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Courtship Behaviour in Drosophila: Sexual Isolation or Sexual Selection?

Courtship Behaviour in Drosophila: Sexual Isolation or Sexual Selection?

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Published by Maurice Dow
Authors: Florian von Schilcher & Maurice A. Dow.
Z. Tierpsychol 43: 304-310, 1977.
Abstract:
The theory of sexual selection and isolation lays heavy emphasis on the role of the female - defined as the sex with the higher parental investment - in these activities. The male investment is, however, not negligible and we hypothesize that, under certain, well defined conditions, sexual isolation will be a function of male behaviour, whereas sexual selection will always mainly be a female prerogative.
We present behavioural data from observations of intra- and interspecific single pair matings of 5 sibling species of the Drosophila melanogaster group - D. melanogaster, D. simulans,
D. mauritiana, D. teissieri, D. yakuba.
The results are consistent with our hypothesis. Males are primarily responsible for sexual isolation, whereas females dominate sexual selection.
Authors: Florian von Schilcher & Maurice A. Dow.
Z. Tierpsychol 43: 304-310, 1977.
Abstract:
The theory of sexual selection and isolation lays heavy emphasis on the role of the female - defined as the sex with the higher parental investment - in these activities. The male investment is, however, not negligible and we hypothesize that, under certain, well defined conditions, sexual isolation will be a function of male behaviour, whereas sexual selection will always mainly be a female prerogative.
We present behavioural data from observations of intra- and interspecific single pair matings of 5 sibling species of the Drosophila melanogaster group - D. melanogaster, D. simulans,
D. mauritiana, D. teissieri, D. yakuba.
The results are consistent with our hypothesis. Males are primarily responsible for sexual isolation, whereas females dominate sexual selection.

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Published by: Maurice Dow on Feb 23, 2013
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Z.
Tierpsychol.,
43,
304-310 (1977)
@
1977 Verlag Paul Parey, Berlin und Hamburg
ISSN
0044-3573
I
ASTM-Coden: ZETIAG
Department
of
Zoology, University
of
Edinburgh, Scotland
Courtship Behaviour in
Drosophila:
Sexual Isolation
or
Sexual Selection?
By
FLORIAN
ON
SCHILCHER
nd
MAURICE ow
With one figureReceived: September
3,1976
Accepted: October
27, 1976
Abstract
The theory
of
sexual selection and isolation lays heavy emphasis on the role
of
the
9
-
efined as the sex with the higher parental investment
-
n these activities. The male in-vestment is, however, not negligible and we hypothesize that, under certain, well definedconditions, sexual isolation will be a function of male behaviour, whereas sexual selection willalways mainly be a female prerogative.We present behavioural data from observations of intra- and interspecific single pairmatings
of
5 sibling species of the
Drosophila melanogaster
group
-
. melanogaster, D.simu-
lans,
D. mauritiana, D. teissieri,
D.
yakuba.
The results are consistent with our hypothesis.
8
8
are primarily responsible for sexualisolation, whereas
99
dominate sexual selection.
Sexual isolation and sexual selectionare two of the salient functionsproviding the selective forces which mould courtship behaviour
(BASTOCK
1967;
GHISELIN
974). According to one prominent theory of speciation, twopreviously isolated populations, which come into contact, will evolve reproduc-tive isolating mechanisms if their hybrids have lower fitness than offspringresulting from crosses within the populations. Initially the barriers will bemostly postmating, but as these are rather inefficient, premating isolatingmechanisms will be selected for
(MAYR
1963). Sexual selection in the Dar-winian sense within one species remains
a
hotly debated issue in theoreticalbiology since
DARWIN
roposed it as an evolutionary mechanism differentfrom natural selection
(CAMPBELL
972). Empirical support for the existenceof sexual selection within and sexual isolation between species comes fromnumerous examples of its action in diverse species of animals
(WILSON
976).However, these observations leave the problem of the justification
of
DAR-
WIN’S
distinction untouched.In the choice of a mating partner
-
ithin and between species
-
he
9
should generally be more critical than the
8
(BATEMAN
948;
TRIVERS
972).In the following we report and discuss results which relate to these issues.They were obtained in intra- and interspecific matings with
Drosophila
me-
lanogaster
and its sibling species.
 
Courtship Behaviour in
Drosophila:
Sexual Isolation or Sexual Selection
305The courtship behaviour of
D.
melanogaster
and
D.
simulans
has beenwell described
(MANNING
959a). The
8
orients to the
0,
taps her with
his
front legs, extends one wing 90' and vibrates it, licks her genitalia andattempts to copulate.
Drosophila
??
can show
a
number of rejection move-ments, like kicking with their hindlegs, wingflicking and extrusion of theovipositor
(CONNOLLY
nd
COOK
973). Among the sensory channels involvedin
Drosophila
courtship, auditory channels
(BENNET-CLARK
nd
EWING
967),and air-borne chemical
(SHOREY
nd
BARTELL
970) and contact chemical ones
(MANNING
959 b), are outstanding in their importance for successful1 copula-tion. The latter have been shown to function in species isolation
(MANNING
1959 b), whereas airborne chemical channels play a role in intraspecific sexualselection
(EHRMAN
969; AVERHOFFnd
RICHARDSON
976). The function ofthe auditory information given by courting
6
8
to the
??
remains an un-resolved problem as far as the above discussed dichotomy is concerned
(SCHIL-
CHER
1976a).Recent experiments
on
the sensory basis of the rare type mating advan-tage in
D. melanogaster
have been interpreted by the authors to constituteevidence for an auditory control
of
this kind of intraspecific sexual selection
(PETIT
and
hTouAuD
1975). The evidence, however, rests heavily
on
theassumption that
??
can perceive auditory signals from wingless
8
6.
Thereis not much support for this contention
(SCHILCHER
976b) and the resultsmight be due to
an
unspecific effect of aristae amputation
on
female dis-criminatory activity.There are now5known sibling species of
D. melanogaster
-
. simulans,
D.
erecta,
D.
mauritiana, D. teissieri
and
D.
yakuba
(TSACAS
ers. comm.).
D. melanogaster
and
D. simulans
are cosmopolitan
(PATTERSON
nd
STONE
1952) and are therefore sympatric with all the other species.
Materials and Methods
Stocks were cultured on standard cornmeal-molasses medium, seeded with live yeast, at
25fZOC
and under
12
:
12
LD
conditions. Pairs
of
3
to
4
days old flies were introducedwithout etherization into cylindrical perspex arenas (2.5 cm in diameter,
1.1
cm high) withcentral partitions separating the sexes. After removal of the partition, courtship interactionswere observed with
a
binocular microscope. We recorded the duration
of
the first courtshipbout, the male and female sexual behaviour patterns occurring during the interaction and thesex which terminated the bout. The durations
of
the first bouts were dichotomized
as
less thancr equal to
1
s,
and greater than
1
.
The start of courtship was defined
as
the first orientation
of
the
8
to
the while within one bodylength of her. The bout was scored
as
terminatedif the
8
was no longer oriented or was preening. The bout was ?-terminated if she flewaway or rejected the
8
ust before the end
of
it. The
8
ended the courtship if he turned away,did not follow the
9
or preened, when these behaviours were not preceded by
a
visible femalerejection.
D.
melanogaster
and
D. simulans-
8
8
were paired with
99
of
all
6
species.
As
therewere no significant differences between the two species
of
8
8
in their behaviour towardsconspecific
99,
he results were pooled. The results from the interspecific crosses were pooled
on
the same grounds.
Results
As
can be seen in Table
1,
8
6
more often terminated interspecific court-ship bouts and
??
terminated more intraspecific bouts
(x'
=
54.95, df
=
1,
p
<
0.001).
Table
1
also shows that intraspecific courtship interactions lastedlonger than interspecific ones
(x
=
36.14, df
=
1,
p
<
O.OO1).
Table
2
showsthat the frequency of the different male behaviours depends
on
whether the
0
is conspecific or not
(x2
=
53.6,
df
=
3,
p
<
0.001).
Partitioning the chi-
Z.
Tierpsvchol..
Bd.
43.
Heft
3
20
 
306
FLORIAN
ON
SCHILCHER
nd
MAURICE
ov
terminating
sex
Table
f;
The number
of
courtship interactionswhich were terminated by each sex and their duration
durationintraspec ificinterspecif ic
18
3
1
30
4
64 42 26
Discussion
courtshipintraspecificinterspecific
Since
6
6
more frequently end interspecific encounters, and as these are
also
much shorter than intraspecific ones, we can conclude that in the formerthe
8
6
are not highly motivated.
As
most of the interspecific courtships areterminated after the first orientation and/or tap, and as the sequence orienta-tion-vibration-licking-attempted copulation has been interpreted asindicative of increasing male sexual excitation (BASTOCKnd MANNING
955),
this is further evidence for the
6
8
ow state of sexual arousal in encounterswith heterospecific
99.
Contact
-
nd/or airborne chemical factors will mostprobably be responsible for this inale discrimination of heterospecific
99.
If
one hypothesises that the
99
switch off the interspecific courtship by somemeans which we cannot readily perceive, then one has to explain why they act
so
conspicuously when ending intraspecific interactions. Especially since inthe former the selective disadvantage conferred by
a
forced mating would bemuch more serious. In rare instances we did observe long and rather intensiveinterspecific interactions and as
99
do occasionally terminate these, the maledisplays might still be used as a fail-safemechanism for sexual isolation;however, the selective forces maintaining this mechanism might come from therequirements
of
sexual selection within the species.
It
therefore represents aselective epiphenomenon or a preadaptation rather than a true adaptation.Our observations clearly demonstrate that the choice of a mating partnerwithin the species is a female prerogative
to
a large extent
at
least.The theory according to which speciation is a consequence of geographicisolation, predicts that isolating mechanism will be more efficient, the earlierin the sequence
of
epigamic interactions between the sexes they act (MAYR
1963).
The parental investment theory predicts that the
99
(the sex with thehigher parental investment) should be the principal factor which maintainssexual isolation. In all cases where the
8
(the sex with the lower parentalinvestment) initiates courtship through a physical approach to the
0,
the two
Behavior
0
T
v
L
AC
1
3
8
27
*
19
30
1s
4

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