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The Energy Cost of Song in The Canary, Serinus Canaria
The Energy Cost of Song in The Canary, Serinus Canaria
doi:10.1006/anbe.2003.2250
Although sound production requires energy, it has been unclear how much singing increases metabolic
rate in passerine birds. We measured the rate of oxygen consumption of two breeds of canary that sang
inside a respirometry chamber. Metabolic rate increased with the proportion of time that birds spent
singing. Average metabolic rate during singing at 15–20C was 1.05–1.07 times that of standing quietly in
the same temperature range or 2.2–2.6 times basal metabolic rate (BMR). Whether an increase in
metabolic rate during song of this order would represent a fitness cost to free-living passerine birds would
depend upon the circumstances. Singing rather than perching during the day would raise metabolic rate
only slightly. Singing at night or at dawn, instead of sleeping with a metabolic rate closer to BMR, would
cause a greater increase in metabolism. Birdsong could act as a condition-dependent signal, since birds
that are easily able to achieve energy balance could afford the cost of singing, but those close to their
energy limits might not.
2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
Birdsong is a conspicuous example of a display that 2001). Field observations of birds imply that singing has
can function both to deter rivals and to attract mates an energy cost. Birds sing more when in good body
(Catchpole & Slater 1995). The possible costs of birdsong condition or following supplementary feeding and less
are not well understood. Most models of animal signal- after cold nights or following adverse experimental treat-
ling assume that displays should be costly if they are to ments (Lambrechts 1996). However, these data do not
carry honest information about the quality of the signal- show whether singing itself is energetically costly or
ler (Zahavi 1975; Clutton-Brock & Albon 1979; Grafen whether time that could have been spent singing, at
1990a, b; Godfray 1991). Hypotheses on how animal however low a rate of energy expenditure, was better
displays may have evolved assume explicitly or implicitly spent foraging during periods of energy shortage.
that signals should be costly (Fisher 1930; Zahavi 1975; Measurements of the rate of oxygen consumption of
Grafen 1990a, b). The costliness of signals is also an captive animals show that sound production can cause
important assumption of stochastic dynamic program- large factorial increases in metabolism for insects,
ming models of animal signalling (e.g. Hutchinson et al. anurans and bats (Ryan 1988; Speakman et al. 1989;
1993). Prestwich 1994; Kotiaho et al. 1998). However, one can-
Song production obviously requires energy. Muscles not infer from these data how energetically costly singing
associated with control of respiration, the syrinx, the should be for passerine birds, because resting metabolic
upper vocal tract and movements of the bill are all used rates and sound production mechanisms differ between
during singing (Goller & Larsen 1997; Suthers et al. 1999; taxa. Singing inside respirometry chambers appears to be
Larsen & Goller 2002). Singing could also be costly in energetically costly in Carolina wrens, Thryothorus ludovi-
other ways: time spent singing is not available for other cianus (Eberhardt 1994), but not in zebra finches, Taen-
activities, singing might attract predators, and song learn- iopygia guttata, Waterslager canaries, Serinus canaria, or
ing and production involves specialized areas of the brain European starlings, Sturnus vulgaris (Oberweger & Goller
that could be costly to develop or maintain (Gil & Gahr 2001; Franz & Goller 2003). Singing by free-living com-
mon nightingales, Luscinia megarhynchos, increases their
Correspondence: S. Ward, School of Biology, Bute Medical Buildings, overnight rates of mass loss, implying that song has an
University of St Andrews, St Andrews, Fife KY16 9TS, U.K. (email:
energy cost in this species (Thomas 2002). In contrast,
sw29@st-andrews.ac.uk). J. R. Speakman is at the Aberdeen Centre for
Energy Regulation and Obesity, School of Biological Sciences, Zoology
begging by nestling birds is thought to be energetically
Building, Tillydrone Avenue, Aberdeen B24 2TZ, U.K. and Division of cheap (Chappell & Bachman 2002) as is crowing by
Appetite and Energy Balance, Rowett Research Institute, Greenburn cockerels, Gallus gallus domesticus, and junglefowl, G.
Road, Bucksburn, Aberdeen, AB21 9SB, U.K. gallus spadiceus (Chappell et al. 1995; Horn et al. 1995).
893
0003–3472/03/$30.00/0 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
894 ANIMAL BEHAVIOUR, 66, 5
Conclusions that bird vocalization is energetically singly (to prevent aggression between birds) in cages
costly (Eberhardt 1994) and that it is energetically cheap measuring 6030 cm and 35 cm high except during
(McCarty 1996) have both been questioned (Gaunt et al. breeding, when a pair of birds shared a double cage. The
1996; Verhulst & Wiersma 1997; Weathers et al. 1997). male birds could see and hear each other. The Fife
Whether singing is energetically costly or not therefore canaries could also see females. The diet was mixed seed
remains controversial, both because some of the empiri- (Super Canary for Fife canaries and Roller Mix for roller
cal data are contradictory and because it is unclear how canaries, Haiths, Cleethorpes, U.K.) fed ad libitum and
much energy expenditure must increase during an supplemented with broccoli, egg food (Sluis, Woodlea
activity before this will impose a fitness cost. Verhulst & Birds, Maldon, U.K.) mixed with hardboiled egg, soaked
Wiersma (1997) proposed that whether a particular seed (Easysoak, Haiths, Cleethorpes, U.K.) and vitamins
activity is energetically costly should best be considered (Daily Essentials, The Birdcare Company, Nailsworth,
by calculating its marginal cost, that is, how much of the U.K.). The birds had constant access to water and crushed
available energy that activity requires. A problem with oyster shell. Water baths were provided once a week. The
this approach, however, is that it is unclear how to photoperiod (daylight fluorescent strip lights) was
determine how much energy is available to an animal. 10:14 h light:dark in October, was increased evenly to
Previous studies of the energy cost of singing inside 16:8 h in January, remained constant until June and was
respirometry chambers have not included measurement decreased evenly to 10:14 h by October. Room air tem-
of a possible anaerobic component to metabolism during perature was 15–20C. The two breeds of canary were kept
vocalization (Weathers et al. 1997), for which the oxygen in separate rooms. Data were collected between December
debt would be repaid by increased aerobic metabolism and May. Bird husbandry and our experimental pro-
between songs. Nor has it been assessed whether birds cedures were approved by the Home Office, as was
sang as loudly inside respirometry chambers as when they returning the birds to local canary breeders at the end of
were not enclosed. It is important that song volume is not our work.
reduced inside respirometry chambers, since louder
sounds contain more energy. The energy cost of singing
Respirometry
was found to increase with song amplitude in a single
European starling that sang both quiet and loud songs We used as subjects the 12 male Fife canaries that sang
(Oberweger & Goller 2001). most frequently in their home cages, and all nine male
We measured the energy cost of singing by two breeds roller canaries. Each bird spent 30–90 min inside the
of canary (roller and Fife Fancy; body masses given in respirometry chamber during an experiment. The
Table 1) from the rate of oxygen consumption of birds respirometry chamber was placed in the room in which
singing inside a respirometry chamber. Each song bout the birds were normally kept. Food (broccoli and mixed
lasted several minutes and included both singing and seed) was provided inside the chamber. Eight Fife canaries
pauses between songs, so our measurements would and six roller canaries sang inside the chamber.
include any increase in gas exchange between songs that To measure the rate of oxygen consumption of the
compensated for an anaerobic component to metabolism canaries, we used an open-flow respirometry system con-
during song. The structure of the song differs substan- nected to an oxygen analyser (model 1100A or Xentra,
tially between the two breeds of canary that we studied. Servomex, Crowborough, U.K.). Air was pumped (Charles
The song of the Fife Fancy is louder, higher-pitched and Austen, Byfleet, U.K., diaphragm pump) through an
contains more phrases than that of the roller (Güttinger ABS plastic chamber (1712 cm and 15 cm high, Ensto,
1985; Mundinger 1995). We compared the cost of singing Briticent, Christchurch, U.K.) with a 1-cm-diameter
between these two breeds to assess whether song com- wooden perch 3 cm from the floor. The upper 9 cm of the
plexity influenced the energy cost of singing. To ensure chamber was transparent. We used a wet type gas flow
that the measurements were representative of song in meter (model DM3A, Zeal, London, U.K.) to measure the
more normal situations, we compared the characteristics flow rate of ambient air into the chamber. The meanSD
of song recorded inside the respirometry chamber and in flow rate of dry air at standard temperature and pressure
the birds’ home cages. We recorded the rate at which (STPD) across experiments was 1480450 ml/min
birds moved so that we could separate the energy costs of (N=206). Flow rates varied by less than 0.5% within
singing from those of moving. We also measured basal experiments, and these changes were taken into account
metabolic rate (BMR) so that we could express metab- in the calculations of the rate of oxygen consumption.
olism during singing as a multiple of BMR. Our results Gases were dried (silica gel) before and after passing
allow us to assess whether singing is energetically costly through the flow meter and after passing though the
or cheap in passerine birds. chamber. The zero point of the oxygen analyser was set
each week using oxygen-free dry nitrogen gas (BOC,
Guildford, U.K.) and the span was set before each exper-
METHODS iment using ambient air. The output from the oxygen
Birds and Husbandry analyser was sampled at 30 Hz using a microcomputer
with an analogue-to-digital converter (PC-ADH24, Bede
We purchased nine male roller canaries from local Technology, Sunderland, U.K.) and averaged every 2 s
breeders and purchased or bred 49 male Fife Fancy by customized software written in BASIC. The ambient
canaries (hereafter called Fife canaries). We kept the birds oxygen content of the air that was pumped through the
Table 1. Metabolic rates (W) of Fife canaries and roller canaries at night in the thermoneutral zone (BMR), and during the day while sitting (on a perch with the tarsi covered by the body
feathers and the body in a relatively horizontal posture), standing (in the singing posture with the tarsi exposed and the body held relatively vertically), eating and singing
Metabolic rate (W) Factorial increase t test, sing versus stand
Fife canaries
Dor31 20.1 0.25 — 0.72±0.06 0.74±0.07 0.84±0.10 0.39±0.09 3.41 — 1.18 3.91 22 0.0007 0.64
(13) (4) (14) (14)
DP37 17.1 0.28 — 0.55±0.08 0.57±0.04 0.50±0.03 0.21±0.09 1.82 — 0.92 −1.37 6 0.22 −0.49
(6) (4) (6) (6)
DW2 17.7 0.33 — 0.60±0.07 0.70±0.09 0.62±0.09 0.40±0.09 1.89 — 1.03 0.41 5 0.70 0.18
(3) (11) (7) (7)
J6 22.4 0.31 0.54±0.05 0.60±0.05 0.62±0.07 0.61±0.10 0.24±0.09 1.98 1.14 1.02 0.22 2 0.85 0.15
(2) (8) (11) (3) (3)
J9 21.5 0.23 — — 0.59±0.11 0.73±0.07 0.35±0.07 3.13 — — — — — —
(17) (7) (7)
J22 23.7 0.35 0.52 — 0.73±0.09 0.80±0.06 0.18±0.06 2.28 1.54 — — — — —
(1) (21) (2) (2)
LY4 21.5 0.37 — 0.62±0.05 0.77±0.07 0.71±0.09 0.33±0.11 1.95 — 1.14 2.91 15 0.01 0.60
(6) (4) (19) (19)
TW47 28.3 0.25 0.46±0.05 0.66±0.02 0.78±0.09 0.73±0.04 0.56±0.17 2.93 1.59 1.11 4.19 14 0.0002 0.80
(2) (7) (15) (10) (10)
Mean 21.5±3.5 0.30±0.05 0.51±0.04 0.62±0.06 0.69±0.08 0.69±0.11 0.33±0.12 2.46±0.67 1.42±0.25 1.07±0.10 (6) — — — 0.31±0.47
(8) (8) (3) (6) (8) (8) (8) (8) (3) (6) (6)
Roller canaries
Var 19.8 0.29 — 0.73±0.16 0.75±0.17 0.68±0.11 0.39±0.09 2.32 — 0.93 −0.69 7 0.51 −0.25
(10) (19) (21) (21)
208 21.9 0.35 — 0.69±0.08 0.72±0.09 0.71±0.08 0.54±0.15 2.00 — 1.02 0.3 7 0.8 0.11
(5) (7) (17) (17)
1725 19.9 0.32 0.43±0.05 0.51±0.09 0.58±0.06 0.62±0.01 0.36±0.10 1.92 1.44 1.21 1.23 4 0.3 0.52
(5) (4) (11) (2) (5)
109 18.7 0.25 0.40±0.03 0.52±0.06 0.53±0.06 0.56±0.06 0.32±0.07 2.20 1.40 1.06 1.08 6 0.32 0.40
(5) (17) (11) (5) (5)
23 21.7 0.22 — 0.56±0.06 0.56 0.62±0.11 0.54±0.17 — — 1.11 0.63 5 0.56 0.27
(16) (1) (5) (5)
209 17.9 0.38 — 0.64±0.09 0.75 0.71±0.08 0.44±0.18 — — 1.11 2.43 23 0.02 0.45
(14) (1) (24) (24)
Mean 20.0±1.6 0.30±0.06 0.41±0.02 0.61±0.11 0.65±0.10 0.64±0.07 0.43±0.09 2.11±0.15 1.42±0.02 1.05±0.07 — — — 0.25±0.29
(6) (6) (2) (4) (6) (4) (6) (4) (2) (6) (6)
Only data from bouts of sitting, standing and singing during which the birds did not move are included, except for the data given in italics for two roller canaries (birds 23 and 209) for which
movement rates did not differ between singing and standing birds. Proportion of song refers to the proportion of time for which a bird sang during a song bout. The t tests compare metabolic
rate between perching and singing within individual birds. Sample sizes are given in parentheses. Means are given±SD. The mean values given for each breed of canary are the means of the
mean values for each bird. The durations of measurements of metabolic rate across Fife canaries were 3.0±2.4, 6.6±1.5, 6.8±1.6 and 5.9±2.1 min for sitting, standing, eating and singing,
respectively. The durations of measurements of metabolic rate across roller canaries were 7.5±2.6, 5.2±3.0, 6.4±5.2 and 6.5±1.0 min for sitting, standing, eating and singing, respectively.
WARD ET AL. THE ENERGY COST OF CANARY SONG
895
896 ANIMAL BEHAVIOUR, 66, 5
chamber was measured before and after each bird was categorized as singing, standing (in the singing posture
placed in the chamber. We used these measurements to with the tarsi exposed and the body held relatively
compensate for any drift in the output of the oxygen vertical), sitting (with the legs and feet covered by the
analyser during an experiment. body feathers and the body in a relatively horizontal
We calculated the rate of oxygen consumption by posture) or eating. We defined song duration as the
multiplying the difference between the fractional oxygen period during which a bird produced a series of phrases in
content of ambient and excurrent air by the incurrent rapid succession. Song duration was always shorter than
flow rate (corrected to STPD). Carbon dioxide was not the equilibration time of the chamber. Our measure-
absorbed; this maximizes the accuracy of calculated ments of the energy cost of singing therefore refer to the
energy expenditure when the respiratory quotient is not average metabolic rate during bouts of songs. Song bouts
measured (Koteja 1996; Speakman 2000). We converted began or ended when the bird perched, sat, ate, moved (if
rates of oxygen consumption to energy equivalents using the bird had not otherwise moved during the singing
an oxycalorific value of 20.92 J/ml (derived from Brody bout) or sang at a rate that differed by more than 50%
1945 for a respiratory quotient of 1). We measured basal from that during the rest of the bout. Movement was
metabolic rate (BMR) by placing each bird in the assessed from the rate at which birds moved their feet (i.e.
respirometry chamber overnight at 30C (within the to hop between the perch and the chamber floor or to
thermoneutral zone for passerine birds of similar mass to rotate by 180 upon the perch).
canaries; Calder & King 1974) inside a constant tempera-
ture incubator (model INL-401N-010, Gallenkamp,
Song Duration and Amplitude
Loughborough, U.K.). BMR was calculated from the
lowest metabolic rate over 5 min. We recorded Fife canary song using a Tandberg TM6
To determine the washout characteristics of the microphone connected to a computer (Dell Dimension
respirometry chamber, we reduced the oxygen content of 4100, Creative Sound Blaster AudioPCI 64V soundcard)
the gases inside the chamber to 20.8% (a level similar to running Goldwave (version 4.23, Goldwave Inc., St
that experienced by the birds during experiments) using John’s, Canada) software with a sampling rate of 22 kHz
nitrogen gas (BOC). We then pumped ambient air and 16-bit resolution. The centre of the microphone was
through the chamber at rates within the range used 5 cm in front of and 2 cm below the bird’s bill when the
during experiments until the oxygen content of the bird sang facing the microphone while standing on
excurrent gases returned to that of ambient air. A Fife the perch inside the respirometry chamber. We used the
canary that had died of natural causes was placed on the video equipment described above to assess when the birds
perch inside the chamber to make the internal volume sang in the same position relative to the microphone
and air flow characteristics of the chamber as similar as inside their home cages. We recorded song from eight
possible to those during experiments with living birds. A Fife canaries standing in this position relative to the
meanSD of 5200200 ml (N=3) of ambient air was microphone inside the respirometry chamber and from
passed through the chamber before the oxygen content seven of these birds in their home cages.
of the gases leaving the chamber recovered by 95% from We calculated the proportion of time spent singing
the level to which it had been perturbed. We therefore during each bout of song from the summed durations
assumed that the equilibrium rate of oxygen consump- of all songs divided by the duration of the bout. The
tion of a bird for a given behaviour was reached after that duration of each Fife canary song was measured from the
behaviour had been performed consistently for long digitized recordings to the nearest 0.1 s using Avisoft-
enough for 5200 ml of gas to have passed through the SASLab Pro (Berlin, Germany, version 3.94c). The dur-
chamber (hereafter called the chamber equilibration ation of each roller canary song was measured from the
time). The meanSD chamber equilibration time was videos, with a stopwatch. Comparison of the meanSD
3.81.1 min (N=206 experiments). The combined time duration of Fife canary songs that were measured both
taken for gases to pass from the respirometry chamber to with the stopwatch and from the digitized recordings
the oxygen analyser and for the analyser to respond to showed that song duration was measured to the nearest
any change in concentration of the gases was 3–5 s, 0.40.1 s (N=230 songs) using the stopwatch. The pro-
depending on the flow rate. This response time was portion of singing during song bouts measured using the
taken into account when we matched the data on the stopwatch differed by 23% (N=8 bouts) from that
oxygen content gases leaving the chamber with bird measured from the digitized recordings.
behaviour. We did not use an instantaneous correction All eight Fife canaries included three types of phrase
(Bartholemew et al. 1981) to calculate the rate of oxygen during the first 5 min of song that we recorded from each
consumption, because this was unnecessary when bouts bird in his home cage. These were a series of elements
of behaviour were long enough to allow us to measure with continuously descending pitch, an ‘A’ phrase (Vallet
equilibrium rates of oxygen consumption. & Kreutzer 1995; Vallet et al. 1998), and a series of
elements in which descending and low pitch notes alter-
nated (Fig. 1). To measure the peak amplitude and the
Bird Behaviour
frequency of the peak amplitude of these elements, we
We recorded bird behaviour on VHS video (JVC GR-A used the spectral characteristics function from the log-
X280 video camera connected to a Panasonic NV-FS90 arithmic power spectrum in Avisoft-SASLab Pro. We
video recorder) during experiments. Bird behaviour was sampled 3–15 repetitions of each element of the three
WARD ET AL. THE ENERGY COST OF CANARY SONG 897
1 t12 =1.96, P=0.07; bird 23: t6 =0.1, P=0.9). When the data
(a) from these two birds were included in the analysis,
0.8 metabolism during singing was 1.050.07 times greater
than that during standing (Z=1.8, N=6 birds, P=0.03;
0.6 Table 1, Fig. 2b). Metabolism during singing without
moving did not differ from that during eating (Z=0.2,
0.4 N=4 birds, P=0.4). Metabolic rate during singing without
moving was 2.110.15 (N=4 birds) times BMR and
Metabolic rate (W)
Table 2. Relations between the proportion of time spent singing and the metabolic rate of three male Fife canaries
Metabolic rate for continuous song
Mean (95% prediction range)
of thermoregulation changes with temperature. Singing F4,27 =4.3, P=0.01 with Tukey post hoc multiple compari-
caused a factorial increase in metabolism over sitting that sons). The cost of singing measured for Carolina wrens
was intermediate between those relative to standing and may be greater than that of the other species because of
BMR partly because the thermoregulatory costs of sitting difficulties in measuring transient changes in oxygen
are likely to be lower than those of standing birds, consumption in a relatively large respirometry chamber,
because sitting birds covered their legs. The legs can make because the birds moved as well as sang during measure-
a substantial contribution to heat loss (Ward et al. 1999). ments, or because of insufficiently detailed mathematical
Metabolic rate during sitting was also likely to be lower treatment of the data (Frappell et al. 1989; Gaunt et al.
than that of singing or standing because the birds 1996; Ortigues et al. 1997). Movements during song bouts
appeared to be less alert during sitting. The energy cost of were not monitored during Eberhardt’s (1994) study of
song production itself should therefore be assessed by Carolina wrens, so the costs of any movements would
comparing metabolism during song with that during have been included in the measured cost of singing (Horn
standing in the same posture (mean factorial increase et al. 1995; Gaunt et al. 1996). If free-living Carolina
1.05–1.07). Our data thus support the hypothesis that wrens move during song bouts, the cost of movements
song production by canaries is energetically cheap. should be included in the cost of the display, but in this
case the cost of the display would clearly be greater than
Does Singing Generally Increase Metabolic Rate in that of song alone. The increased energy expenditure of
sage grouse, Centrocercus urophasianus, that displayed at
Birds?
higher rates was attributed partially to higher movement
Singing increases the rate of oxygen consumption of rates (Vehrencamp et al. 1989). Song production is likely
Carolina wrens, European starlings, zebra finches and two to make a minor contribution to the cost of display by
breeds of canary singing inside respirometry chambers birds such as skylarks, Alauda arvensis, that sing while
(Eberhardt 1994; Oberweger & Goller 2001; Franz & flying (Hedenström & Alerstam 1996).
Goller 2003; this study). Singing also increases the rate Because consistent results have been obtained from
of overnight mass loss (which is presumably correlated more than one breed of canary and two other passerine
with the rate of consumption of body reserves and there- species (Oberweger & Goller 2001; Franz & Goller 2003;
fore with energy expenditure) of free-living nightingales this study), it is likely that the modest cost of singing,
(Thomas 2002). In contrast, crowing by junglefowl or relative to that of standing during the day, in these
cockerels inside a respirometry chamber does not cause a species is representative of most passerine birds. Winter
measurable increase in the rate of oxygen consumption wren song is louder in relation to the mass of the bird
(Chappell et al. 1995; Horn et al. 1995). Although cock- than that of many other passerines (Brackenbury 1979),
erel crows are loud and inefficient (Brackenbury 1977), and singing by wrens of all genera is considered to be
they were performed for a much lower proportion of the particularly loud (Brewer 2001). Wren song may therefore
time than that for which our canaries sang (0.05–0.06 for be energetically more costly than that of most other
crowing: Chappell et al. 1995; Horn et al. 1995; 0.33–0.43 passerine birds, but further empirical data are required to
for singing: this study). Cockerel crows also contain determine whether this is the case.
substantially less acoustic power in relation to the mass of
the bird (60 mW/kg) than the songs of typical passerine
The Energy Cost of Song and the Dawn Chorus
birds such as the song thrush, Turdus philomelos, winter
wren, Troglodytes troglodytes, and European robin, Erith- Singing appears to increase metabolism only slightly
acus rubecula (870, 600 and 300 mW/kg, respectively; over that of a bird standing during the day, but the
Brackenbury 1979). The substantially higher proportion factorial increase over BMR is much greater. The
of time spent singing and greater volume of passerine additional energy cost of singing at night (rather than
song in relation to their body mass probably explain why roosting with a metabolic rate closer to BMR) would be
the cost of passerine song has been detected over other substantially greater than that of singing during the day
fluctuations in metabolism, but the cost of crowing by (rather than standing). This is because the energy cost of
junglefowl and cockerels has not. the activity that singing would replace would be much
The measurements of metabolic rate of singing lower at night than during the day. Similarly, singing
Carolina wrens (Eberhardt 1994) have generally been during the dawn chorus could cause a greater factorial
regarded as anomalously high (Gaunt et al. 1996; increase in energy costs than singing during the day if
Oberweger & Goller 2001; Thomas 2002). The reported birds sing rather than sleep at this time. The energy cost
factorial increase in the metabolism of Carolina wrens of singing at night or at dawn would presumably be a
during song (3.61.1BMR, with a maximum value of factorial increase of between 1.4 (the difference between
9BMR: Eberhardt 1994) was greater than that of Fife sitting and singing during the day in canaries) and 2.2–
canaries (2.40.6BMR, this study), roller canaries 2.6 (the difference between BMR and singing in canaries).
(2.40.4BMR: this study; Oberweger & Goller 2001), The extent of the increase in metabolism during singing
zebra finches (2.00.2BMR: Oberweger & Goller 2001) at night would depend on the difference in temperature
or European starlings (2.10.06BMR: Oberweger & (and hence thermoregulatory costs) between the micro-
Goller 2001; ANOVA with species, or breed of canary, as a habitats in which birds would roost or sing, changes in
factor and the mean factorial increase in metabolism thermoregulatory costs due to altering posture and
during singing of individual birds as independent data: altered alertness. Sound production itself would make a
WARD ET AL. THE ENERGY COST OF CANARY SONG 901
minor contribution to metabolism. Our results are there- also be less effective than singing, since a bird can only be
fore consistent with the energetic cost hypothesis for the in one place at a time, but woodland birds can often
evolution of the dawn chorus (McNamara et al. 1987) be heard simultaneously in all directions for greater
although they do not exclude other explanations for the distances than they can be seen.
dawn chorus such as enhanced sound transmission
(Brown & Handford 2003) or reduced foraging success
(Kacelnik & Krebs 1983) at this time. Acknowledgments
Franz, M. & Goller, F. 2003. Respiratory patterns and oxygen Oberweger, K. & Goller, F. 2001. The metabolic cost of bird-
consumption in singing zebra finches. Journal of Experimental song production. Journal of Experimental Biology, 204, 3379–
Biology, 206, 967–978. 3388.
Frappell, P. B. R., Levin, H. A. & Baudinette, R. V. 1989. Under- Ortigues, I., Dussap, C-G. & Anglaret, Y. 1997. Comparison
standing respirometry chambers: what goes in must come out. of various methods of calculating the instantaneous res-
Journal of Theoretical Biology, 138, 479–494. piratory gaseous exchanges from discrete measurements in
Gaunt, A. S., Bucher, T. L., Gaunt, S. L. L. & Baptista, L. F. 1996. respirometry chambers. Journal of Theoretical Biology, 185, 489–
Is singing costly? Auk, 113, 718–721. 501.
Gil, D. & Gahr, M. 2002. The honesty of bird song: multiple Prestwich, K. N. 1994. The energetics of acoustic signalling in
constraints for multiple traits. Trends in Ecology and Evolution, 17, anurans and insects. American Zoologist, 34, 625–643.
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