© 2008 The Authors Entomologia Experimentalis et Applicata 128: 109–116, 2008
Journal compilation © 2008 The Netherlands Entomological Society 109
DOI: 10.1111/j.1570-7458.2008.00695.x
Blackwell Publishing Ltd
Differential effects of jasmonic acid treatment of
Brassica nigra on the attraction of pollinators,
parasitoids, and butterflies
Maaike Bruinsma*, Harm IJdema, Joop J.A. van Loon & Marcel Dicke
Laboratory of Entomology, Wageningen University, PO Box 8031, 6700 EH Wageningen, The Netherlands
Accepted: 13 February 2008
Key words: Apis mellifera, syrphid flies, Cotesia glomerata, Pieris rapae, host plant selection, flower
visitors, octadecanoid pathway, pollination, Lepidoptera, Pieridae, Hymenoptera, Braconidae
Abstract Herbivore-induced plant defences influence the behaviour of herbivores as well as that of their
natural enemies. Jasmonic acid is one of the key hormones involved in both these direct and indirect
induced defences. Jasmonic acid treatment of plants changes the composition of defence chemicals
in the plants, induces volatile emission, and increases the production of extrafloral nectar. However,
few studies have addressed the potential influence of induced defences on flower nectar chemistry and
pollinator behaviour. These have shown that herbivore damage can affect pollination rates and plant
fitness. Here, we have investigated the effect of jasmonic acid treatment on floral nectar production
and the attraction of pollinators, as well as the effect on the behaviour of an herbivore and its natural
enemy. The study system consisted of black mustard plants, Brassica nigra L. (Brassicaceae), pollinators
of Brassica nigra (i.e., honeybees and syrphid flies), a specialist herbivore, Pieris rapae L. (Lepidoptera:
Pieridae), and a parasitoid wasp that uses Pieris larvae as hosts, Cotesia glomerata L. (Hymenoptera:
Braconidae). We show that different trophic levels are differentially affected by jasmonic acid-induced
changes. While the herbivore prefers control leaves over jasmonic acid-treated leaves for oviposition,
the parasitoid C. glomerata is more attracted to jasmonic acid-treated plants than to control plants.
We did not observe differences in pollinator preference, the rates of flower visitation by honey-
bees and syrphid flies were similar for control and jasmonic acid-treated plants. Plants treated with
jasmonic acid secreted less nectar than control plants and the concentrations of glucose and fructose
tended to be lower than in nectar from control plants. Jasmonic acid treatment resulted in a lower
nectar production than actual feeding damage by P. rapae caterpillars.
Introduction
Induction of defence responses in plants can alter the
behaviour of associated insects. This is well studied for
foliar herbivores (folivores) and their natural enemies
associated with vegetative plants (e.g., Dicke et al., 1990;
Turlings et al., 1990; Shiojiri et al., 2002; Bruinsma & Dicke
2008). However, the effects of induced defences on
flowering plants, and consequently pollinator behaviour,
have received much less attention. Root and foliar
herbivory may indirectly affect plant fitness by reducing
resources for reproduction through reduction of root
volume or photosynthetic area. Floral herbivory (florivory)
directly affects plant fitness by reducing the number of
gametes (Poveda et al., 2003). Observed effects of foliar
herbivory on flowering plants include a decrease in
pollinator visitation, an increase in secondary metabolites
in leaves, flowers, or nectar, fewer and smaller flowers,
and decreased pollen production (Strauss et al., 1996,
2004; Lehtilä & Strauss, 1997; Mothershead & Marquis,
2000; Hambäck, 2001; Smallegange et al., 2007). Root
herbivory can increase pollinator visitation (Poveda et al.,
2003, 2005) and florivory can decrease the number of flowers,
nectar production, seed set, and pollinator visitation
(Krupnick et al., 1999; Adler et al., 2001).
Although these examples show that herbivory can
increase or decrease pollinator visitation, depending on type
of damage, plant and pollinator species, the mechanisms * Correspondence: E-mail: maaike.bruinsma@wur.nl