
the pellets were not readily swallowed, a little water was given through
a dropper. Sometimes gentle stroking of the throat seemed to aid
when swallowing wa§ especially slow.
Three series of experiments were carried out. In series I, each
pigeon was fed 1 gram of glucose;. in series II, 2 grams; and in series
III, 3 grams. In terms of grams per kilogram of body weight, the
average amount of glucose fed was 4 grams, 7 grams, and 10 grams
in the respective series.
The ordinary clinical test for carbohydrate tolerance is 100 grams or
about 1.4 grams per kilogram of body weight, if the average weight for
man is taken to be 70 kilograms. This amount was fed by Cummings
and Piness (20), Hiller and Mosenthal (21), Jacobsen (22), Tachau (23)
and Strouse, who has also fed 2 grams and 2.8 grams per kilo to normal
men. Jones gave rabbits an average dose of 7.87 grams per kilogram.
BLOOD SUGAR OF PIGEONS
161
From the foregoing it will be seen that the amounts given to the
pigeon exceed those usually given man in similar experiments. In
spite of this, the smallest dose used in the present experiments which
is equivalent to one of 1 .75 grams for a man weighing 70 kilograms, had
very little effect on the concentration of sugar in the blood of the
pigeon. In a man such an amount would in all probability raise the
concentration of blood sugar to 200 mgm., and probably induce gly-
cosuria.
2. Time of the maximimi. Since it was desired to determine the
principal points in the entire curve, that is, to follow the curve to its
return to the initial value, and since the number of samples of blood
which could be drawn safely in any one experiment was limited be-
cause of the injury to the heart which would result from repeated
TABLE 6
Time of maximum of alimentary glycemia
BLOOD SUGAR
BLOOD SUGAR PER 100 CC. AFTER FEEDING GLUCOSE
BIRD
GLUCOSE FED
AFTER
INANITION
1 hour
2 hours
3 hours
4 hours
5 hours
grams
mgm.
mgm.
mgm.
mgm.
mgm.
61
3
160
155
240
400
335
320
62
3
180
185
235
305
210
215
64
2
210
205
245
315
275
260
65
2
185
200
215
240
250
205
66
1
170
190
200
215
240
200
67
1
165
170
185
205
225
190
punctures at short intervals, it was necessary to determine the time
elapsing between the administration of the glucose and the maximum
sugar concentration in the blood. For this purpose, as indicated in
table 6, hourly determinations were made after the sugar was fed. It
will be seen from the results given in this table that the maximum may
be assumed to occur between the third and -fourth hours. This last
interval was therefore allowed to elapse after feeding and before drawing
the first sample, and the sugar level at this time may be assumed very
nearly to represent the maximum attained.
When 3 grams of glucose were fed the maximum occurred at or about
the third hour. When 2 grams of glucose were fed the maximum
occurred in one case at the third hour and in the other case at the fourth
hour. After the feeding of 1 gram of glucose, the maximum occurred
at about the fourth hour. From these results, given in table 6, it will
162
HANNAH ELIZABETH HONEYWELL
appear that the greater the amount of glucose fed the earlier the maxi-
mum will be reached.
In this connection it is interesting to note, as Strouse has pointed out,
that a heavy dosage often has the effect in man of delaying the onset of
the maximum rather than accelerating it as in the pigeon. There
must be, then, some fundamental difference between the carbohydrate
economy of the pigeon and that of man.
3. Course of alimentary hyperglycemia. In each case samples of
blood were drawn just before the administration of the glucose and
again after the lapse of 4, 6 and 24 hours. The results are collected in
J.GO
l¥
tables 7, 8 and 9 and summarized in the accompanying curves (fig. 1).
In the first series one gram of glucose was given each bird. As noted
before, this is about double the ratio of the test meal usually given
man for diagnostic purposes. From table 7 it will appear that there is
no change or as occurs more frequently, only a slight rise at the end of
the first period. The average for the series gives a rise of 18 per cent
at this time.
In the second and third series there is a different manifestation. In
the second series 2 grams, and in the third 3 grams were given each bird.
The average rise at the end of 4 hours in the second series was 45 per
cent, and in the third, 93 per cent. From table 8 it will be seen that
BLOOD SUGAR OF PIGEONS
163
m the second series only five birds had returned to their previous level
at the end of 24 hours; and in the third series, table 9, one alone had
returned in that interval to the level which existed before the ingestion
of the glucose.
While the average at the end of the inanition period may vary some-
what for the different groups, the average at the end of 24 hours after
feeding the glucose is approximately that of the normal birds. Be-
cause of the relatively low initial values found in the second and the
third series, the final values found for these series are distinctly higher
TABLE 7
Effect of ingestion of glucose upon the sugar in the blood
Series I
GLU-
BLOOD SUGAR PER 100 CC.
COSE
AFT EI
I FEEDING GLUCOSE
BIRD
DATE
WEIGHT
FED
PER
KILO
OF IN-
CREASE
REMARKS
Inani-
tion
4
hours
6
hours
24
hours
grams
(jrams
mgm.
mgm.
mgm.
mgm.
10
November 19
250
4.0
175
100
110
250
-37.0
Bird excited by
11
November 19
320
3.1
120
150
150
125
25.0
escape from
12
November 19
240
4.2
155
150
160
150
3.2
cage before
13
December 17
■ 280
3.5
100
275
245
195
175.0
drawing of 24
14
December 17
275
3.6
90
210
300
270
233.0
hour sample
5
December 17
325
3.1
175
170
160
230
- 2.8
9
January 7
272
3.7
185
270
175
265
46.0
17
January 7
312
3.2
265
280
275
275
5.7
18
January 7
290
3.4
310
350
300
250
1.3
19
January 7
330
3.0
290
295
260
155
1.7
Average
259
3.9
190
225
175
180
18.0
than the initial values. Whether or not this is significant we are not
prepared to state definitely, although it would seem to be accidental.
Unfortunately there is very little data available which permits of a
comparison of the course of the alimentary hyperglycemia of different
species. In fact, only three species seem to have been studied from
this point of view. In her recent paper, Jones has made determinations
on the blood of rabbits only at a single period after the ingestion of the
glucose. From the work of Bang (24), the 1-hour period which she
chose would presumably give figures at or near the maximum attained.
Her results do not, however, permit one to follow the course of the curve.
Bang reported a short series of experiments on rabbits which had been
164
HANNAH ELIZABETH HONEYWELL
fed from 5 to 20 grams of glucose. After a S-daj^ period of inanition he
found that the maximum was reached in 1| to 2^ hours, and that in
general the sugar level had returned to normal in 6 hours. The amount
of sugar did not seem materially to modif}^ the time relations of the
curve. The same may be said of a similar but even shorter series,
in which sugar was given without a previous period of inanition.
Fisher and Wishart (25) in experimenting with dogs weighing 8 to
9 kilograms, fed approximately 6 grams of glucose per kilogram, and
found that the maximum blood sugar occurred one hour after the
ingestion of the glucose.
TABLE 8
Series II
BIRD
DATE
WEIGHT
GLUCOSE
FED PER
KILO
BLOOD SUGAR PER 100 CC. .
FEEDING GLUCOSE
A.FTER
PER CENT
OF IN-
Inani-
tion
4 hours
6 hours
24 hours
CREASE
grams
grams
Tngm.
mgm.
mgm.
mgm.
20
November 4
248
8.4
100
150
125
295
50.0
21
November 4
280
7.1
105
300
250
180
185.0
22
December 10
305
6.5
170
290
280
2b0
70.0
23
November 12
310
6.4
155
440
140
145
184.0
14
November 12
315
6.3
210
210
■ 250
240
19.0
25
November 12
215
9.3
170
200
190
120
17.0
26
November 12
250
8.0
155
250
210
160
61.0
27
December 4
340
5.9
150
280
100
130
87.0
28
December 4 .
305
6.5
140
200
285
175
103.0
29
December 10
340
5.9
175
270
160
110
54.0
Aven
lee
290
6.9
165
240
195
180
45.0
The work of many investigators, notably Cummings and Piness,
Hiller and Mosenthal, Hamman and Hirschman (26), Jacobsen and
Strouse, indicates that after the ingestion of 100 grams of glucose the
maximum concentration of sugar in the blood of man occurs normally
in about 30 minutes and that it has returned approximately to its
previous level by the end of the second hour. Jacobsen and Strouse
point out that occasionally the maximum is attained only after a longer
period; and that when this is true the level is apt to be higher than usual,
and the return to the previous value is usually slower.
Strouse particularly calls attention to the fact that in diabetes and
other conditions which may be presumed to alter the carbohydrate
metabolism, such curves are common but that such individuals may be
BLOOD SUGAR OF PIGEONS
165
induced to give the ''normal" or usual curve if given less sugar. On
the other hand normal individuals will give the ''diabetic" curve if
the dose be doubled or tripled.
A study of the curves obtained from pigeons shows that the maximum
occurs from the third to the sixth hour after feeding, and when amounts
were fed which essentially altered the sugar level, the curve did not
return to normal for a much longer period, in some cases exceeding
2-4 hours. Thus they resemble more closely the delayed curves ob-
tained from men rather than the usual or normal one, and, at first
thought, the obvious reason is the very heavy dose of glucose given to
the birds.
TABLE 9
Series III
GLU-
COSE
BLOOD SuUaR per 100 CC.
AFTER FEEDING GLUCOSE
PER
1
BIRD
DATE
WEIGHT
FED
PER
KILO
1
OF IN-
CREASE
REMARKS
Inani-
tion
4
hours
6
hours
mgm.
24
hours
grams
grams
mgm.
mgm.
mgm.
8
December 18
300
10.0
55
215
200
205
291.0
20
December 18
262
11.4
135
320
150
160
137.0
Excited by pres-
4
December 18
290
10.3
100
160
300
22Q
200.0
ence of strangers
33
December 20
330
9.0
175
335
200
110
91.5
when third sam-
12
December 20
317
9.4
155
240
210
225
55.0
ple was shown.
11
December 20
285
10.5
150
320
195
175
113.0
Struggled and died
36
December 21
280
10.7
110
225
205
260
104.5
during drawing
37
December 21
285
10.5
160
200
265
180
65.5
of last sample.
38
December 21
340
8.8
115
265
110
195
130.4
9
December 21
340
8.8
180
330
220
205
210
180
84.0
Average
303
9.8
1
135
260
93.0
In order to determine this point, that is, whether the pigeons would
respond to a smaller dose and whether or not a maximum occurring
during the first hour had been overlooked, four birds were fed 0.4
gram of glucose each, after an inanition period of 48 hours. This is
the amount of glucose which, for the weight of the bird, approximates
the usual test dose for man. Blood sugar determinations were made
30 minutes, 1 hour and If hours after the ingestion of the glucose. The
results are shown in table 10. Since the variations were all within the
limits of experimental error, it may be concluded that the pigeon does
not respond to as small a dose as does man and that there is a fundamen-
tal reason for the difference in the alimentary glycemia curves shown
by the two species.
166
HANXAH ELIZABETH HONEYWELL
In his series on normal men Strouse obtained an average increase of
42 per cent after an ingestion of 100 grams of glucose, or 1.4 grams per
kilogram. The pigeons show an average increase of only 18 per cent
after the ingestion of 1 gram or about 3.5 grams per kilogram, and it
was not until they had been given 2 grams or 7 grams per kilogram that
they approached the percentage increase reported by Strouse for men.
In addition it should, perhaps, be pointed out that the resemblance
is more apparent than real for, as mentioned above, the effect of the
size of the dose upon the time elapsing between the administration of
the dose and the occurrence of the maximum is in the opposite sense
in the pigeon and in man. It would thus seem that the mechanism
of storage of sugar is somewhat different in the two groups.
Post-mortem examinations of two pigeons which were killed after
inanition and before feeding showed that the crop and gizzard were
TABLE 10
Effect of varying amounts of glucose on time of maximum
BIRDS
WEIGHT
AMOUNT OF
GLUCOSE FED
BLOOD SUGAR IN MGM. PER 100 CC.
FEEDING GLUCOSE
AFTER
PER KILO
Inanition
h hour
1 hour
U hours
grams
grams
1
250
1.6
180
165
180
175
2
370
1.1
220
225
230
225
3
280
1.4
195
200
190
205
4
350
1.1
185
175
180
175
empty, while the intestine contained only a small amount of fluid.
Conditions were the same in pigeons which were examined at the end
of the fourth and sixth hour after feeding. The contents of the ali-
mentary tract were not tested for the presence of sugar. Conse-
quently, while the indications as they stand are that the delay in
reaching the maximum is not due to delay in absorption, but rather to
some peculiarity in the mechanism of storage, one is not justified in
definitely drawing such a conclusion until a study of the contents of the
alimentary canal has been made in parallel with blood sugar deter-
minations.
Since concentration of the sugar in the bird is normally so high as
compared with mammals, in this particular resembling the diabetic,
and since the curve obtained from birds somewhat resembles that ob-
tained from diabetic man, there may possibly be some relationship be-
BLOOD SUGAR OF PIGEONS 167
tween the absolute initial height of the sugar concentration and the
form of the curve of alimentary hyperglj^cemia. However, as noted
above, it would seem more probable that in the birds the storage
mechanism is somewhat different from that common in mammals
and that further work must be done before a satisfactory correlation is
possible.
SUMMARY
1. Each bird has a characteristic sugar level about which it varies
from day to day. In this it resembles the rabbit and dog.
2. These individual variations are caused by variations in the ex-
ternal and internal environment of the bird.
3. A series of inanition values for blood sugar is given and com-
pared with the values found on full diet. From these figures it is
concluded that 48 hours' inanition has practically no effect on the blood
sugar of the pigeon.
4. It has been found that, in general, when from 1 to 3 grams of
glucose are fed to the pigeon the maximum rise in the blood sugar
occurs in 3 to 4 hours.
5. It is indicated that the greater the amount of glucose given the
earlier will the maximum be reached.
6. When 1 gram of glucose or less is fed to pigeons there is very
little modification of the sugar in the blood. When 2 or 3 grams are
fed there is a manifest rise in the blood sugar which gradually
approaches its former level.
The author wishes to acknowledge her indebtedness to Prof. E. L.
Scott for his assistance in the execution of this work.
BIBLIOGRAPHY
Warburg: Hoppe-Seyler's Zeitschr., 1910, Ixvi, 305; Ixxvi, 331.
Bierry: Comptes Rendus, 1919, clxix, 1112.
Hollinger: Deutsche. Arch. f. klin. Med., 1908, xcii, 217.
Mosso: Fatigu, New York, 1904.
MacLean: Biochem. Journ., 1919, xiii, 135.
Pike and Scott: Amer. Naturalist, 1915, xHv, 321.
Scott and Hastings: Proc. Soc. Exper. Biol, and Med., 1920, xvii, 67.
Kramer and Coffin: Journ. Biol. Chem., 1916, xxv, 423.
Jones: Journ. Biol. Chem., 1920, xliii, 507.
Strouse: Arch. Int. Med., 1920, xxvi, 751.
Thallinger: Journ. Amer. Med. Assoc, 1921, Ixxvi, 295.
168 HANNAH ELIZABETH HONEYWELL
(12) BoEHM AND Hoffmann: Arch, f, Exper. Path. u. Pharm., 1878, viii, 271.
(13) Pavy: Journ. Physiol., 1899, xxiv, 479.
(14) Cannon : This Journal, 1914, xxxiii, 356.
(15) Shaffer: Journ. Biol. Chem., 1914, xix, 285.
(16) Scott: This Journal, 1914, xxxiv, 271.
(17) Scott and Honeywell: This Journal, 1921, Iv, 362.
(18) Flemming: Journ. Physiol, 1920, liii, 236.
(19) Rogers: This Journal, 1916, xli, 555.
(20) CmiMiNGS AND PiNESs: Arch. Int. Med., 1917, xix, 777.
(21) HiLLER AND MosENTHAL : Joum. Biol. Chem., 1917, xxviii, 197.
(22) Jacobsen: Biochem. Zeitschr., 1913, Ivi, 471.
(23) Tachau: Arch. f. klin. Med., 1911, civ, 437.
(24) Bang: Der Blutzucker, 1913.
(25) Fisher and Wishart: Journ. Biol. Chem., 1912, xiii, 49.
(26) Hamman and Hirschmann: Arch. Int. Med., 1917, xx, 761.
VITA
Hannah Elizabeth Honeywell was born in Walton, N. Y., April 3,
1888. She graduated from Walton High School in 1906, received
the degree of Batchelor of Arts from Mount Holyoke College in 1910,
and the degree of Master of Arts from Columbia in 1917.
Her publications are:
A Study of the Sugar in the blood of Normal Pigeons. (With
E. L. Scott.) American Journal of Physiology, 1921, Iv, 362.
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