Biological Journal of the Linnean Society , 2003, 78 , 149–154. With 1 figure
© 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 78 , 149–154 149
Blackwell Science, LtdOxford, UKBIJBiological Journal of the Linnean Society0024-4066The Linnean Society of London, 2003
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Original Article
SPERM COST IN A HERMAPHRODITE POLYCHAETE WORMG. SELLA and M. C. LORENZI
*Corresponding author. E-mail: gabriella.sella@unito.it
Increased sperm allocation delays body growth in a
protandrous simultaneous hermaphrodite
GABRIELLA SELLA* and MARIA CRISTINA LORENZI
Dipartimento di Biologia Animale e dell’Uomo, Università di Torino, Italy
Received 31 July, 2001; accepted for publication 31 October, 2002
Sex allocation theory for simultaneous hermaphrodites predicts that hermaphrodites have a fixed amount of repro-
ductive resources to allocate to both sex functions and that they trade off their allocation to both sexes depending on
environmental conditions. Ophryotrocha diadema is a simultaneous hermaphroditic polychaete worm which has a
protandrous phase prior to the hermaphroditic phase. The ability of adolescent males to compete with mature her-
maphrodites for egg fertilization and the costs of an increase in male expenditure during the protandrous phase were
tested in experiments where adolescent males had the possibility to fertilize hermaphrodites’ eggs. Results document
that adolescent males were able to compete with hermaphrodites for egg fertilization and caused the hermaphrodites
fitness losses of 31.4%. Adolescent males which fertilized hermaphrodites’ eggs had a slower growth rate than males
which had no possibility to fertilize eggs. This result indicates that an increased male expenditure is a cost for ado-
lescent males and that, during the protandrous phase, resources are allocated to reproduction at the expense of
somatic development. © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 ,
149–154.
ADDITIONAL KEYWORDS: male expenditure – Ophryotrocha diadema – Polychaeta – protandry – simulta-
neous hermaphroditism – sperm cost – trade-off.
INTRODUCTION
Most research on simultaneous hermaphrodite
resource allocation is focused on how sex allocation is
partitioned between male and female functions: sex
allocation theory predicts that hermaphrodites have
a fixed amount of reproductive resources to allocate
to both sexual functions; moreover, they allocate
resources to either sex function based on environmen-
tal conditions (Charnov, 1982; Fischer, 1984; Angeloni,
Bradbury & Charnov, 2002). Theory predicts that
within such a fixed amount, allocation to male func-
tion (which may be very costly, Dewsbury, 1982; Greeff
& Michiels, 1999; Wedell, Gage & Parker, 2002)
increases as mating group size increases. Empirical
evidence from Raimondi & Martin (1991) supports
this statement: barnacles allocate proportionally more
of their reproductive resources to female function
when they occur in small mating groups than when
they are in large ones. Similar findings were described
by Trouvé et al . (1999) and by Schärer & Wedekind
(2001) in platyhelminth parasites.
Little is known about other life-history trade-offs in
hermaphrodites. Reproductive investment may limit
other life functions. For example, increased sperm
production reduces lifespan in Caenorhabditis ele-
gans (Van Voorhies, 1992) and may also lead to a
delay of the onset of oogenesis and hence to an
increase in minimum generation time and a decrease
in population growth (Hodgkin & Barnes, 1991;
Barker, 1992). In the colonial ascydian Botryllus
schlosseri , a cyclical hermaphrodite with sexual and
asexual reproduction, a negative correlation has been
demonstrated between asexual growth and sexual
investment (Yund, Marcum & Stewart-Sevage, 1997).
A trade-off between growth and male reproductive
success was also shown in two gonochoric fish species
(reviewed by Taborsky, 1994).
Evidence of a relationship between reproduction
and growth rate in hermaphrodites is lacking. De
Visser, Ter Maat & Zonneveld (1994) showed that
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