Impotentie brengt een constant ongemak met zich mee, net als fysieke en psychologische problemen in uw leven cialis kopen terwijl generieke medicijnen al bewezen en geperfectioneerd zijn
JOURNAL OF AVIAN BIOLOGY 33: 269 – 275, 2002
The natural plasma testosterone profile of male blue tits during
the breeding season and its relation to song output
Katharina Foerster, Angelika Poesel, Hansjo¨rg Kunc and Bart Kempenaers
Foerster, K., Poesel, A., Kunc, H. and Kempenaers, B. 2002. The natural plasmatestosterone profile of male blue tits during the breeding season and its relation tosong output. – J. Avian Biol. 33: 269 – 275.
In male birds, the gonadal hormone testosterone (T) is known to influence territorialand mating behaviour. Plasma levels of T show seasonal fluctuations which vary inrelation to mating system and social instability. First, we determined the natural Tprofile of male blue tits Parus caeruleus
during the breeding season. We found thatplasma levels of T increased at the onset of nest building. Thus, the increase incirculating T was not associated with territory establishment, nor with the fertileperiod of the males’ mates. In most individuals, T levels dropped to values close tozero during the period of chick feeding. Second, we investigated the relationshipbetween plasma levels of T and male age, size, and singing behaviour. During themating period, T levels did not differ between 1 yr old and older males and did notcorrelate with body size or condition. However, song output during the dawn chorustended to be positively correlated with T levels. Therefore, if high T levels are costly,song output might be an honest indicator of male quality in blue tits. Finally, weshow that plasma levels of T are significantly higher during the night than during theday. This pattern has also been observed in captive non-passerine birds, but itsfunctional significance remains unknown.
. Kunc and B
, Max PlanckResearch Centre for Ornithology
Testosterone (T) plays an important role in driving
1990). In all temperate passerines with paternal care, T
male reproductive behaviour in birds. Temporal pat-
levels drop rapidly during the incubation phase and are
terns in the plasma levels of this sex steroid are trig-
low during the period of nestling feeding (e.g. Ball and
gered by several factors including photoperiod and
Wingfield 1987, Vleck and Brown 1999). T levels may
food availability (Wingfield 1983), as well as social
show repeated peaks or remain elevated over the entire
stimuli (Wingfield et al. 1990). Different T profiles
breeding season, when multiple clutches, extra-pair cop-
during the breeding season have been related to the
ulations, or polygyny are frequent (e.g. Hegner and
social systems and the mating strategies of the observed
Wingfield 1986, Beletsky et al. 1995, Wada et al. 1999).
bird species (Wingfield et al. 1990, Beletsky et al. 1995,
As a consequence of the T-mediated trade-off between
Vleck and Brown 1999). Outside the breeding season,
sexual behaviour and parental care (Hegner and
when competition for nesting sites and copulation part-
Wingfield 1987, Beletsky et al. 1995, Raouf et al. 1997),
ners is low or absent, plasma levels of T are low
high T levels during the late breeding season enable
(nonbreeding baseline, sensu
Wingfield et al. 1990).
males of these species to acquire additional breeding or
When breeding territories are established and when
copulation partners, at the cost of reduced paternal
mating takes place, T levels rise up to 20-fold to a
breeding baseline (e.g. Ball and Wingfield 1987, Vleck
T is known to affect also the development of mor-
and Brown 1999, Wada et al. 1999). Within the breed-
phological, sexually selected traits (e.g. Zuk et al. 1995,
ing season, large fluctuations in T levels are observed,
Eens et al. 2000, Evans et al. 2000, Peters et al. 2000).
for instance during social challenges (Wingfield et al.
Similarly, the role of T in the expression of behavioural
traits involved in sexual selection is well established.
served on 5 March, first eggs were laid on 5 and 6
Males with elevated T levels defend larger territories
April, and the last eggs (except for a few replacement
(Silverin 1980, Wingfield 1984, Chandler et al. 1994),
clutches) between 29 April and 8 May. The period
are more aggressive (Beletsky et al. 1990, but see
from 5 March to 8 May includes nest building and
Romero et al. 1998), and show more elaborate
egg laying and was therefore defined as the mating
courtship displays (Enstrom et al. 1997, Hill et al.
. We refer to the periods before and after the
1999) than males with low T levels. Males with high
mating period as the territory defence period
T levels also sing more (e.g. Silverin 1980, Ketterson
chick feeding period
et al. 1992, Hunt et al. 1997), although some studies
Blue tits were captured at night (20:00 – 00:00 hours)
have not found this effect (Beletsky et al. 1995, Saino
when they slept inside the nestbox and were placed
and Møller 1995). Thus, it is evident that elevated T
separately in cotton bags. Blood samples (20 – 100 ml)
levels as observed in males before and during the
from the brachial vein were taken within 30 min of
females’ fertile period enable behaviours necessary to
capture. During the mating period some males were
acquire and defend nest sites and copulation partners.
caught at day in food-baited potter traps. When the
However, males vary greatly in their peak level of
chicks were 8 – 9 days old, parents were caught inside
circulating T, and this might be related to their repro-
the nestbox when feeding young. In this case, blood
ductive success. Few studies have investigated the ef-
samples were taken within 10 min of capture. After
fect of this variation on the intensity of sexually
centrifugation, plasma was stored at − 70°C until
selected traits and sexual behaviour (Erickson 1970,
Searcy and Wingfield 1980, Borgia and Wingfield
Individuals were colour-banded, sexed and aged (1
yr old or older bird, based on plumage characteristics,
In this study, we investigated the natural plasma T
Svensson 1992). We measured tarsus length to the
levels in a population of blue tits Parus caeruleus
nearest 0.1 mm using callipers and body mass to the
during the breeding season. This species is mostly so-
nearest 0.1 g using a digital balance. We calculated
cially monogamous, but polygyny is not uncommon
body condition as the residuals of a regression of
(Kempenaers 1994). Extra-pair copulations are fre-
quent and females choose extra-pair mates that areolder and have longer tarsi than their social partner(Kempenaers et al. 1997). Blue tits are non-migratory
Song recording and analysis
and males start singing and defending their territoryas early as January/February, long before nest build-
The complete dawn song was recorded once for 28
ing and egg laying. Only during the fertile period of
individual males when their females were fertile and
the mate, do males show a distinct dawn chorus. A
previous study on the same population showed that
Windows. For a detailed description of recording
song output was related to female fecundity and is
methods, song analysis, and song variables see Poesel
therefore probably under sexual selection (Poesel et al.
et al. (2001). Here we used two measures of song
output: mean strophe length (duration of strophes in
The aim of this study was twofold. First, we de-
seconds), and mean percentage performance time
scribe the temporal changes in natural plasma levels
(strophe length divided by the sum of the strophe and
of T in male blue tits during the breeding season.
following pause lengths). As mean percentage perfor-
Second, we relate plasma levels of T during the pe-
mance time declined with recording date (Poesel et al.
riod when T levels are elevated with male characters
2001), we corrected all analyses for recording date.
that are related to their reproductive success, namely
Versatility (number of song type switches during the
age, size, and song characteristics.
entire dawn chorus multiplied by the number of dif-ferent song types) was used as a measure of songcomplexity. For 10 of the recorded males, the T levelwas measured during their females’ fertile period
(song recordings on days 2 to 15, T levels measuredon days
Study area and field methods
− 3 to 10, 0 = day of the first egg).
We studied a population of blue tits breeding in nest-boxes in a mixed deciduous woodland at Kolbeter-
berg in Vienna (48°13%N, 16°20%E), Austria. Duringthe breeding season, nestboxes were checked every
An enzyme immunoassay (EIA) was used to assess
second day to determine the onset of nest building,
the concentration of T in the plasma samples col-
laying, hatching, and fledging. In three study seasons
lected in 1998. In general, we followed the protocol of
(1998 – 2000), the first nest building activity was ob-
Palme and Mo¨stl (1993). Five to 20 ml of plasma was
diluted 1:10 with double distilled water, extracted with
5 ml diethyl ether and reconstituted in 50 ml assaybuffer. Mean recovery was 84%. The assay sensitivity
Temporal changes in plasma levels of T
was 0.06 ng/ml for a plasma sample of 20 ml.
The natural profile of circulating T in blue tit males
Plasma samples collected in 1999 and 2000 were
during the breeding period changed from low non-
analysed in a radio immunoassay (RIA). Seven to 50 ml
breeding levels in late winter to high levels at the time
of plasma was diluted to 100 ml with assay buffer. We
of nest building (Fig. 1a). High T levels were observed
used a modification (Fusani et al. 2000) of the RIA
from 17 March onwards, two weeks before the first
procedure of Wingfield and Farner (1975). The sampleswere reconstituted in 160 ml assay buffer. Mean recov-
females in the study area were fertile. Individual males
ery was 90% and the sensitivity of the assay was 0.03
showed elevated T levels as early as 27 days before their
ng/ml for a plasma sample of 20 ml. The intra-assay
social mate became fertile (Fig. 1b). T levels remained
variation was 2.8%. All samples were measured in a
elevated during egg laying, but dropped when males
All measured plasma levels of T fell within the range
of the two standard curves (EIA: 0.3 – 80 pg/tube, RIA:0.1 – 200 pg/tube). When analysing hormone levels cor-rected for mean recovery, our conclusions did notchange, and we report uncorrected values.
In 1999 and 2000, some of the resident males in the
study area received T-releasing or control implantsduring the mating period. We implanted self-dissolvingpellets with a release time of two months during the lastweek of February and the first week of March (K.
Foerster and B. Kempenaers unpubl.). T levels duringthe mating period did not differ between untreatedmales and males with control implants in these twoseasons (n = 23, x¯ = 1.96 91.48 ng/ml and n=6, x¯=1.61 91.73 ng/ml, resp.; t=0.497, P\0.6). Also, Tlevels of all males (untreated and control-implanted)measured during the mating period did not differamong the three study seasons (each individual usedonly once, Kruskal-Wallis test, H = 2.165, df = 2, P \0.3), or between the two T assay methods (EIA: n = 14,x¯ = 1.74 90.43 ng/ml and RIA: n=15, x¯=1.9590.34ng/ml;
Therefore, we pooled all data obtained during themating period.
Statistical analyses follow Sokal and Rohlf (1995) andwere performed using SPSS 10.0 and StatXact 3 forWindows. Plasma levels of T during the mating periodwere normally distributed after log transformation. Weapplied this transformation when testing for correla-tions of T levels with sampling date and time, and with
Fig. 1. Natural profile of plasma testosterone levels in resident
male characteristics. Untransformed data are presented
male blue tits during the breeding season. Data are from 68
as means 9SE, transformed data as back-transformed
individuals in three seasons. Circles: T levels measured during
means with upper and lower 95% confidence limits. We
day, triangles: T levels measured during night. Multiple data
applied two-tailed tests and used Bonferroni corrections
points are labelled with numbers. (a) T levels in relation tosampling date (1 = 1 Jan). Filled circles: socially polygynous
in the case of multiple comparisons.
males, filled triangles: birds not found breeding. The shaded
Some males were caught repeatedly over the three
area indicates the mating period. (b) T levels in relation to egg
years of this study and all data are presented in Fig. 1.
laying of the social mate (0 = day when the first egg was laid).
The shaded area indicates the maximum fertile period of blue
For each statistical test we used only one data point per
tit females, starting at day − 5 and ending at day 15 (maxi-
were feeding nestlings (Fig. 1a). Four polygynous males
chick feeding (partial correlation: n = 25, r = 0.44, P =
that were sampled during the nestling phase had T
levels comparable to those of monogamous males (Fig.
Samples that were collected at night had higher T
concentrations than samples collected at day (matingperiod only, night: n = 22, x¯ = 2.31 90.29 and day:
This study shows that the profile of plasma levels of T
n = 7, x¯ = 0.39 90.10; U=4.0, PB0.001). Also, T lev-
in male blue tits during the breeding season follows the
els at night were more variable than T levels measured
typical pattern of socially monogamous temperate
during day (mating period only, Conover test for equal-
passerines. The increase in T levels was not associated
ity of variances with unequal means, T = 506.0, P =
with territory establishment, or with the fertile period
0.01). During the mating period, T levels did not vary
of the males’ mates. T levels during the mating period
with date or sampling hour (ANCOVA: day/night sam-
seem to correlate with song output during the dawn
chorus, which is a sexually selected trait (Poesel et al.
test for differences in T levels between the three periods
The natural profile of plasma levels of T in male blue
of the breeding season, we compared only samples
tits showed a single, distinct peak at the time whencompetition for breeding sites and copulation partners
obtained at either night or day. At the time of territory
is high. T levels increased at the beginning of nest
defence in January and February, all samples were
building, long before females were fertile. This is similar
collected at night, and their T levels were significantly
to results from other studies showing that T levels
lower than those of samples obtained at night during
increase at the onset of nest building (e.g. Silverin et al.
the mating period (territory defence: n = 9, x¯ = 0.11 9
1986, Schoech et al. 1991) or, in migratory species,
0.03 and mating period: n = 19, x¯ = 2.32 90.32; U=
when females arrive at the breeding sites (e.g. Silverin
0.0, P B0.001). During the mating period, most birds
and Wingfield 1982, Hunt et al. 1997). Blue tit males in
were captured at night, while all birds were captured at
our study population occupied their territories during
day when feeding chicks. However, considering only
winter (own obs.). Therefore, most males that later
daytime samples, T levels still dropped significantly
bred on the study plot had already established a terri-
from the mating period to the chick feeding period
tory long before the onset of nest building (either by
(mating period: n = 7, x¯ = 0.39 90.10 and chick feed-
occupying the same area as during the preceding spring
ing: n = 42, x¯ = 0.16 90.06; U=57.5, P=0.008).
or by establishing a territory sometime during winter).
However, some 1 yr old males arrived in the study areaas late as March and tried to occupy territories. Aggres-
Testosterone and male characteristics
sive male-male interactions were frequent at the timewhen T levels started to rise, but also occurred earlier
To correlate male characteristics and T levels, we con-
(own obs.). Mate guarding was less intensive during
sidered only samples taken at night during the mating
early nest building compared to when females were
period. Samples taken during the day were too few for
fertile (K. Foerster and B. Kempenaers unpubl.) Copu-
meaningful analyses. Male T levels did not differ be-
lations were observed almost exclusively during the
tween 1 yr old males and older individuals (1 yr old:
female’s fertile period (own obs.). Thus, high T levels in
n = 12, x¯ = 1.92 [CI 1.19, 3.11] and older males: n = 9,
blue tit males were less correlated with mate guarding
= 1.76 [CI 1.12, 2.76]; t = 0.29, P \0.7). T levels were
than with territorial behaviour, as predicted by the
not correlated with male size (tarsus: n = 22, r = 0.17,
‘‘challenge hypothesis’’ (Wingfield et al. 1990). How-
P \0.4) or male condition (n=17, r = −0.05, P\
ever, territories are also defended during the period of
0.8). Song output during the dawn chorus measured as
mean percentage performance time tended to increase
Elevated T levels were observed over a period of 4.5
with T level (partial correlation corrected for recording
weeks, which is comparable to the pattern found in
date: n = 7, r = 0.86, P = 0.03; significance level after
other temperate monogamous species (Wingfield et al.
Bonferroni: P = 0.017). Mean strophe length and song
1990, Vleck and Brown 1999). Birds with prolonged
versatility were not significantly related to T level (n =
competition over access to extra-pair mates, with multi-
7, r = 0.62, P \0.1 and n=5, r =0.42, P\0.4, re-
ple clutches, or polygyny maintain high T levels over a
spectively). Mean percentage performance time was
long period (Hegner and Wingfield 1986, Vleck and
positively related to male body mass and condition in
Brown 1999, Wada et al. 1999). In our population
the prebreeding period (partial correlation: n = 28, r =
extra-pair paternity occurs frequently (59.5% of all
0.51, P = 0.007 and r = 0.51, P = 0.007, respectively;
broods in 1998 contained at least one extra-pair young,
significance level after Bonferroni: P = 0.0125), and also
own unpubl. data). However, females produced only
tended to correlate with male body mass later during
one clutch (with the exception of a few replacement
clutches) and laying was highly synchronous (breeding
non-significant results should be interpreted cautiously,
synchrony index for 1998 – 2000: 69 – 72% see Kempe-
as they might be caused by high individual variation
naers 1993). Therefore, the availability of possible ex-
tra-pair mates for males was limited to a short time
Although blue tit females prefer older and larger
interval that overlapped greatly with the fertile phase
males as extra-pair partners (Kempenaers et al. 1997),
of the social partner. Thus, in this population few
we did not find any evidence that these characteristics
females were fertile during the chick-feeding period
were related to T levels during the mating period.
and males did not face the usual trade-off between
However, we found that males with higher T levels
mating and parental effort. In other species, T has
during this period tended to have a higher song output
been shown to play an important role in the regula-
at dawn. Furthermore, males with higher song output
tion of the behaviours involved in this trade-off (Belet-
were in better condition before and during the breed-
sky et al. 1995, Raouf et al. 1997). Males with high T
ing season. Song is a dynamic behavioural trait that
levels during the nestling stage feed nestlings less, or
plays an important role in mate choice in passerines
not at all (e.g. Silverin 1980, Ketterson et al. 1992,
(Andersson 1994). In the blue tit, song output might
Schoech et al. 1998, Hunt et al. 1999). In blue tits,
be under sexual selection, since males with a higher
both monogamous and polygynous males had low lev-
song output were paired to females that laid earlier in
els of circulating T during the nestling stage, compara-
the season (Poesel et al. 2001). Several studies have
ble to levels before the mating season (with two
shown that individual variation in the avian song con-
exceptions, see Fig. 1) and provided about half of the
trol system and syringe control is influenced by T (e.g.
food to the nestlings (own unpubl. data).
Luine et al. 1980, Smith et al. 1997). Also, T affects
We found that plasma levels of T were higher and
the frequency of spontaneous song during the day (e.g.
more variable during the night than during the day.
Ketterson et al. 1992, Hunt et al. 1997, but see Saino
Higher T levels during night or a peak at the end of
and Møller 1995). In this study we show that within a
the dark phase have been reported in domestic ducks
period of continuous song (dawn chorus), males with
(Balthazart 1976), domestic fowl (Bachman et al. 1987,
higher T levels tend to have a higher song output. To
Schanbacher et al. 1974), and ring doves (Balthazart et
our knowledge only one other study has so far related
al. 1981). These studies describe circadian patterns ofT levels in captive birds, but the functional signifi-
a song characteristic (other than song rate) to T levels
cance of the observed differences is unclear (Balthazart
during the breeding season: Galeotti et al. (1997)
et al. 1981). To our knowledge, our study provides the
found that male barn swallows Hirundo rustica
first data on free-living individuals and on a passerine.
high T levels produced longer rattles than males with
Our results may have been influenced by the sampling
low T levels. Interestingly, that study also showed that
methods; handling time during day was generally
a characteristic of the rattle (peak amplitude fre-
shorter than during night, when birds were trans-
quency) was related to body condition. Similarly, we
ported to a shed before bleeding. However, several
found a relation between song output and condition in
studies have shown that T levels were not influenced
significantly by long handling times (Wingfield et al.
It is probably costly for a male to maintain high T
1982, Silverin 1998, Kitaysky et al. 1999).
levels over a long period, as T has been shown to be
Some studies indicate that the relative concentration
immunosuppressive in birds (Verhulst et al. 1999, Pe-
of circulating T during the period when T levels are
ters 2000, but see Hasselquist et al. 1999). If high T
elevated correlates with the intensity of aggressive and
levels are costly and the intensity of a sexually selected
sexual behaviour (Searcy and Wingfield 1980, Borgia
trait depends on T, then this trait might signal male
and Wingfield 1991, Enstrom et al. 1997). However, it
quality (Folstad and Karter 1992). Thus, our data
is still unclear whether the individual variation in T
indicate that song output during the dawn chorus
levels during the mating period affects male attractive-
might provide an honest signal of male quality.
ness to females (Hill et al. 1999). Few studies haveinvestigated correlations between T levels and male
– We thank Katharina Peer and AgnesTu¨rk for help in the field and Cheryl Bishop and Etie¨nne
characteristics in free-living birds. Single measurements
Vermeirssen for analysing plasma samples. Erich Mo¨stl (Vet-
of T levels may depend strongly on time of day (this
erinary University, Vienna) and Tom Van’t Hof (Max Planck
study) and on the state of the animal prior to sam-
Research Centre for Ornithology, Andechs) generously pro-
pling. Social interactions quickly raise plasma levels of
vided resources and help in their labs. Raphael-ThomasKlumpp and Alfred Fojt from the Institute of Silviculture,
T in some bird species (Wingfield and Wada 1989,
Vienna, provided access to their facilities in the study area. We
Wingfield et al. 1990, Wikelski et al. 1999). Thus, if
thank Virginie Canoine, Leonida Fusani, Tom Van’t Hof, and
only one sample is taken per individual, large fluctua-
two anonymous referees for helpful comments on the
tions over time might mask relations of T levels with
manuscript. We are grateful to Hans Winkler from the Kon-rad Lorenz Institute for Comparative Ethology (KLIVV),
male characteristics. We were not able to take multiple
Vienna, for continuous support during this study. KF was
measurements during the mating period. Thus, our
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Gary Swan, Ph.D. Director, Center for Health Sciences SRI International Curriculum Vitae Education B.S., Psychology, University of Washington, Seattle, Washington M.A., Psychology, State University of New York at Stony Brook, New York Ph.D., Psychology, State University of New York at Stony Brook, New York Academic Positions / Employment Clinical Instructor, Dept. of Psychia
“Product Management and Development for Insurance Companies” 4th – 6th June 2009 | Zagreb / Croatia organised by the European Actuarial Academy in cooperation with the Croatian Society of Actuaries. provisional programm 1. Introduction Product management in the European Insurance Industry is one of the most important – if not only – tools for insurance compani