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and Vernet- in 1888 ascended together one of the highest points of the Jura.
They found that there was a distinct rise in the rectal temperature. Marcet,
however, does not look upon this result as conclusive ; he attempts to explain
the rise of temperature as due to congestion of the haemorrhoidal vessels. It
must be pointed out, however, that the increased circulation due to exercise
would probably not cause congestion, and, whether it did or not, the rise in
the temperature of the rectum indicates a rise in the temperature of the
internal parts of the body. Further, Marcet himself shows that cooling the
under surface of the chin causes a fall in the temperature of the mouth, and
this was probably the cause of the low readings observed in his first ascents.

ObernierS found that a walk for thirty-five minutes, when the
external temperature was ll°-2, raised the rectal temperature from 37°
to 38°. A walk of five miles raised the temperature of Ogle's mouth
from 37° to 37°"4o.'^ Similar results have been obtained by others.^

Similar results to the above have been obtained upon animals. The
temperature of a dog during the first hour of work upon a treadmill was
raised l°-8, but although the work was continued the temperature quickly
fell (U. Mosso).'^ In the case of two stallions three years old, Liska^ found
the temperature before work 37°"8 and 38°-0 respectively ; after
fifteen minutes' work, 39°-5 and 39°; and again, after twenty minutes'
rest, 37°"7 and 38°. Siedamgrotzky ^ found that exercise raised the
temperature of horses by an amount varying from 0°"3 to 1°, while
Hobday^ found in the case of healthy omnibus horses that the rectal
temperature was generally raised 2° or more by hard work, and in sheep
and pigs the exertion of running caused a similar rise in temperature.

Further details of the production of heat in muscle will be given
later.

In the case of insects the effect of muscular activity is very marked.
Thus iS^ewport '^ found the temperature of the abdomen of a very active humble-
bee {Bombus terrestris) to be 23°, when the air was 19°'3 ; four of these active
bees placed in a glass bottle raised the temperature of the air from 19°*3 to
23°-6.

The influence of mental work. — Mental activity is said to have an
effect both upon the general temperature of the body and upon the local
temperature of the brain and head. Thus Davy ^^ found that mental

^ Arch. d. sc. jjhys. et naf., Geneve, tome xxxvi. p. 247.

- Marcet, Croonian Lectures, Brit. Med. Journ., London. 1895, vol. i. p. 1367.'

3 "Der Hitzschlag," Bonn, 1867, S. 80.

■* St. George's Hosp. Eep., London, 1866, vol. i. p. 232.

^ Crombie, Indian Ann. Med. Sc, Calcutta, 1873, voL xvi. p. 579 ; Roger, " Ee-
cherches cliniques sur les maladies de I'ent'ance," tome i. p. 227 ; Speck, Arch. d. Ver. f.
gemeinsch. Arb. z. Ford. d. luissensch. Heilk., Gottingen 1862, Bd. vi. S. 161-324; Cuny
Bouvier, Arch. f. d. ges. FhysioL, Bonn, 1869, Bd. ii. S. 386.

^ Ellenberger, " Vergleichende Physiologie der Haussaugethiere," 1892, Bd. ii. Tli. 2,
S. 87.

■^ Deutsche Ztschr. f. Thiermed., Leipzig, 1875, Bd. i. S. 87.

* Journ. Comp. Path, and Therap., Edin. and London, 1896, vol. ix. p. 286.

^ Phil. Trans., London 1837, pt. 2, p. 259.
10 Ihid., 1845, pt. 2, p. 319 ; 1850, p. 443.



8o8



ANIMAL HEAT.



work in England and in the tropics raised his temperature 0°"27 and
1°"1 respectively, and an increase varying from 0°"1 to 0°*7 has been
observed after similar exertion by Speck,^ Eumpf ,^ and Gley ^ ; the
temperature was taken in the rectum, axilla, or mouth. Clifford
Allbutt,"* however, in a long series of observations, found that mental
work had no effect upon the temperature.

Cavazzani^ states that in the case of a man whose skull had been
trephined over the right temporo-occipital region, a thermometer placed
in the dura mater showed a rise of two-tenths of a degree during mental
activity. A. Mosso^ maintains that intense psychical processes may
cause so much heat to be set free in the brain that its temperature may
remain for some time 0°'2 to 0°'3 above the temperature of the rectum.
In a curarised dog the action of cocaine may produce a rise of as much
as 4° in the temperature of the brain (37° to 41°). In man, Lombard ^
found that mental activity caused a slight rise in the temperature of the
head, especially in the occipital region.

It is probable, however, that this local rise of temperature is not due,
as Mosso believes, to very active combustion in the ganglion cells, but to
vascular changes consequent upon the mental activity. Hill and
Nabarro,^ have show^n that the blood from the venous sinuses of the
skull is less venous in colour than that of the femoral vein, that the
metabolism of the brain is very low, and that it is scarcely increased
during an epileptic fit. The average differences between the gases
in samples of blood from the carotid artery and from the torcular
Herophili of dogs were as follows : —





Normal.


Tonic Fit.


Clonic Fit.


Art.


Tore.


DiflF.


Art.


Tore.


Diff.


Art.


Tore.


Diff.


Carbon dioxide
Oxygen .


40-86
16-81


44-74
13-39


+ 3-87
-3-42


44-98
15-17


49-04
10-22


+ 4-06
-4-95


30-59
15-77


33-58
11-46


+ 2-99
-4-31



It is probable, therefore, that the temperature of the brain is not
perceptibly greater than that of the blood. The cerebral circulation
changes passively with every alteration of the general arterial or venous
blood pressure,^ and this is apparently the explanation of Lombard and
Mosso's results. Moreover, the experiments of Helmholtz,^^ Heiden-
hain,^^ and Eolleston ^^ have failed to demonstrate the formation of heat
in nerve.



^ Arch. f. ex'per. Path. u. Pharmakol., Leipzig, 1882, Bd. xv.
- Arch. f. d. ges. Physiol., Bonn, 1884, Bd. xxxiii. S. 601.
^ Compt. rend. Soc. de hiol., Paris, 1884, p. 265.
•* Note communicated to the writer.
^ Arch. ital. da hiol., Turin, 1893, tome xviii. p. 328.

^ Proc. Poy. Soc. London, 1892, vol. li. p. 83; "Die Temperature des Gehirns,"
Leipzig. 1894.

^ Arch, de ^ihysiol. norm. etjMth., Paris, 1868, tome i. p. 670.
^ Journ. Physiol., Candiridge and London, 1895, vol. xviii. p. 218.
'■• Roy and Sherrington, ihid., 1890, vol. xi. p. 85 ; Hill, ihid.. 1895, vol. xviii. p. 15.
^^ Arch. f. Anat., Physiol, u. ivissensch. Med., 1848, S. 158.
1^ Stud. d. physiol. Inst, zu Breslau, Leipzig, 1868, Bd. iv. S. 250.
^" Journ. Physiol., Cambridge and London, 1890, vol. xi. p. 208.



INFLUENCE OF FOOD. 809

The influence of food.— The investigations of many observers ^ show
that the effect of food upon the temioerature of the body is to cause a shght
rise, or, in the case of the evening meals, to postpone for a short time
the customary fall of temperature at that time. The rise is often in-
appreciable and rarely exceeds half a degree; the maximal effect is
seen about one hour and a half after the meal. A draught of cold water
(10°) lowers the temperature about half a degree.^

In the case of the horse the effect of food is to cause a rise of 0''-2
to 0°'8, which persists for three or four hours.

Maurel ^ states that in the rabliit food is the chief cause of the daily
variation in temperature, for if the animal be kept without food during
the day but be fed during the night, the temperature shows a rise to the
maximum, not at the usual time, in the evening, but in the morning.
This is denied by Carter,^ who observed an evening rise in the tempera-
ture of rabbits which had fasted three days.

Bernard ^ determined the temperature of the blood of the portal and
hepatic veins under different conditions as regards the nutrition of the
animals, and came to the conclusion that more heat was produced in the
liver during digestion. The following are some of his results : —





Blood of Portal
Vein.


Blood of Hepatic
Vein.


Blood of Right
Side of Heart.


Dog-


-After fasting for four days


.37° -8


38°-4


38°-8


M


Beginning of digestion .


39°-9


39°-5




>J


In full digestion .


39°-7


41°-3


39°-2



The effect of starvation upon the temperature of animals has been
studied chiefly by Chossat,^ and Bidder and Schmidt.'^ The first observer
made experiments on twelve pigeons, and he found that the rectal
temperature gradually fell until a short time before death ; during the
period of inanition the daily variation in temperature became more
marked, and towards the end of life a rapid fall in temperature occurred.
The results are shown in the table on p. 810.

On the day of death the temperature of the pigeon fell to 26°-2.
Similar experiments on turtle-doves, hens, rooks, rabbits, and guinea-
pigs gave the following temperatures:— 22^-9, 28°-2, 34°-3, 27°-0, and 23°-9
respectively on the day of death.

Bidder and Schmidt experimented upon a cat, and found that after

1 Davy, PUl. Trans., London, 1845, pt. 2, p. 319; iUd., 1850, p. 444; Damroscli,
Deutsche Klinilc, Berlin, 1853, S. 317 ; Jitrgensen, "Korperwarme des gesundeu Menschen,"
Leipzig, 1873, S. 21 ; Deutsches Arch. f. Min. Med., Leipzig, 1867, Bd. iii. S. 165 ;
Ringer and Stuart, Proc. Roij. Soc. London, 1877, vol. xxvi. p. 194 ; Ogle, St. George's
Hosp. Rep., London, 1866, vol. i. ; Crombie, Lidian Ann. Med. Sc, Calcutta, 1873, vol.
xvi. p. 581.

" Liebermeister, " Handbuch d. Path. u. Therap. des Fiebers," Leipzig, 1875, S. 123 ;
Wunderlich, "Medical Thermometry"; Siedamgrotzky, Deutsche Ztschr. f. Thiermed.,
Leipzig, 1875, Bd. i. S. 87.

^ Compt. rend. Soc. de bioL, Paris, 1884, p. 588.

* Journ. Nerv. and Ment. Dis., N.Y., 1890, vol. xvii. p. 785.

^ " Lecons sur la chaleur animale," Paris, 1876.

^"Recherches experimentales sur I'inanition," Paris, 1843, quoted from Gavarret,
" De la chaleur etc.," p. 394.

^ "Die Verdauungssafte und der Stoffwechsel," Leipzig, 1852, S. 322.



8io



ANIMAL HEAT.



355 hovirs' hnnger the rectal temperature fell from 39°-08 to 38°-4 ; after
369 hours, to 38°-l ; after 393, to 35°-5 ; after 415 hours, to 33^-7; and
again, after 426 hours, to 32°-4, when the animal died.





Rectal Tejiperatuee.


Daily Variation.


Midday.


Midnight.


Normal pigeons ....

Complete inanition, first period
,, ,, second ,,
,, ,, third ,,


42°-22
42° -1
41°-9
41° -4


41°-48
39°-8
38° -7
37°-3


0°-74
2° -3
3°-2

4°-l



In the case of the fasting man Tanner,^ no fall in temperature was
observed after thirty days' fast; the temperature of his mouth was
36°-9 (98°-4) on the twenty-fifth day, and 37°-l (98°-8) on the thirtieth
day. It is uncertain whether the fast was perfectly genuine, for Tanner
took a certain amount of Hquid. Noyes - recorded a temperature of
34°'4 (94°) in the case of a partly demented man, who had taken no food
for forty-five days, but it is to be noted that the condition was compli-
cated by paralysis of the lower limbs.

The influence of sleep.— The heat of the body falls during the night
and early morning, the time of inactivity and rest, but, according to
Barensprung ^ and Wunderlich,^ sleep in itself has no influence on the
temperature. Crombie,^ on the other hand, found that sleep during the
day caused a fall in temperature of about half a degree, but was
rapidly followed by a rise after awaking. Hunter ^ found that during
sleep the temperature fell about eight-tenths of a degree. The observa-
tions of Jiirgensen and Liebermeister '^ show that the temperature of a
man asleep is not lower than his temperature at a similar time of day
when he is awake and lying still. Inactivity causes a fall in temperature,
and sleep is a condition in which inactivity is most marked. Lieber-
meister ^ found that, by contracting the habit of sleeping each afternoon
for ten days, the mean temperature of his axilla fell to about 36'''5,
whereas it had previously been for that time of day 37°*3. Observations
by U. Mosso ^ also show that sleep during the daytime causes a fall in
the rectal temperature of man.

The influence of sex. — -Very little difference in temperature can
be observed in the two sexes.^^ Women may have a slightly higher
temperature, but the difference does not exceed half a degree ; their
temperature, however, appears to be more liable to variations. Davy ^^

^ Brit. Med. Journ., London, 1880, vol. ii. p. 171. ^ Ibid., p. 557.

^ Arch. f. Anat., Physiol, n. wissensch. Med., 1851, S. 163.

■* "Medical Thermometry," p. 109.

•' Loc. cit., p. 585.

'^ Phil. Trans., London, 1778, vol. l.Kviii. pt. 1, p. 20; "Works," Palmer's edition,
London, 1837, vol. iv. p. 144.

'' Liebermeister, " Handbuch d. Path. n. Therap. des Fiebers, " 1875, S. 87.

^ Hid., S. 92.

" Arch. ital. de hiol., Turin, 1887, tome viii. p. 177. See also this article, p. 802.
^^ Wunderlich, "Medical Thermometry."
" Med. Times and Gaz., London, 1864, vol. ii. p. 337.



INFLUENCE OF RACE.



8ii



concluded that the temperature of women and female animals was lower
than that of the male, but his observations were made upon only three
or four individuals. Thus he states that the temperature of three
healthy men varied between 37°'2 and 37°"5, that of three women
between 36°-5 and SG"'? ; in the case of three cocks and three hens the
results were 42°-4 and 42°1 respectively. Barensprung ^ found no
marked difference, the average temjDerature of eighteen women being
37°'25. As the result of seventy or eighty observations, Siedamgrotzky ^
gives the temperature of stallions, mares, and geldings as 37'''8, 38°-2,
and 38°"05 respectively ; the average temperature of a large number of
ducks was found by Martins^ to be 41°-96 for the male, and 42°-27 for
the female. Singleton * determined the rectal temperature of fifty dogs
and of fifty bitches ; the average for the former was 38°"9, for the latter
38°"7. The observations were made at similar times of the day, but
upon animals of different breeds.

The influence of race — The natives of tropical countries appear to
have a temperature slightly higher than that observed in the inhabitants
of mild or cold climates, but the difference is to be ascribed mainly to
the climate. Davy,^ from observations made upon natives in the
Cape of Good Hope, Isle of France, and Ceylon, found the temperature
to be about 0°-6 higher than the average in temperate climates ;
Crombie^ made fifty-two observations on Hindus, Mohammedans, and
East Indians in Bengal, and found that the average temperature from
10 A.M. to 10 P.M. was between 37°-2 and 37°-8, that from 10 p.m. to
10 A.M. between 36°-7 and 37°'2. Both of these observers also found
that the temperature of Europeans living in the same district was about
half a degree higher than the average in England. Joussef^ made
numerous observations on natives and Europeans living in tropical
climates, and came to the conclusion that the axillary temperature is
generally 0°-7 to 0°-8 higher than that observed in temperate climates.
The following figures show that climate, and not race, is the important
factor : —



Natives of Tropics.


Europeans.


Hindus ...


37°


•85


Officials at Chandernagore .


38°


•16


Cochin-Chinese .


37°


•60








Chinese


37°


•85








Negroes of Senegal


37°


•70


Sailors at Senegal


37°


•75


,, Congo


37°


•80


,, Antilles


37°


•70


,, Antilles .


37°


•80


Soldiers at ,, .


37°


•75



Similar results to the above were obtained by Maurel.^
The temperature of natives in South Africa was found by Livingstone ^
to be 36°^7, when the temperature of the air in the shade was 42°^2,



1 Arch. f. Anat., Physiol. %. luissensch. Med., 1851, S. 155.
- Deutsche Ztschr. f. Thiermed., Leipzig, 1875, Bd. i. S. 87.
^ Ellenberger, " Vergleichende Physiol, der Haussaugethiere, '
85.

■* See p. 790.

^ "Researches," London, 1839, vol. i. p. 169.
^ Indian Ann. Med. Sc, Calcutta, 1873, vol. xvi. p. 591.
'' Arch, de vied, nav., Paris, 1883, tome xl. pp. 123, 426.
* Bull. Soc. d'anthrop. de Paris, 1884, tome vii. p. 380.
^ "Travels and Researches in South Africa," 1857, p. 509.



1892, Bd. ii. Th. 2,



8i2 ANIMAL HEAT.

but his own temperature was 37°'8, owing probably to the difference in
clothes. Thomson ^ found the mean temperature of natives in Iceland
to be 37°'27, and Eijkman^ states that the average temperature of
Europeans living in Batavia is 37°"02, that of the Malays 36°-93.

The influence of menstruation and pregnancy.'' — Normal men-
struation and pregnancy in healthy women have no marked influence
upon the general temperature of the body. During labour the temper-
ature rises somewhat during the pains, but falls again between the pains.
Immediately after delivery a slight fall in temperature occurs.

Individual peculiarities in temperature. — Observations on men,
and especially on animals, show that the mean temperature of different
individuals is not the same, even when the conditions are as far as
possible equal.^ The mean temperature in the axilla of different men
may vary from 36^-5 (97°-7) to 37°-25 (99°-05). In animals even
greater differences are found.^

The influence of the temperature of the surroundings. — The
temperature of man and other warm-blooded animals is only slightly
influenced by the temperature of their surroundings. This fact is well
shown by the records of the temperature of men and animals in the
tropics and Arctic regions, where the extremes of the temperature of
the air occur, in the former +59°C., in the latter — 55°C. During a
voyage from England to Ceylon, Davy^ made observations upon the
temperatures of seven healthy men under 30 years of age ; he found
that the average temperature under the tongue was about 36°'9 (98°'4)
when the temperature of the air was 15°-6 (60°), and 37°"32 (99°-2)
when the air was 26° 4 (79°-5). From these and other observations,'^
he concluded that the temperature of man increases in passing
from a temperate into a warm climate, and that the inhabitants of
warm climates have a slightly higher temperature than those of
mild climates. Eeynaud and Blosville^ found the mean temperature
of eight men to be 37°'58 (100°), when under the torrid zone,
the temperature of the air varying from 26° to 30° (79*-86°), and
37°"11 (99°) in the temperate zone, with an external temperature
varying from 12° to 17° (53°-62°). The average temperature of the
mouth was found by Eattray^ to be 37°"25 (99°) in the tropics, with an
external temperature of 25°, as compared with 36°'8 (98°'3), the average
temperature in England during the summer heat (18°).

These and further observations, made by Brown-Sequard and others,^"

^ "Ueber Kraiikheiten und Kranklieitsverlialtnisse auf Island," Schleswig, 1855, S. 24.

" Firchow's Archiv, 1895, Bd. cxl. S. 125.

■^ Numerous references on this subject will be found in Wunderlich's "Medical Ther-
mometry," New. Syd. Soc. Translation, p. 105. See also Biirensprung, Arch. f. Anat.,
Physiol., ti. wissensch. Med., 1851, S. 157 ; Probyn Williams and Lennard Cutler, Lancet,
London, 1895, vol. i. p. 932 ; Giles, Brit. Med. Journ., London, 1894, vol. ii. p. 70. As
regards animals, see Hobday, Veterinary Rec., London, 1896, vol. viii. p. 488.

* This article; p. 789.

s This article, p. 790.

^ "Researches," London, 1839, vol. 1. p. 161.

■^ Phil. Trans., Loudon, 1850, p. 437.

^ "Animal Heat," article by Edwards in Todd's " Gyclopfedia," vol. ii. p. 659.

" Proc. Roy. Soc. London, 1870, vol. xviii. ]). 526.

^^ Brown-S(jquard, Journ. de laphysiol. de Vhomme, Paris, 1859, tome ii. p. 152 ; Gress-
well, Brit. Med. Journ., London, 1884, vol. ii. p. 164 ; Mantegazza, Presse mM. beige,
Bruxelles, 1863, tome xv. p. Ill ; Maurel, Bull. Soc. d'anthrop. de Paris, 1884, tome vii.
p. 371 ; Joussct, Arch, de m&l. nav., Paris, 1883, tome xl. p. 124 ; Pinkerton, Journ.
Anat. and Physiol., London, 1881, vol. xv. p. 118; Edoux and Souleyet, Compt. rend.
Acad. d. sc, Paris, 1838 tome vi. p. 456.



INFLUENCE OF EXTERNAL TEMPERATURE.



813



show that the effect of tropical heat is to raise the mean temperature
of the human body, but the increase is generally less than one degree.
Crombie^ found, as the result of 1288 observations upon himself, that
the temperature of the mouth was about 0°"23 higher in Bengal than the
average in England, but the difference was greater during the first few
weeks of residence in the hot climate.

On the other hand, some observers maintain that residence in a
tropical climate does not raise the temperature of the body ; thus
Boileau ^ states that the normal axillary temperature is between 36°"7
and 37°"2, Thornley^ and Furnell* that it is invariably the same as in
England, 36°-9.

JS'umerous careful observations recently made by I^euhauss ^ during a
voyage round the world, show the effect of external heat upon the daily
temperature, pulse, and discharge of urine. The following are some of the
results : —



Temperature
OF Air.


Six A.M.


Twelve
Noon.


Ten P.M.


Six P.M.


Remarks.


Temp
Zo
Min.

ir-5

Tropica
23°-9


ERATE

Max.
13°-6

L Zone.
26°-6


36°-6
55

36°-9
60


36°-9
55

37°-3
68


36°-8
56

37°-l
64


37°-l

62

37°-3

72


Teniperatm-e i.i^ jj(^^^ ^^ ^^

Temperature in"| ,^ „, ,

rectum I Mean of twenty-
Pulse J fi^<^ d^y^-



The influence of the different seasons of the year — ISTo marked
effect upon the heat of the body can be ascribed to the different seasons
of the year, apart from that due to variations in external temperature.
The numerous observations made by Davy ^ upon himself tend to show
that the temperature of the mouth is somewhat lower during the winter
months in England, and slightly higher during the summer ; a similar
series taken in the tropics, in Barbadoes, where the mean annual
temperature of the air is 26°'7, and the range throughout the year is
about 8°, shows no marked variation during the different seasons.

Joussef^ found that the cool season caused a fall of two- or three-
tenths of a degree in the average temperature of natives of the tropics.

From Bosanquet's ^ observations of the rectal temperature, it appears
that the highest sustained average temperature occurred in the winter
and early spring months. These determinations were made upon
himself four times a day for a period of three years.

A few observations have been made on the influence of winter and
summer upon the temperature of animals. Thus Edwards^ found in
the case of sparrows that the mean temperature rose progressively from

^ Indian Ann. Med. Sc, Calciitta, 1873, vol. xvi. p. 550.

" Lancet, London, 1878, vol. i. p. 413.

3 Ihid., 1878, vol. i. p. 554. ■* Ihid., 1878, vol. ii. p. 110.

5 Firchow's Archiv, 1893, Bd. cxxxiv. S. 365.

6 Phil. Trans., London, 1845, pt. 2, p. 319 ; and 1850, p. 437.
"^ Arch, de med. nav., Paris, 1883, tome xl. p. 124.

® Lancet, London, 1895, vol. i. p. 672.

^ "Animal Heat," in Todd's "Cyclopaedia," vol. ii, p. 659.



8 14 ANIMAL HEAT.

the depth of winter to the height of summer ; in the month of February
the mean temperature was 40°-8, in April 42°, and in July 43°-77 ; from
this time the temperature began to decline. It was also found that, in
winter, birds could more readily resist the action of extreme cold than
in summer.

Davyi observed the temperature of sheep during summer and
winter, and his results, although they are not sufficiently consistent
for positive conclusions, seem to show that the temperature of the body
is a little higher in the warm weather than in the cold.

The influence of extreme heat and cold. — The experience of
the inhabitants of tropical climates shows that it is possible to live even
in an atmosphere the temperature of which at times exceeds that of the
body, and that the body is able, by means of the cooling effect of the
evaporation of sweat, to prevent its temperature rising a degree above
the normal.

Lining,^ in 1738, found that the temperature of his axilla was 36°-l,
and that of his mouth 36"-7, when the heat in the sun's rays was 51°-1,
in the shade 36°-7, on a hot summer's day in South Carolina. Ellis,^ in
1758, observed that the temperature of his body was not above 36°"1
when he was living in Georgia, and the temperature of the air was
40° -6. Experiments on men and on lower animals have shown that much
greater heat can be borne for short periods. Blagden and Fordyce *
observed their own temperatures after remaining in heated rooms, and
found that the effect varied according to the amount of moisture



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