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Torpor and metabolism in hummingbirds

Article  in  Comparative Biochemistry and Physiology Part A Physiology · December 1982

DOI: 10.1016/0300-9629(82)90275-4

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Cony. Biochem. Physiol. Vol. 7iA. No. 4, pp. 679 to 689. 1982 0300-9629’S?/ I ?0679- I 1$03.00/O
Printed in Great Britain. 0 1982 Prrgamon PressLtd

TORPOR AND METABOLISM IN HUMMINGBIRDS

KKISTINE KRUGER. R. PRINZINGER* and K.-L. SCHUCHMANN

Institut fiir Biologie III, Lehrstuhl Zoophysiologie. Auf der Morgenstelle 28.
7400 Tiibingen 1, W. Germany

(Rrcrirrd 8 Marc,h 1982)

Abstract- -1. Body temperature and diurnal cycle of energy metabolism at different ambient tempera-
tures (+2 to +25 C) were tested for 18 different hummingbird species from different biotopes and of
different body-masses (2.7-17.5 g).
2. The level of resting metabolism (night time) reaches 3(&70”,, of activity metabolism values (day
time). The mean is almost exactly 50”,.
3. The mean metabolism-weight regression line of the night time values follows the equation
M = 0.67 x l+‘“.73 (bf = energy metabolism in kJ/hr, W = body weight in g). That of the day time
values is hf = 0.83 x W”.53.
4. Resting metabolic rate of the hummingbirds is considerably higher than the theoretically expected
value for nonpasserine birds, even when torpor values are taken into account.
5. All species tested show torpor during night time, independent of ambient temperature. feeding
situation or the environment (biotope) of the species.
6. In comparison to the resting metabolic rate. metabolism during torpor decreases by 6&90”,, to a
relatively constant level. This level is not (linearly) temperature-dependent (~1 = 0.008. r = 0.22). never-
theless many species show a minimum at cu 15-20~~‘.
7. During torpor. body temperature normally reaches the level of the ambient temperature but never
falls below 18 -20 C. The normal values during the daytime range from 38 to 40 C and during night time
from 35 to 37 C.

INTRODUCTION only in a state of emergency or is torpor used for

1.
conserving energy as often as possible, i.e., regularly
Torpor is a numbness-like state with a large reduction
each night. not controlled by a physiological de-
of body temperature and metabolism. Entrance into,
ficiency but (perhaps) by a endogenous rhythm’! To
maintenance of and arousal from torpor are active clarify this question we investigated the energy metab-
and regulated processes. Torpor has no ill effect on olism of hummingbirds of different body sizes from
the animal (Swan, 1974). Torpor has been described in different climatic zones (tropics. desert, temperate
many animal groups. Raths & Kulzer (1976) list zone).
among the Mammalia bats, tenrecs, prosimians.
various rodents and shrews. In birds torpor is known
from species of the orders Caprimulgiformes, Apodi- hlATERlALS AND ,METHODS
formes. Trochiliformes and Coliiformes. among
Seventeen different hummingbird species and one hybrid
others.
were used in this study. All birds were wild-caught with the
Hummingbirds, which include the smallest bird exception of the hybrid. and underwent a complete moult
species, inhabit nearly all climatic and altitude zones in captivity. Their exact age is unknown but is estimated to
of America and are one of the most species-numerous be between 3 and 5 years. The hummingbirds were housed
groups among the nonpasserines. Corresponding to at a constant temperature of 20’ C with a 12: 12 hr photo-
their low body size and their mode of life-they feed period (lights on 07:3@19.30 hours). Some birds were kept
mainly on nectar during flight, hovering from blos- individually in small, cloth cages (0.5 x 0.5 ml and others
som to blossom-they show a high energy demand were housed. several birds together. in an aviaiy (3 x 5 m).
but only minimal storage capacity. The ability to They were fed with Drosophi/a and a lo”,, water-solution of
saccharose. fructose and glucose as well as amino acids and
reduce energy consumption drastically during the
aromatic substances. Water for drinking and bathing was
night may be the only possibility for these birds to
available ud lihitum.
survive in, for example, the Andes at a height of Body temperature was measured (under the plumage) in
4000 m and a mean ambient temperature of +_5 C. the axilla of the birds with a digital thermometer (Testo-
For these reasons, investigations of torpor are of therm KG) (accuracy about +O.Z’C). To prevent a warm-
special interest in humming-birds. ing of the birds, and by this a falsification of the results, the
From many previous works on hummingbirds completely motionless hummingbirds (during torpor) were
came the assumption that these birds only undergo layed in a cloth while measurements were taken and not
torpor in the course of acute energy deficiency and touched directly by hand.
During the measurements of metabolism, the birds were
low ambient temperatures. From the considerations
isolated in cylindrical Plexiglas respirometer chambers
stated above, we held another question to be more
(20 x 40cm) containing a perch and a feed tank (Fig. 1).
reasonable: Do hummingbirds practice this ability The respirometer chambers were placed in a tempersture
controlled cabinet, regulated with an accuracy of + 1 ‘C.
* To whom requests for offprints should be sent The air flow through the chambers was adjusted to the

679
 680 KR~ST~NEKRUGER et LI/.

When entering torpor, energy consumption de-

creased once more by 6&90”,, to a very steady level.
Duration and level of the torpor phase are different in
the individual species.
At the beginning of the light phase energy con-
sumption increased very quickly. After a short time
(5-20min.) the hummingbirds had reached their nor-
mal day time level. The rate of metabolic increase
during this awakening phase may amount to over
loo”,, per 5 min.

The respiratory quotient (RQ) during day time had

a constant value of 1 (which means that mainly sugar
was metabolized). After the beginning of the dark
Fig. I. Schematic vie\\ of the plexiglas rcspirometer
phase (end of food intake) RQ remained at this value
chamber used in this study. Shown is a vrr> small hum-
for about 10min and then decreased within about
40 min to a lower mean value of 0.8 (fat metabolism).
which then remained steady during the entire night.
In the morning after food intake, RQ again increased
constant value of 20 l:hr. Oxygen consumption and carbon
dioxide production were measured in an &en flow system to 1 within half an hour after the beginning of the
with a Hartmann & Braun MAGNOS 2 T and URAS 2 T light phase.
paramagnetic 0,.analyzer. and an infrared COz-analyzer. Body mass of the hummingbirds also underwent a
All gas volumes were reduced to standard pressure and distinct diurnal rhythm. On average the birds weighed
temperature conditions (accuracy of all data: _t I”,, of the (dependent on the weight class) about O.lLO.3 g in the
measuring range). evening more than in the morning. This amount was
During the measurements of metabolism the photo cycle stored during the day time as an energy reserve for
was adjusted to provide the birds with the normal diurnal
the night.
cycle. For this reason they were also fed at the same times
as under normal housing conditions (9.00 hr and 16.00 hr):
food was given trd lib. The hummingbirds normally became
quickly familiar with the testing conditions, Usually within
one day they sat calmly on the perch and showed a normal Duriny cccti&y. The mean values of activity metab-
food intake. As they could fly in the respirometer chambers olism increased with falling ambient temperature. The
(hovering-flight). measurements of awake ( =active) birds changes of energy metabolism were different from
do not represent basic energy metabolism. species to species (see Table 1). The mean of the corre-
To test for a possible effect of a circadian rhythm of sponding regression line during the day time is
metabolism on the results, each ambient temperature was
M = -6.67 T, + 515 ( + 151). The metabolic adap-
monitored continuously for at least 24 hr. Each experiment
tation to changing ambient temperature was ac-
lasted 46 days without interruption. During the duration
of the experiment the birds remained in the respiratory complished very quickly. often within a few minutes.
chambers. and were only removed for taking measure- The increasing rate was in this case largely propor-
ments of body temperature and body mass. The results of tional to the temperature fluctuations.
the first day measurements were not evaluated, as it was Dwingg rhr night. The dependence of energy con-
observed that the birds were very nervous at the beginning sumption on the ambient temperature was distinctly
of the testing procedure and that they needed some time to less during the night than during the day. If the
habituate to the changed housing conditions. regression line is calculated without taking into
The measuring apparatus, with six changing channels.
consideration the torpor values. it lies at bl =
allowed the tebting of 5 individuala at the same time. The
- 3.85 T, + 252 () 180). The average temperature
last channel served as a zero-point control.
dependence of night time metabolism including tor-
nor values is remarkablv small (m = -0.84). In many
RESL’LTS hummingbird species the slope of the regression line
(ru) even inverted (see Table 1) during the night. Al-
together there existed greater interspecific differences
in night time metabolism than in day time metab-
As expected, there were greater fluctuations in olism.
energy metabolism during the day time. caused by the Durirly! torpor. A temperature dependence of energy
fact that the hummingbirds could fly in the respirat- metabolism during torpor could not be confidently
ory chamber. Nevertheless the values show a temper- confirmed statistically. This means that the level of
aLure-dependent characteristic mean (Table 1). With torpor metabolism is not linearly temperature-depen-
the beginning of darkness, energy consumption of all dent. The mean regression line follows the equation
hummingbirds decreased very quickly to an essen- M = 0.008T,+ 48 (i 36). Nevertheless. for many
tially lower level, remaining relatively constant during species there seemed to be a minimum of energy con-
the entire night provided that the bird did not sumption at about 15-20 C (the range of deepest ob-
undergo torpor. The values of night time metabolism served body temperature). There was also no statisti-
range between 30-70”” of the mean day time values cally significant correlation to be found between
with an average of almost exactly 50”,,. ambient temperature and the occurrence of torpor
 Torpor and metabolism in hummingbirds

IOOO-
O.UNOERUOOOIf 1000 R.MULSRNT

;
:
coo-

600.
-
800
1
:
2
~ 400.

1’5 21 3 9 15

1000
1 U.BENJRMINI

800~ ,“,

oJ

15 21 3 9 I5

10007
TSCIIULbS

I5 2’1 i i 1’5
1000,
CH.OENONE

3 iI 1'5 I5 21 3 9 15

""li Ui:iEIPNDRI '2' 1000

1 HYBRID

800

z i

200
1

y-1
15 21 3

Fig. 2.
682 Fig. 2-cont.
1000 F.NELLiVORR ;

---
1s 2, ; 1; 21 3 9 ‘5.
R.CUPPIPENNIS
1000
O.ESTELLR I

800

CT ;
: 600 : 600
_
:
7 1

z
400 400
z

“00 200

0 0

'oool O.ESTELLR 2

-
;
:
800

600. I
1
0
:
800

600
I
;
7 i

IS 21 3 9 is
1000
E.JUGULRRIS

0
1
800
I 800
I

: 600

400
r

200

1000

tloo-

;
: 630.
;
7

400.
x

200

Fig. 2. Diurnal cycles of energy metabolism at dmerent amblent temperatures. measured contmuously
on consecutive days. The black and white bars represent the dark light cycle. 25 (-. ‘0 c.
15 c. ----- 10 c. 5 c.
 Torpor and metabolism in hun~nlln~birds 683

fr = 0.22) or the length of the torpor phase (I’ = 0.1) The mean metabolism-weight regression line of
(see Table 1). In the same way as during the resting the day time values follows the equation M =
time (night), metabolism adjusted during torpor to a 0.83 x W”.5h (!\I = metabolism in kJ)hr and L+’=
very steady level. This adjustment did not usually body weight in g). That of the night time values is
begin until about 13.OCOI.00 hr. Before the beginning M = 0.67 x I+” ‘3. During the day time the heavier
of the light phase. torpor normally first passed over to hllmmingbirds were calmer than the smaller species.
normal sleep, but several birds also awoke directly This results in a lower regression exponent for the day
from torpor shortly before day-break. time values.

During torpor the minima1 body temperature of all
tested hummingbirds lay between I8-70°C and during
night time (normal sleep) between 3S-37°C (see
Table 3). Values beyond 18’C did not occur. Table 3
shows only those values where comparable day and
night time data were available.
When disturbed during torpor. the hummingbirds
showed a number of typical displays occurring at cer-
tain phases of torpor. In the deepest phase they
showed no visible reaction at all. However. a short
time after a waking-stimulation (for example a touch)
they strcched their wings abruptly and gave shrill
cries (like the syueak of a mouse) that are never heard
from waking birds. At the same time we observed
clamp-reflexes of the birds’ feet. When placed on :I
perch they grabbed it and remained sitting in an
upright position. During the deep phase of torpor and
during the awakening period the claws were closed
and no clamp-refIex occurred. During undisturbed
awakening from torpor there also occurred several
distinctively different phases of display in which
breathing frequency, body stance and movement of
the hummingbirds changed characteristically from
minute to minute until the bird was awake.
Both during day and night. and at all temperatures
tested. the metabolism of all the hummingbirds was
considerably higher than the theoretically expected
value for nonpasserine birds. This fact could also be
~~n~rrned for periods during which torpor occurred.
DISCUSSION
From the in~~estigations of other authors, the fol-
lowing equations were obtained (defined as above):
M = 0.1 SS x I+“.” for passerines (Dawson & Hud-
son. 1970); M = 0.081 x Cyo.” for nonpasserines
(Aschoff & Pohl. 1970). It is commonly assumed that
the basal metabolic rate of passerine birds is higher
than that of nonpasserines. Prinzinger & HCnssler
(1980)investigated small representatives of nonpasser-
ine birds and found no difference in the metabolism-
weight relationship in comparison to passerine birds
of the same weight. The calculated equation (M =
0.138 x 14’O-lh) lies between those given above. but
closer to that of the passerines. The slope of all three
lines is about the same. The values for our much
smaller hummingbirds are significantly higher although
the slope is again the same. The correlation between
basal metabolic rate and body weight al night time
with an ambient temperature of 2S’C (calculated in-
cluding torpor values) follows the equation hl =
0.2 i 2 x f4’“-T’. Hummingbirds therefore. though
belonging to the nonpasserine birds. have ;I higher
basal metabolic rate than all other birds species inves-
tigated thus far. The values are equal to those of
mammals of the same weight. for example shrews
(Fans & Sicart. 19X0) (Table 5). This means that the
level of n?et~lbolism does not depend on the system-
atic order ;I bird belongs to inonpasserincs~~ passer-
ines) but rather on its body mass. The corresponding
level of metabolic rate of hummingbirds and Croci-
durinae indicates that this rule is valid over ;I very
Nide range; while in the same lveight class. humming-
birds and ~hitet[~~~thed shrews belong to {cry difkr-

Table 1. Mean energ! tnctabolism (J g,hr) at direrent ambient temperatures during &I! (Iqt number).
during night (2nd number1 and during torpor (3rd numberl

715 604 507 J9.1 \ = -15.3 r: -i-S47 I -0.96)

37x 791 234 203 ; = -I 1.6 X +3x0 I - 0.90)
60 36 44 * ?’ =- 1.6 x +71 t-0.65)
6X9 636 - 419 479 42x y= -13.6 x +734 (-0.86)
261 311 778 203 180 y = -5.4 x +318 (-0.84)
101 76 15 26 26 y= -4.0 x fill ( - 0.90)
475 450 - 416 477 375 y= -3.5 x f491 ( - 0.64)
217 242 - 205 231 I44 y= -3.2 x +X5 i-0.65)
85 75 51 26 63 y = -1.9 x +8x t -0.64)
- 456 - 5’0 313 300 y= -13.5 x +624 I-0.81)
- 308 - 228 165 140 y= -11.3 x +408 t -0.98)
.- * 25 68 4 = -2.1 x +99 I - 0.50)
532 9x
598 - 6’6 - 471 Y= -3.3 x -+605 I - 0.44)
249 390 .- ‘97 - 235 ; = -2.9 x +33-! i-0.35)
* 61 - * * *
- 615 ._ 533 535 478 y = -8.2 x -+638 ( - 0.94)
- 349 228 ‘28 170 y = -10.7 x l-431 ( - 0.92)
52 * * * *
684 KRETINE KRUGER et al.

?‘;thic I -- cont.

Species, sex. mesn body mass tg)

35x 48? 446 y = -C.l x +547

_ 194 244 751 y = -1.3 x +XS
35 30 36 y=O.? x +28
596 358 ‘14 y = -20.9 x +814
260 1.51 120 y = -9.8 X +377
* I3 * *
_ SlZ 5’79 513 y = 4.8 x +415
_ 215 2% 254 y = 3.5 X i 171
_ 29 3s 102 ]i=CW x 1-62
_ 421 4’10 431 y = -2.8 x +a4
_ 2417 2% 1’1 )’ = -2.0 X +1x0
47 -il 97 y = 1.2 x 1-311

All)
185
29
_ 398
225
_ *
370 306
17S 141
I‘ 10
367 310
726 188
* *
45x 3X6 314
196 344 ‘03
* * *
_ 3% 411
228 224
_ 52 35
_ 439
166
9
298
173
*
7132
If>,?
I6
_,2h 767
* 157
37 15
396 231
2% I SO
35 X
240
13%
*

Mean day time regression

Mean night time regression
(without torpor valuesf ?;” -3.x5x +‘Sl(i_lRO)
Mean torpor regression y= 0.008 x +3Y (t?h)

ent taxa. These results atso show fh;tt values for the
relation h = LI x M/h amnor be caiculated exactly
over a wide range of body weights. and values cannot
be extrapolated correctly beyond the actual measur-
ing range as the position of the line is shifted in the
direction of zero by great changes in body weight and
the factor LIincreases when bodk mass decreases, It is
of interest that even Hhen torpor metaboiic values are
included in the calcukttion. the metabolic rate of
htzmmirtgbirds is much higher than that of heavier
birds.
These tiny birds cannot compensate for their higher
metabolic demands by using their normal ability to
save energy. This means that they have to use their
ability to fall ir~io torpor as often as possible. even at
it high ambient tompertlture and with sufEcient food
 Torpor and metabolism in hummingbirds 685

IO00 1000
0.UNOERH0001, R.RLEYRNOR, 2

.
01

0 IO 20 30

1000
CH.MELLISUGUS

800 800

; ;
; 600 ; 600

; ;
7
7
_

z
4oc r
400

200
-- --ii 200

-----. 0
.
c

0 IO 20 30

1
1000 1000 ".BENJRMlNI

800 800

u 0
; 600 : 600
; ;
2 3
400 400
z r

200

0 I'0 i0 40 0 IO 20 70

1000 IO00
0.CRISTRTUS 2 T.SCITULUS

BOO 800

z
0 ;
: 600 : 600

; ;
1
_’ _

z 400 r 400

200 200

0 0

-- I
b I’0 20 li0 0 IO 20 30

1000 ,000
R.RLEXRNOR, I CH.OENONE

801 800

CT ;
: 6OC : 600
; ;
7 7
r 400 400
z

200
--N 200

0 0

Fig. 3.
 Fig. 3-~011~.
I DO0

IO 20 30
0

lOOal R.NIGRICOLLIS ““1 E.JUGULRRIS

800
- i

--- r----- , 30
7-----

30 10 20
0 :3 20 0
T DC
T LOCI

Fig. 3. Energy metabolism (M) as a function of ambient temperature (T,) of different hummingblrd
species during day (-). during night (N) and during torpor (0) (see also Table 1).
 Torpor and metabolism in hummingbirds 6X7

Table 2. Occurrence of torpor at different ambient temperatures

Ambient temperature (‘C) 5 IO I5 20 ‘5

Number of birds in torpor II 15 13 14 14

No torpor observed 6 4 7 5 6
Percentage of birds in torpor 65 79 65 74 70

Table 3. Mean body temperatures at different physiological stages

Mean body temperature ( ‘C)

Species day time night time torpor Ambient temperature

L. clemenciae 39.6 35.7 19.6 15

0. esfella 40.8 35.7 20.3 15
U. benjamini 39.0 28.0* IO
0. c,risratus 40.0 20.8 15
Ch. ocnonc 39.0 35.0 18.0 I5

* Bird not in a deep torpor.

supply. Torpor occurred in all hummingbird species use of torpor. In contrast to mammals (e.g. shrews)
investigated. No correlation between occurrence of hummingbirds can consume food only during the day
torpor and body mass was found. The lowering of time, thus they have greater difficulty maintaining
metabolic rate was the same in all weight classes their energy balance during the night.
(about 80-909k of the active metabolic rate). Inhabi- According to calculations by Berger & Johanson
tants of deserts fell into torpor as often as those of (1980) a hummingbird can save up to 85”,, of energy
tropical, temperate or cold zones. Length of torpor. as expenditure in torpor. compared to the homoiother-
well as occurrence, is not correlated with the biotope mic stage, a fact which is confirmed by our own
of the different species. Torpor seems to be a common results. The maximum amount of this energy saving is
characteristic of the entire family of Trochilidae and is at relatively high ambient temperatures. of 15--20 C.
not limited only to those living in extreme biotopes. At these temperatures torpor is. therefore, very effec-
Probably the evolution of this ability was mainly tive for the birds. The minimum energy consumption
dependent upon the low body mass of these smallest lies in this range, because the birds can decrease their
representatives of the birds whose body temperature. body temperature passively to correspond to this level
of 4&42’C, lies above even that of the mammals. of ambient temperature; below this level they again
Even in tropical regions with favourable living con- need more energy to regulate their body temperature
ditions the small hummingbirds can only achieve a to the minimum value of about 18-20 C. which then
positive energy balance sufficient for survival with the are above ambient temperature.

Table 4. Metabolism-weight relationship at different ambient temperatures during da\

and night (mean values of I7 different species) (,$I in kJ hr. I$’ (body mass) in gl

Ambient Weight-metabolism regression (coefficient r)

temperature day time night time

5 ‘C n-r = 0.926 x W0 ‘” (0.88) iLf = 0.273 x U.” ‘- (0.69)

IO c M = 0.975 x u’“.SD (0.81 I hf = 0.23 I x 12’”T6(0.67)
I5 c M = 0.866 x W” 54 (0.84) I!{ = 0.2X6 x 12’” ” (0.64)
20 c M = 0.726 x W” ‘.’ (0.87) nr = 0.221 x U’” I(1 (0.X5)
25 c R,f = 0.708 x w” SJ (0.84) R1 = 0.21’ x LZ’[’-6 (0.79)

Mean M = 0.803 x W”.sb .If = 0.670 x 14’” -’

Table 5. Energy consumption of shrews (S) and hummingbirds (H) which have
similar size and which were measured under nearly identical conditions (during
resting period at an ambient temperature of 25°C) [shrew data from Fons 81
Sicart (1980)]

Mean body mass Mean resting metabolism

Species (g) (J/g x hr)

Suncus etruscus(S) 2.5 320

Ocreatus underwoodii (H) 2.7 300
Crocidura russula (S) 8-10 150
Lampornis clemenciae (H) 8.3 120
688 KRISTINE KRUGER et al.

During torpor the hummingbirds do not abandon temperature in primarily homoiothermic animals that
temperature regulation but the whole metabolic rate occasionally ~mo~sebirds~ or continuously (humming-
is regulated to another (lower) level. It is of interest in birds, shrews) live near the subsistence leve1. These
this regulation that there is no adjustment of new. animals can lower their body temperature secondar-
periodically changing nominal levels, but rather an ily, thereby saving a great amount of energy without
adjustment of a single nominal value of 18-2YC. Up having to give up their ability to regulate tempera-
to this point no regulation takes place. When this ture. On the contrary. their ability to regulate tem-
point is reached regulation begins automatically and perature. expanded as metabolism, can be regulated
so body temperature does nat fall below this level. on several different levels, depending on the actual
During torpor, metabolic rate is also reduced to a demands. The theory of torpor being a secondary
very low level which is obviously independent of character is confirmed by the fact that newborn
ambient parameters (e.g. ambient temperature). It shrews do not develop the ability to fall into torpor
seems to represent a minimum levef for maintaining until they have developed hom~iother~~i~ reactions at
the vital functions of the hummingbirds. Prinzinger et the age of about one week (Nagel, 1977).
~rl. ( 1981) investigated the red-backed mousebird
(Co[ius L.U.~~LIIZOIK~) and found that during times of REFERENCES
food scarcity these birds decrease their body tempera-
ASCHOFF J. & POHL K-L(1970) Der Ruheumsatz von Viigeln
ture and metabolic rate continuously. When falling als Funktian der Tageszeit und der KijrpergrBBe. J. Orn.
short of a critical body mass of about 5Og they 111. 38-47.
undergo a nightly torpor by reducing their metabolic BERGER M. & JOHANSEN K. (1980) Die Stadien der Kg&es-
rate abruptly to &out 5”. of normal values. They also tarre bei Kol~br~~-~Anpassung van &mung und Kreis-
maintain a minimal critical body temperature of lauf. !‘erlz. dt. rnoi. Ges. 19x0. 307,
about 18~‘C. Unlike hummingbirds, mousebirds do C’AKPEE~TERF. L. (1974) Torpor in an Andean humming-
not faI1 into torpor when food supply is sufficient. bird-~ Its ecological significance. Scipn(:r 183, 545-547.
CAKPENTCR F. L. (1976) Ecology and evolution in an
They are essentially heavier (55-70 g) than humming-
Andean hummingbird, Unit>. Cal$ Prrhls 2001. 106,
birds and so have a more favorable relation of body
I-75.
mass to energy expenditure. DAWSON W. R. & HUDSON .I. W. (1970) Birds. In Cotnpuru-
In small mammals torpor occurs in nearly the same tier Physiology uf‘ Thermoreyulutir>n, Vol. 1 (Edited by
way as in birds. It seems to be a common pattern that WHITTOW G. c.). pp. 223-310. Academic Press, New
is only slightly changed to fit the special demands of York.
different groups of animals. In bats (e.g. Il$j*oris Fms R. 6t SICART R. (1980f Contributions i la connais-
m_~oris. Heldmaier. 1970) a periodic occurrence of tor- sance du m&abolisme Cnergetique chez deux Crociduri-
por could aiso be found even when they were main- nae, S*mcus etruscw (Savi. I882) et Crocidura russula
(Hermann, 1780) (Insectivora, Soricidae). Mammalia 40,
tained under constant ambient conditions (DD). He
299-311.
shows that this is not only a simple reduction of
FREY H. & VOGEL P. (1979) Etude de la torpeut chez
metabolism but a number of complicated regulatary Suncus etruscu~ (Savi, 1822) (Soricidae, Lnsectivora) en
processes in which body temperature and metabolism captivito. Raw suissa Zocrl. 86, 23-36.
can be regulated to different levels. In bats, as in HA~NSWORTH F. R. & WOLF t. L. (1970) Regulation of
1~L~mmingbirds. the same parallel changes of O2 con- oxygen consumption and body temperature during tor-
sumption, decrease of RQ to 0.75 and decrease of por in a h~Immingbird~ E. ~~~~~~~~~~j,~. S&ncr 168,
body temperature to wa’tues of 2@ 22 C can be ob- 348 -369.
served. The same minimum of body temperature dur-
ing torpor can also be observed in white-toothed
shrews whose body rnzs (3-2Og) corresponds to that HELDMAIER G. (1970) Variations of body temperature: and
of hummingbirds. Nngel (1977) induced torpor in metabolism during entrance into cold lethargy in the bat
Sunc~~,s c’trusr~rs by reducing food supply. and also My&is 0lyutis. Bijdr. Die&. 33. 1455.
found that body temperature is not reduced below a K&YSEK C. & HIX:SNHK A. (1964) Le rhythme mycthemeral
lrvcl of’ 18 30 C. Frey & Vogel (1979) found that S. de la depense d’energiquc. Etude de physiolopie com-
r~uzc~~rs falls into torpor when food supply is reduced. pat+“. 3. Pki_sitti.. PrU+a 19, 3-t 16.
as well as spontaneously in a diurnal rhythm occur- KLEIBER M. (1931 ;32f Body size and metabolism. ~~~~~~~~f~~~r
ring mostlv at night between 1.00 hr and 6.00 hr. The 8, 315.
KING J. R. Csr FARNEK D. S. (196t) Energy metabolism.
small Crodidurinae. S. c*t~trscus and C~*r)cidl~r~r ~trssultr,
thrrmorcgulation and body temperature. In Bioloc/y trnd
have a metabolic ratt‘ that lies high ahox the theor- Cmqwuriw Physinltry~~q/Birds. Vol. 2 (Edited by MAK-
etically expected values. The values for energy expen- SHAI.L A. J.). pp~ 215-188. Academic Press, New York.
diture (Fans & Sicart, 1980) correspond remarkably LASIEWSKY R. c. & LAWSON W. R. (1967) A re-examina-
well to those of hummingbirds with the same body tion of the relation between standard metabolism rate
size. and body mass in birds. Cot&w 69, 13 13.
Torpor as a means of energy saving is used by NACZL A. I 1975) Torpor in European ~iii~e-to~?thed
many diRerent groups of animals in times of food shrews. EupmWm% 33, 1455. f456.
shortage. as well as periodically in the resting phase of PKIXZINGEK R. (1936)Tcmperatur- und StaKwechs&zgul:t-
tinn der Dohle Corrus rrrorrrduiu, Rabenkrghe Cr)rclrs
the diurnal cycle. Body temperature, metabolism and
(‘clro~l~und Elster Picu pica. Corvidae. .4/t:. OUI. Ges.
behaviour are changed in a very similar way accord- BuyerI? 15, l--47.
ing to a common pattern with only slight variations. PKI~UZINGEKR. & HGNSSLER I.(1980) Metabolism weight
Therefore. we can agree with Raths & Kulzcr (1976) relationship in some small nonpasserine birds. Esp~+Gn-
that torpor seems to he a polyphyleticnlly evolved tit1 36, 1299-l 300.
ability. It allo~vs a lowering of metabolism and body
 Torpor and metabolism in hummingbirds
Body temperature and metabolism in the Redbacked
Mousebird (C&us casttrnorusf during fasting and torpor.
Camp. Bj~)~h~~~. ~~ys~o~. 69A, 689-692.
RATHS P. & KULZER E. (1976) PhysioIogy of hibernation
and related lethargic stages in mammals and birds. Bon-
ncr Zool. Monogr. 9.
SWAN H. (1974) Thernzoregultrtmn und Biomrryrfic’s. Else-
vier. Amsterdam.
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FREYH. (1979) Influence de ta temperature et de la nour-
riture disponabi~ sur la torpeur chez la ~1Lls~~reigne
musette (Crucidura rzrssufa) en captivite. Bttl!. Sot. vcfud.
Sci. Nat. 356. 325-332.
WITHERS W. (1977) Respiration, metabolism and heat
exchange of euthermic and torpid pooruills and hum-
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