Systemic Induction of Salicylates in Salix myrsinifolia(Salisb.)

We studied the induction of salicylates in mature leaves ofSalix myrsinifolia Salisb. (Salicaceae) following severe woundingby a specialist leaf-beetle Phratora vitellinae L. (Chrysomelidae).Levels of individual salicylates and aromatic amino acids andtheir total levels were determined in leaves at different developmentalstages. Induction of salicylates depended on: (1) the individualcompound; (2) the developmental stage of the plant organ; and(3) the genotype of the plant. Induction of salicylates wassystemic: levels of salicylates rose in unwounded young immatureand mature leaves, but no local response was detected in woundedleaves. In addition, there were clear clonal variations in boththe constant and induced levels of salicylates: clones withthe highest levels of salicylates were also most capable ofincreasing this level in response to herbivore attack i.e. notrade-off between constant and induced levels was detected.Furthermore, the levels of three aromatic amino acids, Phe,Tyr and Trp, increased in immature leaves of herbivore-affectedplants, which may indicate induction of enzymes of the shikimatepathway by wounding. The increase in salicylates was suggestedto be a consequence of an increased rate of synthesis ratherthan that of translocation. The induced levels of salicylatesdid not affect the subsequent feeding of highly specializedP. vitellinae. However, the ability to increase levels of salicylatesmay reduce grazing by generalist herbivores which are not ableto tolerate high levels of salicylates. Copyright 2001 Annalsof Botany Company Salix myrsinifolia, dark-leaved willow, Phratora vitellinae, salicylates, phenolic glycosides, herbivory, induced defence     

Impacts of nutrient enrichment and sediment on phytoplankton community structure in the northern Baltic Sea

A three-week mesocosm experiment was conducted in order to study the effects of bottom sediment and nutrient enrichment on phytoplankton and zooplankton community structure in the Archipelago Sea, northern Baltic Sea. The transparent polyethylene enclosures included the whole water column and varied in volume from 30 to 40 m³. There were two types of enclosures: some with natural sediment as a bottom and others with a plastic bottom. The experiment was a 2 × 2 factorial design with presence of sediment and nutrient enrichment as treatment factors. Both the sediment presence and nutrient enrichment significantly increased water nutrient concentrations and the rate of primary production. However, external nutrient enrichment and the presence of sediment stimulated the growth of different phytoplankton groups, indicating that the effect of sediment was not related to nutrient fluxes alone, but involved more complex interactions. External nutrient enrichment was primarily channelled to picoplanktonic cyanobacteria, the biomass of which increased four- to fivefold due to enrichment. The presence of sediment increased the biomass of cryptophytes, chrysophytes and prasinophytes, but decreased the biomass of N₂-fixing cyanobacteria. Zooplankton biomass increased during the experiment, but was not affected by the treatments. The study shows that sediment plays a significant role in phytoplankton dynamics, underlining the importance of including sediment in shallow-water mesocosm experiments. Handling editor: J. Padisak     

Shading enhances the quality of willow leaves to leaf beetles – but does it matter?

Plant shading is commonly recognised as a factor, which increases susceptibility of plants to attack by herbivorous insects. In this study we experimentally investigated the effect of host plant shading on two willow-feeding leaf beetles, Galerucella lineola feeding upon Salix phylicifolia and Phratora vitellinae feeding upon Salix myrsinifolia . Both beetle species were more abundant on potted willows growing in open habitats than on the same clones placed under the shade of trees. However, in the laboratory the food preference by adults and larval performance showed that the shaded willows are actually better food for both beetle species. On the contrary, when larvae were reared in the field under natural abiotic conditions, we found no difference in larval performance, or if any, even better performance in open habitats. Apparently, higher and more variable daily temperatures in open habitats accelerated the growth of the larvae. When adults were let to emigrate from or immigrate to potted willows, which had been grown in the same conditions but placed either in the open or shady habitats, adults preferred exposed willows. Invertebrate predators were more abundant in open habitats, but we found no differences in leaf beetle mortality by natural enemies between the habitats. Although the larval performance appeared to be approximately equal in the two habitats during the unusually warm study period, we suggest that under suboptimal temperatures the better abiotic conditions of open sites can easily override the better food provided by shaded habitats. The selection of abiotic habitat thus plays a significant role in the adaptive habitat and host plant selection of these beetles within the gradient of shadiness.     

Effect of nutrient enrichment on bacterioplankton biomass and community composition in mesocosms in the Archipelago Sea, northern Baltic

Effects of nutrient enrichment on the biomass and communitycomposition of heterotrophic bacteria and picocyanobacteriawere studied in large (42 m³) mesocosms in the brackish-waterArchipelago Sea (Baltic Sea) in late summer 2000 using cellcounts and denaturing gradient gel electrophoresis (DGGE) ofpolymerase chain reaction (PCR)-amplified 16S rRNA gene fragments.The identity of the major DNA bands was determined by sequencing.The obtained sequences were related to - and -proteobacteria,actinobacteria, verrucomicrobia and cyanobacteria. Nitrogenand phosphorus additions increased the biomasses of heterotrophicbacteria and picocyanobacteria and caused significant changesin their community composition judging from the DGGE bandingpatterns. Most verrucomicrobial bands had their highest relativeintensity in the control treatment and their lowest in the highernutrient addition treatment, whereas most Synechococcus-relatedbands had their lowest relative intensity in the lower nutrientaddition treatment. The responses of proteobacteria and actinobacteriawere more variable. The presence of both freshwater and marinesequences among the closest relatives to our sequences highlightsthe intermediate character of the Archipelago Sea between afreshwater and truly marine environment.     

Evidence for host race formation in the leaf beetle Galerucella lineola

We examined preference and performance of four Finnish Galerucella lineola F. populations on alder and willow. In standardized two‐choice laboratory feeding trials with alder and willow, only two naturally alder‐associated G. lineola populations accepted alder. Two conspecific willow‐associated populations preferred willow. These preferences seem to be unstable, however, because they can be modified by the beetles’ experience. Thus, there probably is not a complete host preference‐based isolation of alder‐ and willow‐associated G. lineola beetles in nature. In performance experiments, larvae of all four populations survived better on willow than on alder. This may indicate that willows are the ancestral hosts for G. lineola. Nevertheless, larvae of the two alder‐associated G. lineola populations survived better on alder than larvae of the two willow‐associated populations. On the other hand, larvae of the two willow‐associated populations survived better on willow than larvae of the two alder‐associated populations. This performance trade‐off suggests that G. lineola encounters different selective pressures on alders and willows. On both of them, selection probably disfavours those G. lineola genotypes that are the most successful and abundant on alternative hosts. This may reduce the effects of gene flow that is likely to occur as a consequence of incomplete host preference‐based isolation of alder‐ and willow‐associated G. lineola populations. Data from pupal weights support the idea that alder‐ and willow‐associated G. lineola populations may be genetically differentiated. Pupae of the two alder‐associated populations were heavier than those of the willow‐associated populations irrespective of whether larvae had fed on alder or on willow. Overall, our results indicate host race formation in G. lineola. This process may be enforced by the variable abundance of alders and willows in local communities.     

Species-specific differences in phytoplankton responses to N and P enrichments and the N:P ratio in the Archipelago Sea, northern Baltic Sea

A nutrient enrichment experiment was conducted in order to studythe role of nitrogen (N), phosphorus (P) and the N:P ratio onthe early summer phytoplankton community in the ArchipelagoSea, northern Baltic Sea. The phytoplankton community was, interms of chlorophyll a and total biomass, primarily N-limited,but the individual species varied in their responses to thenutrient supply. The recorded overall N limitation was due tofast growth responses of a few N-limited species such as thediatom Chaetoceros wighamii (Brightwell) and the mixotrophicchrysophyte Uroglena sp. Another dominating diatom, Skeletonemacostatum (Greville) Cleve was most clearly P-limited. The N:Pratio had the strongest effect on Uroglena sp., which grew exponentiallyin the enrichments with a high N:P ratio. This can be explainedby the ability of the species to feed on P-rich bacteria, whichgives it a competitive advantage in P-limited conditions. Thespecies-specific differences in the responses to the nutrientenrichments can generally be explained by differences in thespecies physiology and they were consistent with the theoryof resource competition.     

Tritrophic interactions: willows, herbivorous insects and insectivorous birds

Insectivorous birds can increase plant growth by consuming herbivorous insects and reducing insect damage. However, plant traits such as the level of chemical defense may affect the quantity and quality of insects, and alter the foraging behavior of birds. Therefore, I predicted that plant traits can also modify the effect of birds on leaf damage and plant growth. This study compared the effect of insectivorous birds on the herbivory and growth of two chemically different willow species, weakly defended Salix phylicifolia and strongly defended S. myrsinifolia under two fertilization levels. Half of the willows were protected from birds using a translucent gill-net, which did not limit access by insects. The effect of birds on the densities of leaf-chewing insects and leaf damage was considerable on unfertilized S. phylicifolia but less obvious on fertilized ones. The effect of bird predation was negligible on S. myrsinifolia, which had very low insect densities in all treatments. Birds increased the growth of the experimental willows, but the effect was clear only in unfertilized S. phylicifolia. I suggest that birds avoided foraging on willows with low populations of insects and little visible damage. The study shows that bird predation can alter the patterns of insect densities we see on willows, emphasizing the importance of considering multitrophic effects when studying plant-insect interactions. Received: 25 May 1999 / Accepted: 9 August 1999     

Interactions between willows and insect herbivores under enhanced ultraviolet-B radiation

We studied the effects of elevated ultraviolet-B radiation on interactions between insect herbivores and their host plants by exposing two species of phytochemically different willows, Salix myrsinifolia and S. phylicifolia, to a modulated increase in ultraviolet radiation in an outdoor experiment and monitoring the colonisation of insect herbivores on these willows. We examined the effect of increased ultraviolet-B (UV-B) radiation on (1) the quality of willow leaves, (2) the distribution and abundance of insect herbivores feeding on these willows, (3) the resulting amount of damage, and (4) the performance of insect larvae feeding on the exposed plant tissue. Six clones of each of the two willow species were grown in eight blocks for 12 weeks in the UV-B irradiation field. The clones were exposed to a constant 50% increase in UV-B radiation (simulating 20-25% ozone depletion), to a small increase in UV-A radiation or to ambient solar irradiation. We allowed colonisation on the willows by naturally occurring insects, but also introduced adults of a leaf beetle, Phratora vitellinae, a specialist herbivore on S. myrsinifolia. Increased UV-B radiation did not affect any of the measured indices of plant quality. However, numbers of P. vitellinae on S. myrsinifolia were higher in plants with UV-B treatment compared with UV-A and shade controls. In laboratory tests, growth of the second-instar larva of P. vitellinae was not affected by UV-B treatment of S. myrsinifolia, but was retarded on UV-B treated leaves of S. phylicifolia. In addition, naturally occurring insect herbivores were more abundant on willows exposed to elevated UV-B radiation compared to those grown under control treatments. In spite of the increased abundance of insect herbivores, willows treated with elevated UV-B did not suffer more herbivore damage than willows exposed to ambient solar radiation (shade control). The observed effects of UV-B on herbivore abundance, feeding and growth varied significantly due to spatial variation in environment quality, as indicated by the UV-treatment x block interaction. The results suggest that (1) environmental variation modifies the effects of UV-B radiation on plant-insect interactions and (2) specialist herbivores might be more sensitive to chemical changes in their secondary host plants (S. phylicifolia) than to changes in their primary hosts (S. myrsinifolia).