Abundance and diversity of the phyllosphere bacterial communities of Mediterranean perennial plants that differ in leaf chemistry

Summary.  We studied the epiphytic bacterial communities of the summer leaves of eight perennial species naturally occurring in a Mediterranean ecosystem. The species differ in essential-oil content (from rich in essential oil to non-producers) and composition, and also in life form (from herbaceous species to tall shrubs). We compared the epiphytic bacterial communities on the basis of (i) their abundance, (ii) their metabolic profile (derived by use of the BIOLOG Ecoplate system) and (iii) richness and diversity of substrates that they use, as a measure of functional diversity. Among all species, the aromatic Melissa officinalis was the most abundantly colonized. The bacterial communities on the leaves of the aromatic Myrtus communis, Calamintha nepeta and Melissa officinalis, and also of Cistus incanus catabolized all 31 substrates offered; those on the evergreen-sclerophyllous species, Arbutus unedo and Quercus coccifera, catabolized only 14 and 17 substrates, respectively. Carbohydrates were consistently used abundantly by all communities, whereas carboxylic acids were most variably used. On average, the group of aromatic plants scored higher regarding bacterial abundance, and richness and diversity of substrates used by the bacterial communities on their leaves; the lowest values for both substrate-use indices were recorded in A. unedo. Bacterial abundance or richness or diversity of substrates used did not vary with leaf oil content. Abundance was positively correlated with both substrate-use indices. Results support claims that the antimicrobial effects of essential oils are not exerted so much under natural conditions as reports based on biassays with pathogens usually show. Although essential oils play a part in the microbial colonization of the phyllosphere, it is not likely that inhibition of phyllosphere bacteria is essential oils’ primary role, at least in the Mediterranean environment.     

Preferences of six leaf beetle species among qualitatively different leaf age classes of three Salicaceous host species

Summary. As Salicaceous plants produce new leaves for a prolonged period of time, they expose a wide range of differentially aged leaves to herbivores during the growing season. In this work, I show that young leaves of three Salicaceous species, Populus tremula L., Salix phylicifolia L. and S. pentandra L., contain more nitrogen than conspecific old leaves. In P. tremula and S. pentandra young leaves also contained more low-molecular weight secondary compounds, phenolic glucosides. Leaves of S. phylicifolia did not contain phenolic glucosides in detectable amounts. Furthermore, in P. tremula and S. pentandra young leaves contained less polymeric digestability-reducing phenolics, condensed tannins, than old leaves. In S. phylicifolia, higher concentrations of condensed tannins were found in young leaves. In laboratory feeding trials with six leaf beetle species, young leaves of the studied plants were invariably preferred in all tested herbivore × host species combinations. In particular, it is remarkable that three leaf beetle species with known different overall relationships to phenolic glucosides equally preferred more glucoside-containing young S. pentandra leaves over conspecific old ones. Four beetle species were found to prefer young leaves of S. phylicifolia despite the higher content of condensed tannins in young leaves. These results indicate that the general preference of leaf beetles for young leaves of Salicaceous plants probably does not primarily result from variable distribution of secondary compounds. Apparently, the preference for young leaves is fundamentally due to variation in leaf nutritive traits, such as nitrogen content. Received 9 February 2001.     

Host-plant leaf surface compounds influencing oviposition in Delia antiqua

Summary. Delia antiqua (Diptera: Anthomyiidae) females lay eggs between the leaves of onion plants or in the soil around the base of the plants, then the maggots feed on the onion bulb and roots causing rapid secondary infection by fungi and bacteria. It is well known that the first sensory modality used by the onion fly is vision, therefore the shape (vertical narrow cylinders) and colour (yellow) of the plant play a crucial role in the recognition of a potential host plant. In the past it has been shown that n-dipropyl disulfide (Pr₂S₂), a typical component of onion volatiles, is an important chemical host plant cue. We extracted host leaf surface to verify if Pr₂S₂ is the major chemical oviposition stimulant and to determine if other as yet unknown substances may play a role in host-plant selection. We confirmed that the females laid more eggs around onion plants with leaves than when only the onion bulb was present and that the odour of chopped onion stimulates oviposition. Extraction of the surface of onion leaves revealed that only the apolar fraction contained substances that stimulate egg-laying in D. antiqua. GC-EAD analysis indicated that a minor constituent, Pr₂S₂, is perceived by the olfactory receptor on the antennae of the onion fly females. This confirmed the importance of Pr₂S₂ as oviposition stimulant. Contact with the polar fraction did not stimulate egg-laying behaviour in this Delia species. We discuss the oviposition strategy of D. antiqua in comparison with its closely related species, D. radicum, in which the oviposition behaviour is stimulated mainly through contact with the cabbage leaf surface and only partially by the host volatiles.     

Pyrrolizidine alkaloids and pentacyclic triterpene saponins in the defensive secretions of Platyphora leaf beetles

Summary. Field collected exocrine defensive secretions of nine neotropical Platyphora species were analyzed for the presence of plant acquired pyrrolizidine alkaloids (PAs) and pentacyclic triterpene saponins. All species secrete saponins. In addition, five species feeding on Tournefortia (Boraginaceae), Koanophyllon (Asteraceae, tribe Eupatorieae) and Prestonia (Apocynaceae) were shown to sequester PAs of the lycopsamine type, which are characteristic for species of the three plant families. The PA sequestering species commonly store intermedine, lycopsamine and their O^3′-acetyl or propionyl esters as well as O⁷- and O⁹-hydroxyisovaleryl esters of retronecine. The latter as well as the O^3′-acyl esters were not found in the beetles’ host plants, suggesting the ability of the beetles to esterify plant derived retronecine and intermedine or its stereoisomers. Despite the conformity of the beetles’ PA patterns, considerable inconsistencies exist regarding the PA patterns of the respective host plants. One host plant was devoid of PAs, while another contained only simple necines. Since the previous history of the field collected beetles was unknown this discrepancy remains obscure. In contrast to the Palearctic chrysomeline leaf beetles, e.g. some Oreina species which ingest and store PAs as their non-toxic N-oxides, Platyphora leaf beetles absorb and store PAs as the toxic free base (tertiary PA), but apparently avoid to accumulate PAs in the haemolymph. This suggests that Chrysolina and Platyphora leaf beetles developed different lines of adaptations in their parallel evolution of PA mediated chemical defense. Received 30 November 2000; accepted 5 February 2001     

First-instar monarch larval growth and survival on milkweeds in southern California: effects of latex, leaf hairs and cardenolides

Growth rate and survival of first-instar larvae of Danaus plexippus, a milkweed specialist, depended on milkweed species, and was related to the amount of latex produced from wounds, leaf cardenolide concentrations and the presence of leaf hairs. Larval growth was more rapid and survival was higher on leaves of Asclepias californica with experimentally reduced latex, and this species has characteristically high latex, low- to mid-range cardenolide concentrations, and very hirsute leaves. Similarly, growth was higher on reduced latex leaves of both A. eriocarpa (a high latex/high cardenolide, hirsute species) and A. erosa (glabrous fleshy leaves, high latex/high cardenolides). There were no differences in either survival or growth rate between larvae on reduced latex or control leaves of the low latex/low cardenolide A. fascicularis with soft glabrous leaves and both survival and growth rate were higher on this species than the other species tested. Larval growth rates on leaves with reduced latex were similar among ten milkweed species tested to date but differed from growth rates on intact leaves suggesting that latex and possibly included cardenolides are both important in first-instar monarch larval growth, development and survival. We show for a range of ecologically important milkweeds that experiments on cut plant material (no latex outflow) lead to higher growth rates compared to intact plants. Such laboratory assays based on detached leaves will be misleading if the objective is to determine the impact of treatments such as Bt-maize pollen on monarchs on field plants.     

Variation in composition of predator-attracting allelochemicals emitted by herbivore-infested plants: Relative influence of plant and herbivore

Summary During foraging, natural enemies of herbivores may employ volatile allelochemicals that originate from an interaction of the herbivore and its host plant. The composition of allelochemical blends emitted by herbivore-infested plants is known to be affected by both the herbivore and the plant. Our chemical data add new evidence to the recent notion that the plants are more important than the herbivore in affecting the composition of the volatile blends. Blends emitted by apple leaves infested with spider mites of 2 different species,T. urticae andP. ulmi, differed less in composition (principally quantitative differences for some compounds) than blends emitted by leaves of two apple cultivars infested by the same spider-mite species,T. urticae (many quantitative and a few qualitative differences). Comparison between three plant species — apple, cucumber and Lima bean — reveals even larger differences between volatile blends emitted upon spider-mite damage (many quantitative differences and several qualitative differences).     

A field colonization experiment with meiofauna and seagrass mimics: effect of time,distance and leaf surface area

From a conservation point of view, it is essential to know how fast an ecosystem can recover after physical disturbance. Meiofauna and especially harpacticoid copepods are abundant in seagrass beds and are therefore useful to study ecosystem recovery after disturbance. In the western Caribbean coast, a fragmented Thalassia testudinum seagrass bed was selected to conduct a colonization field experiment by means of plastic seagrass mimics. Meiofauna colonization, with special emphasis on harpacticoid copepods, was followed in relation to: (1) colonization time (2, 4, 6, 10, 14 and 21 days); (2) distance to source of colonizers (close and far series) and (3) leaf surface area to colonize (small, medium, large). Colonization was recorded after 2 days with average meiofauna densities of 480 ind/100 cm² (close) and 1350 ind/100 cm² (far) of leaf surface area, while on average 400 ind/100 cm² were collected from the natural seagrass plants. In this early phase, the meiofauna diversity was high, with on average 8 taxa. A longer period of colonization (21 days) showed an increased meiofaunal density and diversity (average density: 3220 ind/100 cm², 13 taxa). Increasing meiofauna colonization with time is probably related to the development of a biofilm making the leaf more attractive for meiofauna. The effect of distance was not so pronounced as that of time. Total absolute densities were highest in the far series (5 m away from natural seagrass patch), mainly because of nematode densities. Meiofauna diversity was lower in the far series than in the close series (at the border of the natural seagrass patch). A larger individual leaf surface area did not affect the overall meiofauna densities but had a significant positive effect on copepod densities. Larger surface areas promoted the presence of epiphytic copepod families such as Tegastidae and Dactylopusiidae. Overall, we found a rapid recovery of meiofauna in fragmented seagrass beds with primary colonizers (both nematodes and benthic opportunistic copepods) originating from the sediment and later colonizers as epiphytic copepods and their nauplii from the local seagrass regeneration pool.     

Glucosinolates and other metabolites in the leaves of <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> from natural populations and their effects on a generalist and a specialist herbivore

Because many secondary metabolites in plants act as defense against herbivores it has been postulated that these compounds have evolved under selective pressure by insect herbivores. One explanation for the within-species variation in metabolite patterns in a particular species is that different populations are under selection by different herbivores. We tested this hypothesis, using Arabidopsis thaliana plants that originated from dune and inland areas. We analyzed Arabidopsis thaliana leaves using NMR spectroscopy and multivariate data analysis. Major differences in chemical composition were found in water-methanol fractions and were due to higher concentrations of sinigrin and fumaric acid in dune plants. Inland plants showed lower levels of glucose. Quantitative analysis of glucosinolates was performed with HPLC. Individual plants and populations demonstrated differences in glucosinolate composition and concentration. In growth chamber experiments, the generalist herbivore, Spodoptera exigua grew significantly better on the inland plants, while the specialist herbivore Plutella xylostella performed equally well on plants of both origins. Aliphatic glucosinolate as well as total glucosinolate concentrations negatively correlated with larval mass of Spodoptera exigua. No significant correlations, however, were found between larval mass of Plutella xylostella and glucosinolates in the leaves. A specialist and a generalist herbivore were responding differently to plant secondary chemistry, as was also found in several other studies. This is an important indication that differences in glucosinolate concentrations among populations may result from differential selection by different guilds of herbivores.     

Within-plant allocation of a chemical defense in Secale cereale. Is concentration the appropriate currency of allocation?

Summary. The among-leaves allocation of DIBOA, a hydroxamic acid associated with plant resistance, in the shoot of rye (Secale cereale) was evaluated over the vegetative development of the plant. The appropriateness of using the concentration of secondary metabolites, DIBOA in this case, as the parameter to evaluate defense allocation in plants is discussed. Both biological and statistical arguments are put forward to suggest that allocation of chemical defenses should refer to absolute content and not to concentration. Results showed that leaf age was significantly linked to leaf concentration of DIBOA, young leaves having higher concentrations. In contrast, leaf content of DIBOA, our proposed currency of allocation, was not significantly higher in younger leaves. Furthermore, a regression analysis showed that the DIBOA content of leaves was better explained by the leaf relative biomass (proportion of shoot biomass) than by leaf biomass itself. It is suggested that, rather than leaf age, leaf relative biomass is the major factor determining DIBOA allocation in rye shoots. It is proposed that studies addressing within-plant defense allocation should use chemical defense content as the currency, emphasizing the major factors driving this process and its underlying mechanisms. Likewise, it is proposed that studies aiming at characterizing optimal patterns of plant defense should use chemical defense concentration as the currency, and be accompanied by evaluations of the actual resistance against herbivores of the plant parts analyzed, together with the effect on plant fitness. Received 19 February 1999; accepted 28 April 1999.     

Mother really knows best: host choice of adult phytophagous insect females reflects a within-host variation in suitability as larval food

Non-random distribution patterns of specialized phytophagous insects on their hosts may depend on intraspecific differences in plant tissue quality, including nutrients and secondary compounds. Secondary compounds are involved in plant resistance, but are also important for the recognition and acceptability of plants as resources by specialized insects. If individuals within a plant species vary in their content of such secondary substances, there may also be qualitative differences between them. In such cases, natural selection will favor insects with the ability to distinguish and prefer the more suitable plants. In Sweden, the leaf beetle Gonioctena linnaeana Schrank (Coleoptera, Chrysomelidae) is highly specialized on one host, the native willow Salix triandra L (Salicaceae). Field observations reveal that some host plants in a population harbor many feeding larvae, causing severe defoliation, whereas neighboring plants may have few or no feeding larvae. Our hypothesis is that the distribution pattern of G. linnaeana larvae in this population results from qualitative differences between individual host plants in combination with the ability of G. linnaeana females to distinguish between plants that are suitable and not suitable for offspring performance. We examine whether larval survival differs depending on diet and whether the content of secondary chemical compounds explains female preference. Based on the higher survival rate of larvae reared on leaves from preferred hosts, we conclude that G. linnaeana females have evolved a behavior that maximizes offspring performance and thus positively affects female fitness. A chemical survey of the plants indicates that luteolin-7-glucoside and an unidentified flavonoid are important for separating the preferred from the non-preferred plants.     

Multiple feeding stimulants in <Emphasis Type="Italic">Sinapis alba</Emphasis> for the oligophagous leaf beetle <Emphasis Type="Italic">Phaedon cochleariae</Emphasis>

Summary. In earlier investigations on host plant discrimination of leaf beetles glucosinolates were described as feeding stimulants for the Brassicaceae specialist Phaedon cochleariae F. (Coleoptera: Chrysomelidae). However, since these findings could not be confirmed in later studies offering 2-propenylglucosinolate in concentrations corresponding to those detected in host plant leaf material, the identification of feeding stimulants of this leaf beetle species remained unclear. In order to investigate which compounds of the host plant Sinapis alba (Brassicaceae) are involved in feeding stimulation, leaf extracts of different polarities were tested in bioassays with adults of P. cochleariae. Number of feeding beetles and net consumption rates were highest on pea leaves painted with methanol extracts of S. alba, whereas weak feeding responses were also detectable for hexane extracts. In subsequent bioassay-guided fractionations of methanol extracts with semi-preparative high performance liquid chromatography, two distinct fractions, one containing glucosinolates and another containing flavonoids, were found to stimulate beetles to feed to variable degrees. Other collected fractions had zero activity. The combination of both active fractions evoked significantly higher consumption rates and stimulated more beetles to feed than fractions tested individually. At least one compound of each fraction, among these the main glucosinolate of S. alba, 4-hydroxybenzylglucosinolate, act additively. Effects of two different naturally-occurring ratios of glucosinolates and flavonoids on the strength of feeding responses were investigated by use of extracts of two sets of host plants differently exposed to radiation. One set was outdoors-exposed, whereas the second set was kept in the greenhouse. However, the feeding behaviour of P. cochleariae was not affected by the significantly different relative compositions of both compound classes in the host material. In conclusion, mustard leaf beetles need a combination of distinct plant metabolites acting in concert for feeding stimulation, whereby the mere presence of these stimulants, but probably not the ratio of involved compounds, determines their feeding response.     

Species-specific responses of herbivores to within-plant and environmentally mediated between-plant variability in plant chemistry

Allocation of resources to growth and defense against herbivores crucially affects plant competitiveness and survival, resulting in a specific distribution of assimilates and defense compounds within plant individuals. Additionally, plants rarely experience stable environmental conditions, and adaptations to abiotic and biotic stresses may involve shifts in resistance to herbivores. We studied the allocation of phytochemicals in Brassica oleracea (Brussels sprouts) due to leaf age, drought stress and herbivore damage and assessed effects on two lepidopteran herbivores differing in diet breadth: the generalist Spodoptera littoralis and the specialist Pieris brassicae. Glucosinolates as secondary defense compounds and total nitrogen and carbon were quantified and linked to plant palatability, i.e., herbivore feeding preference. Herbivore responses were highly species-specific and partially related to changes in phytochemicals. Spodoptera littoralis preferred middle-aged leaves with intermediate levels of glucosinolates and nitrogen over young, glucosinolate and nitrogen rich leaves, as well as over old leaves, poor in glucosinolates and nitrogen. In contrast, P. brassicae preferred young leaves. Both species preferred severely drought-stressed plants to the well-watered control, although analyzed glucosinolate concentrations did not differ. Both S. littoralis and P. brassicae feeding induced an increase of indole glucosinolate levels, which may explain a reduced consumption of damaged plants detected for S. littoralis but not for P. brassicae. By revealing distinct, sometimes contrasting responses of two insect herbivores to within-plant and stress-mediated intraspecific variation in phytochemistry of B. oleracea, this study emphasizes the need to consider specific herbivore responses to understand and predict the interactions between herbivores and variable plants.     

Distinctive types of leaf tissue damage influence nutrient supply to growing tissues within seagrass shoots

Herbivory is now recognized as an important structuring agent in seagrass meadows but the attack pattern and tissue damage of consumers are highly variable. Nutritional preferences of herbivores and/or easy access to resources may cause differences in biomass loss among tissues that damage the plant in functionally distinctive ways. The two main Mediterranean herbivores, the fish Sarpa salpa (L.) and the sea urchin Paracentrotus lividus (Lmk.), remove higher amounts of intermediate and external shoot leaves, respectively. To test whether this selective feeding can have different consequences on the allocation patterns of nutrient within plants, we simulated the effect of both herbivores by clipping external and intermediate leaves (plus unclipped controls) of Posidonia oceanica (L.) and we measured plant tolerance in terms of shoot growth and leaf nutrient supply to new tissue using isotopic markers. As expected, control treatments displayed high carbon and nutrient supply from external leaves (83% of the total ¹⁵N and 84% of the total ¹³C incorporated by the shoot). When subjected to clipping, the remaining leaves enhanced carbon and nitrogen supply compared with the control by 16% of N and 36% of C—in the intermediate clipping—and by over 100% of N and 200% of C—in the external clipping—to compensate for the nutrient lost. However, only in the case of fish herbivory (intermediate clipping), enhanced supply alone was able to fully compensate for the nutrient losses. In contrast, this mechanism is not completely effective when external leaves are clipped (urchin herbivory). Yet, the consequences of this nutrient loss under sea urchin herbivory are not apparent from the nutrient content of the new tissue, suggesting that there are other sources of nitrogen income (uptake or reallocation from rhizomes). Our study does not only confirm the tolerance of P. oceanica to herbivory, but also constitutes the first evidence of leaf-specific, compensatory nutrient supply in seagrasses.     

A test of elemental defence against slugs by Ni in hyperaccumulator and non-hyperaccumulator Streptanthus species

Summary. Tissues of most plant species contain < 10 μg Ni g⁻¹ but Ni hyperaccumulators contain more than 1000 μg Ni g⁻¹ . Hyperaccumulated Ni can defend plants from some herbivores but the defensive role of lesser Ni concentrations is little explored. We raised five species of Streptanthus (Brassicaceae) native to ultramafic soils, one of which (S. polygaloides) is a Ni hyperaccumulator whereas the others are simply Ni-tolerant, on Ni-amended and unamended green-house soils to create plants differing in Ni concentrations. On high-Ni soil, leaves of the hyperaccumulator contained 3800 μg Ni g⁻¹ whereas leaves of non-hyperaccumulator species contained 41–64 μg Ni g⁻¹. Plants of all species grown on low-Ni soils had < 14 μg Ni g⁻¹. Slugs (Limax maximus) were fed plant material in no-choice tests over a 50-day period and survival and mass changes were recorded. All slugs fed high-Ni leaves of the hyperaccumulator species died within 21 d. Slugs fed high-Ni leaves of the other species did not differ significantly in survival or mass change from those fed low-Ni leaves. In choice tests, slugs (Lehmannia valentiana) offered both high- and low-Ni S. polygaloides leaves did little damage to high-Ni leaves. We conclude that hyperacumulated Ni can defend S. polygaloides from slug herbivory via both toxicity and deterrence, but these defensive effects do not extend to Streptanthus species containing < 70 μg Ni g⁻¹.     

The roles of nicotine and natural enemies in determining larval feeding site distributions of Manduca sexta L. and Manduca quinquemaculata (Haworth) on tobacco

Summary. Field observations indicated that hornworms select feeding sites non-randomly on tobacco. We tested the hypotheses that differences in feeding site locations of larvae of Manduca sexta L. and Manduca quinquemaculata (Haworth) (Lepidoptera: Sphingidae) on tobacco could be explained by differential nicotine concentrations within plants and leaves, species-specific responses to nicotine, or pressure exerted by natural enemies. Results showed that third-instar larvae of M. sexta fed more proximally and centrally on the leaf, whereas M. quinquemaculata fed more distally. Within-plant selection of leaves did not differ; both species selected leaves in the middle region of the plant. Nicotine concentrations in a high nicotine genotype, NC95, varied within each leaf, increasing 2—3 fold from the basal to apical portion of the leaf, and within each plant, increasing 7—10 fold from the first fully expanded leaf to the twelfth (lowest) leaf. In laboratory bioassays, both Manduca species responded to nicotine as a feeding deterrent. Electrophysiological studies demonstrated that gustatory organs of both species responded to nicotine at concentrations found in tobacco leaves and that M. quinquemaculata generally showed a less vigorous response to nicotine than M. sexta. Field mortality of M. sexta due to parasitism by Cotesia congregata (Say) and to parasitism and predation combined differed among feeding sites; predation alone did not. Results suggest that although nicotine concentration and species specific responses to nicotine play a role in determining feeding site locations, pressure exerted by natural enemies, especially parasitism by C. congregata, is more important. Received 22 February 2000; accepted 20 July 2001.     

Controlling bacterial leaf blight of rice and enhancing the plant growth with endophytic and rhizobacterial Bacillus strains

This study was conducted to assess efficacy of biological control against bacterial leaf blight (BLB) of rice produced by Xanthomonas oryzae pv. oryzae. Five endophytic strains (A1, A2, A3, A13 and A15) and two rhizospherial Bacilli (D29 and H8) were tested for their antagonistic activities against BLB in vitro and in vivo. All seven strains showed high potential of antagonistic activity against X. oryzae pv. oryzae and three phytopathogenic fungi in vitro. Test of 16SrRNA gene sequence were assigned isolates A1, A3 and A13 as Bacillus amyloliquefaciens while isolates A2 and A15 as B. methylotrophicus and B. subtilis, respectively. In greenhouse, four strains of displayed 50.29%–57.86% inhibition rate against the pathogen and significantly increased plant fresh weight from 50.03% to 73.11% and dry weight from 64.11% to 86.65% in treated rice plants. In addition, these strains demonstrated strong capability to produce indole-3-acetic acid, siderophores, solubilizing phosphate and also colonize roots. Real-time quantitative polymerase chain reaction revealed that expression of defense-related genes including OsAOS2, OsJMT1, OsNPR1 and OsPR1b were significantly up-regulated in leaves of D29-exposed rice plants, suggesting that treatment of rice with D29 suppressed BLB through systemic activation of the plant defense system. Therefore, data suggest that Bacillus isolates A13, A15, D29 and H8 support effective antagonistic activity against BLB under greenhouse conditions in addition to their potential to promote growth of rice plants.     

Effects of plant morphology on small-scale distribution of invertebrates

Habitat structure influences organism communities by mediating interactions between individuals and species, affecting abundance and species richness. We examined whether variations in the morphology of soft-bottom plants affect their function as habitat and whether complex structured plants support higher macroinvertebrate abundance and species richness. Three Baltic Sea plant species were studied, together with artificial plants resembling each species. In a field collection, we found higher invertebrate abundance on the morphologically more complex plants Myriophyllum spicatum and Chara baltica than on the structurally simpler plant Potamogeton perfoliatus. In a colonization experiment, we found the highest invertebrate abundance on artificial M. spicatum but found no difference between natural plants. Invertebrate taxon richness displayed no consistent relationship with plant structural complexity. The results imply that plant morphology influences small-scale invertebrate distribution, partly supporting the hypothesis that structurally complex plants harbour higher invertebrate abundance.     

Biochemical transformation of birch leaf phenolics in larvae of six species of sawflies

Summary. We investigated the biochemical transformation of individual phenolic compounds of mountain birch leaves in larvae of six birch-feeding sawfly species: Amauronematus amplus, Pristiphora alpestris, Nematus brevivalvis, Priophorus pallipes, Arge sp. and Nematus viridis by comparing the phenolic residues in larval faeces to those of their leaf diet. Partial hydrolysis of hydrolysable tannins, isomerisation of chlorogenic acid and glycosylation of flavonoid aglycones were observed in all studied species. Moreover, we found considerable among-species variation in the composition of phenolic compounds in larval faeces. In addition to foliar phenolics, seventeen non-foliar phenolic metabolites, including eight phenolic acids and nine flavonoid glycosides were detected from the faeces. Of the non-foliar phenolic acids, four were egested species-specifically and only two by all six sawfly species. We also detected differences in the ratios of chlorogenic acid isomers in the faeces of different species, which can indicate different physiological conditions in the guts of studied larvae. In addition to the qualitative differences, quantitative differences were detected in the egestion of chlorogenic acids, possible o-quinone precursors in the larvae. Detected differences, either qualitative or quantitative, could not be explained by seasonal changes in the content of compounds in the leaf diet.     

Insecticidal and growth-reducing activity of foliar extracts from Meliaceae

Summary Thirty-one species in twenty genera of the plant family Meliaceae were assayed for the production of growth-inhibiting phytochemicals, using the generalist herbivorePeridroma saucia. Most species were inhibitory when methanolic extracts were incorporated into artificial diets at concentrations at or below those occurring naturally. In general members of the subfamily Melioideae were more inhibitory than members of the Swietenioideae. Extracts of deciduous species with short leaf lifetimes were significantly more inhibitory than those of evergreen species with longer leaf lifetimes. In a smaller sample of species, evergreen species showed a trend towards having tougher leaves than deciduous species. These results support the resource availability hypothesis of Coleyet al. (1985), and suggest that life history attributes may be of some value in selecting plants for phytochemical prospecting.     

Geese and dietary allelochemicals — food palatability and geophagy

Summary The palatability of plants was studied in young, immature and adult geese of three species,Anser anser, Anser indicus andBranta canadensis, with respect to secondary plant metabolites. In their first 1–4 weeks of life, hand-reared goslings feed on a wide variety of plants, more or less irrespective of their allelochemical contents. Older birds become more selective, but still consume plants which are normally considered to be unpalatable or even toxic for other animals. Choice experiments were performed with pure secondary metabolites which were offered on otherwise highly palatable food items, such as leaves ofTaraxacum officinale. These experiments revealed a similar trend, in that very young goslings discriminate their food much less than older goslings or adult geese. In general, food contaminated with essential oils was rejected, whereas alkaloids, glycosides, amines and sulfur compounds were tolerated to a remarkable degree. In consequence, especially young, but also adult geese must have a high capacity to tolerate and/or to detoxify dietary allelochemicals. Another detoxification mechanism became evident during the experiments: geese ingest soil and mud quite regularly. It could be shown experimentally that the respective soil had a high capacity to bind alkaloids. We assume that geophagy is a means (besides a presumed active detoxification in the liver) to adsorb and thus reduce the contents of dietary allelochemicals.