Recent theoretical work (Vos et al. 2004) predicts that inducible defences prevent strong population fluctuations under high levels of nutrient enrichment. Here we evaluate this model prediction and show that inducible defences in algae stabilize the dynamics of experimentally assembled bi-and tritrophic planktonic food chains. At high phosphorus levels, we observed strong population fluctuations in all food chains with undefended algae. These fluctuations did not occur when algae had inducible defences. At low phosphorus levels, we observed deterministic consumer extinctions, as predicted by stoichiometric theory. Our study thus shows that both biotically and abiotically induced changes in algal food quality affect the stability and persistence of planktonic food chains.

During the upsurge of the introduced predatory Nile perch in Lake Victoria in the 1980s, the zooplanktivorous Haplochromis (Yssichromis) pyrrhocephalus nearly vanished. The species recovered coincident with the intense fishing of Nile perch in the 1990s, when water clarity and dissolved oxygen levels had decreased dramatically due to increased eutrophication. In response to the hypoxic conditions, total gill surface in resurgent H. pyrrhocephalus increased by 64%. Remarkably, head length, eye length, and head volume decreased in size, whereas cheek depth increased. Reductions in eye size and depth of the rostral part of the musculus sternohyoideus, and reallocation of space between the opercular and suspensorial compartments of the head may have permitted accommodation of larger gills in a smaller head. By contrast, the musculus levator posterior, located dorsal to the gills, increased in depth. This probably reflects an adaptive response to the larger and tougher prey types in the diet of resurgent H. pyrrhocephalus. These striking morphological changes over a time span of only two decades could be the combined result of phenotypic plasticity and genetic change and may have fostered recovery of this species.

We studied the occurrence of colony formation within 40 different strains of Scenedesmaceae (Chlorococcales, Chlorophyta) in response to grazing-released infochemicals from the herbivorous zooplankters Brachionus calyciflorus Pallas (Rotifera) and Daphnia magna Strauss (Cladocera). With the exception of two strains, all strains showed similar responses to both B. calyciflorus and D. magna infochemicals, either no response or inducible colony formation. Colony size was found to increase with B. calyciflorus infochemical concentration and could be described by a sigmoid function. The increase in colony size was more pronounced in the Scenedesmus species tested than in Desmodesmus species, which was probably due to higher threshold infochemical concentrations for colony induction in Desmodesmus. Therefore, the adaptivity of colony formation to the herbivory threat only holds above the threshold concentration for colony induction and as long as maximum colony size has not been attained. Taking this into account, our results suggest that inducible colony formation is a common adaptive response of many Scenedesmaceae to the threat of herbivory.

Abstract. Inducible defenses are dynamic traits that modulate the strength of both plantherbivore and herbivore-carnivore interactions. Surprisingly few studies have considered the relative contributions of induced plant and herbivore defenses to the overall balance of bottom-up and top-down control. Here we compare trophic cascade strengths using replicated two-level and three-level plankton communities in which we systematically varied the presence or absence of induced defenses at the plant and/or herbivore levels. Our results show that a trophic cascade, i.e., significantly higher plant biomass in three-level than in two-level food chains, occurred whenever herbivores were undefended against carnivores. Trophic cascades did not occur when herbivores exhibited an induced defense. This pattern was obtained irrespective of the presence or absence of induced defenses at the plant level. We thus found that herbivore defenses, not plant defenses, had an overriding effect on cascade strength. We discuss these results in relation to variation in cascade strengths in natural communities.

1. Theoretical studies have shown that inducible defences have the potential to affect population stability and persistence in bi-and tritrophic food chains. Experimental studies on such effects of prey defence strategies on the dynamics of predator-prey systems are still rare. We performed replicated population dynamics experiments using the herbivorous rotifer Brachionus calyciflorus and four strains of closely related algae that show different defence responses to this herbivore. 2. We observed herbivore populations to fluctuate at a higher frequency when feeding on small undefended algae. During these fluctuations minimum rotifer densities remained sufficiently high to ensure population persistence in all the replicates. The initial growth of rotifer populations in this treatment coincided with a sharp drop in algal density. Such a suppression of algae by herbivores was not observed in the other treatments, where algae were larger due to induced or permanent defences. In these treatments we observed rotifer population densities to first rise and then decline. The herbivore went extinct in all replicates with large permanently defended algae. The frequency of herbivore extinctions was intermediate when algae had inducible defences. 3. A variety of alternative mechanisms could explain differential herbivore persistence in the different defence treatments. Our analysis showed the density and fraction of highly edible algal particles to better explain herbivore persistence and extinctions than total algal density, the fraction of highly inedible food particles or the accumulation of herbivore waste products or autotoxins. 4. We argue that the rotifers require a minimum fraction and density of edible food particles for maintenance and reproduction. We conjecture that induced defences in algae may thus favour larger zooplankton species such as Daphnia spp. that are less sensitive to shifts in their food size spectrum, relative to smaller zooplankton species, such as rotifers and in this way contributes to the structuring of planktonic communities.