Ranking of individual mountain birch trees in terms of leaf chemistry: seasonal and annual variation

Summary. The quality of tree leaves as food for herbivores changes rapidly especially during the spring and early summer. However, whether the quality of an individual tree in relation to other trees in the population changes during the growing season and between years is less clear. We studied the seasonal and annual stability of chemical and physical traits affecting leaf quality for herbivores. Rankings of trees in terms of the contents of two major groups of phenolics in their leaves, hydrolyzable tannins and proanthocyanidins (condensed tannins), were very stable from the early spring to the end of the growing season. There were also strong positive within-season correlations in the levels of some other groups of phenolics in the leaves (kaempferol glycosides, myricetin glycosides and p-coumaroylquinic acid derivatives). The contents of individual sugars and the sum content of protein-bound amino acids showed patterns of seasonal consistency in mature leaves, but not in young developing leaves. The seasonal correlations in leaf water content and toughness were also strongest in mature leaves. The correlations between two years at corresponding times of the growing season were strongly positive for the major groups of phenolics throughout the season, but were more variable for the contents of proteins and some sugars. Leaf toughness and water content showed strong positive correlations in mature leaves. Despite the consistency of tree ranking in terms of leaf phenolics, the relative resistance status of trees may, however, change during a growing season because there was a negative correlation between the content of hydrolyzable tannins (early-season resistance compounds) in leaves early in the season and the content of proanthocyanidins (late-season resistance compounds) late in the season, and vice versa. Thus, assuming that phenolics affect herbivore preference and performance, different plants may suffer damage at different times of the growing season, and the overall variation between trees in the fitness consequences may be low. In addition, the adaptation of herbivorous insects to mountain birch foliage in general, as well as to specific tree individuals, may be constrained by variation in the relative resistance status of the trees.     

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.     

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.     

Systemically-induced response of cabbage plants against a specialist herbivore, Pieris brassicae

Summary. Plant responses to herbivory might directly affect the herbivore (“direct” defences) or might benefit the plant by promoting the effectiveness of natural antagonists of the herbivores (“indirect” defences). Brussels sprouts attacked by Pieris brassicae larvae release volatiles that attract a natural antagonist of the herbivores, the parasitoid Cotesia glomerata, to the damaged plant. In a previous study, we observed that feeding by caterpillars on the lower leaves of the plant triggers the systemic release of volatiles detectable by the parasitoids from upper leaves of the same plant.?The role of these systemically induced volatiles as indirect defence and the dynamics of their emission were investigated in wind-tunnel dual choice tests with C. glomerata. The systemically induced emission of volatiles varied depending on leaf age and on plant age. Systemic induction affected parasitoid effectiveness, as induced plants could be more easily located by parasitoids than non-induced ones.?The role of the systemic induction as a direct defence was investigated through behavioural and feeding tests with P. brassicae. In dual choice assays, 1st instar larvae preferred to feed and fed more on systemically induced than on non-induced leaves. In single choice assays, the leaf area consumed by caterpillars was larger on systemically induced leaves than on non-induced control leaves. However, caterpillars fed on systemically induced leaves attained the same weight as those feeding on non-induced controls. In addition, P. brassicae pupae whose larvae were fed on systemically induced leaves had longer developmental times than those of larvae fed on non-induced leaves. Adult oviposition behavior was not influenced by systemic induction.?We conclude that systemically induced responses in cabbage might reduce P. brassicae fitness both directly, by affecting their development and feeding behavior and indirectly by making caterpillars and pupae more vulnerable to attack by carnivores. The occurrence of a possible relationship between direct and indirect defence is discussed. Received 24 January 2001; accepted 3 May 2001.     

The occurrence of aristolochic acids in neotropical troidine swallowtails (Lepidoptera: Papilionidae)

Summary. Analysis of individuals of 17 troidine species demonstrated the presence of aristolochic acids in these butterflies in varying concentrations. Although aristolochic acids do not occur in Aristolochia galeata leaves, they were present in Battus polydamas larvae reared on these leaves, and thus may be synthesized by the larvae from chemical precursors in the plant. Received 17 August 1999; accepted 20 January 2000     

Seasonal changes in nucleic acids,amino acids and protein content in juvenile Norway lobster (<Emphasis Type="Italic">Nephrops norvegicus</Emphasis>)

The objective of this study was to describe the seasonal variations in nucleic acid contents and amino acid profiles in the muscle of juvenile Nephrops norvegicus. RNA and protein contents, and RNA:protein and RNA:DNA ratios varied significantly between seasons, being highest in spring and lowest in autumn/winter ( P<0.05). Though DNA content increased significantly from autumn to summer ( P<0.05), protein:DNA ratio did not show significant seasonal variations ( P=0.05). In respect to protein-bound amino acid content (BAA), a significant increase was observed from winter to summer ( P<0.05). Both essential (EAA) and non-essential amino acids (NEAA) increased significantly (27.6% and 27.8%, respectively; P<0.05), mainly due to the considerable increase in arginine and proline (59.1% and 225.2%, respectively; P<0.05). A significant decrease was observed in the free amino acid content (FAA) from winter to summer ( P<0.05); and a higher percentage decrease occurred in free non-essential (FNEAA; 27.9%) in comparison to free essential amino acids (FEAA; 21.8%). The significant increase in RNA and BAA contents from winter to spring may be related to protein synthesis. On the other hand, the lowest values obtained in winter may be due to a reduction in feeding activity; in this period the muscle protein must be progressively hydrolysed, which is evident with the higher FAA content. The liberated amino acids enter the FAA pool and become available for energy production.     

Spread of metals through an invertebrate food chain as influenced by a plant that hyperaccumulates nickel

Summary. Hyperaccumulation of metals in the shoot system of plants is uncommon, yet taxonomically and geographically widespread. It may have a variety of functions, including defense against herbivores. This study investigated the effects of hyperaccumulation on metal concentrations across trophic levels. We collected plant material, soil, and invertebrates from Portuguese serpentine outcrops whose vegetation is dominated by the nickel hyperaccumulator Alyssum pintodasilvae. Samples were analyzed for nickel, chromium, and cobalt. Grasshoppers, spiders, and other invertebrates collected from sites where A. pintodasilvae was common had significantly elevated concentrations of nickel, compared to nearby sites where this hyperaccumulator was not found. Chromium and cobalt, occurring in high concentrations in the serpentine soil but not accumulated by A. pintodasilvae, were not elevated in the invertebrates. Therefore, it appears likely that a flux of nickel to herbivore and carnivore trophic levels is specifically facilitated by the presence of plants that hyperaccumulate this metal. The results may be relevant to the development of phytoremediation and phytomining technologies, which use plants to extract metals from the soil. Reveived 22 August 2002; accepted 2 April 2003. R1D=" Correspondence to: A. J. Pollard, e-mail:joe.pollard@furman.edu     

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     

Larval chemical defence and evolution of host shifts in Chrysomela leaf beetles

Summary. Larvae of Chrysomela leaf beetles release for defence volatile compounds belonging to various chemical families. This study focuses on the defensive strategy based on the esterification of isobutyric acid and 2-methylbutyric acid with a wide variety of alcohols taken up from the host plant. To date, only two species are known to produce these repellents C. interrupta, which is associated with Betulaceae and C. lapponica which occurs either on Betulaceae or Salicaceae.? In order to know if other species have developed this chemical defence and how the food plant influences the secretion of these toxins, we targeted by mass spectrometry the presence of iso- and 2-methylbutyric acids and esters of them in the defensive secretions of Chrysomela larvae exclusively associated with Betulaceae or Salicaceae. ?Screening analyses reveal that the synthesis of these compounds is a common character restricted to all the members belonging to the C. interrupta group sensu Brown (1956) regardless of the host-plant family. These results suggest that the biochemical mechanism leading to the synthesis of these compounds could be considered as a synapomorphy meaning that the group is probably monophyletic. ?Defensive secretions of the members of the interrupta group are quantitatively assayed for iso- and 2-methylbutyric acids and their (Z)-3-hexenyl esters. Results reveal a chemical plasticity developed by Chrysomela species associated with Salicaceae. The amounts of iso- and 2-methylbutyric acids derivatives and of salicylaldehyde in their larval secretions depend on the food plant and on its content in phenolglucosides. Received 5 October 1998; accepted 25 November 1998.     

Damage-inducibility of primary and secondary metabolites in the wild parsnip (Pastinaca sativa)

Summary. Induction of secondary metabolites to herbivore damage is a widespread phenomenon among plants and serves to enhance resistance by reducing suitability or increasing toxicity of foliage. Post-damage responses of primary metabolites are less well known; reductions in primary metabolites may increase resistance by decreasing palatability or nutritional suitability for herbivores or by potentiating toxicity of secondary metabolites. In this study, we examined response to simulated herbivory in Pastinaca sativa, the wild parsnip, in both primary and secondary metabolites. We found that induction of secondary metabolites in response to damage is largely restricted to a single class of compounds, the furanocoumarins. These prooxidant compounds are known to be toxic to a wide variety of insect herbivores. The only primary metabolite that responded to damage was total soluble protein, which increased significantly 24 h after damage. After 24 h, the correlation between total furanocoumarins and total sugars was significant and negative (r = − 0.697). This correlation likely reflects an independent response of sugar to damage, rather than a diversion of resources into furanocoumarin production, inasmuch as this correlation at 3 h, after furanocoumarin induction had taken place, was not significant. In other secondary metabolite pathways, damage produced a significant decline in farnesene and a C-16 unsaturated fatty acid, 7,10,13-hexadecatrienoic acid, each of which may potentiate the furanocoumarin defense response. Farnesene may enhance photooxidative activation of the furanocoumarins and 7,10,13-hexadecatrienoic acid may serve as a precursor to jasmonic acid, a key hormone in regulating induction responses. With few key exceptions, quantities of both primary and secondary metabolites in wild parsnip foliage are unaffected by damage. Those that are affected may well play a role in resistance of parsnips to herbivores. Received 1 July 1998; accepted 28 September 1998.     

Constitutive and herbivore‐induced volatiles in pear,alder and hawthorn trees

Summary. Qualitative and quantitative differences among pear cultivars were found in constitutive and Cacopsylla-induced volatiles, depending on experimental treatment of the trees (i.e., uninfested and partly or completely infested by psyllids). Blend differences were also found between pear cultivars and wild-type pear, alder and hawthorn–the latter trees are frequently present in pear orchard hedgerows. ?Interesting differences were found in the presence of methyl salicylate and (E,E)-α-farnesene, two compounds previously found to mediate attraction of predatory bugs towards psyllid-infested pear trees. Methyl salicylate is expressed constitutively and is induced systemically by infestation in the whole plant of all four cultivars. (E,E)-α-farnesene on the other hand showed also systemic induction in Bartlett, NY10355 and Beurré Hardy, but in partially infested Conference trees it was induced locally, only in herbivore-damaged leaves. No methyl salicylate or (E,E)-α-farnesene were identified in honeydew. In field collected headspace samples of alder leaves infested by aphids and leaf beetles we found methyl salicylate but no (E,E)-α-farnesene, whereas in uninfested hawthorn neither were identified. Insight in the variability of damage-related pear volatiles will have important implications for integrated pest management in the field. Received 27 August 2002; accepted 28 November 2002 R1D=" Correspondence to: Petru Scutareanu, e-mail:scuterea@science.uva.nl     

Rutabaga roots, a rich source of oviposition stimulants for the cabbage root fly

Summary. Two recently identified compounds ("CIFs"), present on the leaf surface of Brassica oleracea (cabbage), are the strongest oviposition stimulants known for the cabbage root fly, Delia radicum. Cabbage leaves contain these compounds in extremely low concentrations, and the amount of CIFs obtained from purifying leaf extracts was so small that it limited further research. We were able to purify far more of these two compounds from the roots of Brassica napus var. napobrassica (rutabaga). Apart from being a richer source of CIFs, rutabaga roots are considerably easier to collect and process than leaves. In addition, we isolated and identified a new CIF compound from the roots that is also very active in stimulating oviposition in the cabbage root fly. Received 26 May 2000; accepted 26 July 2000     

Integrating phenological, chemical and biotic defences in ant-plant protection mutualisms: a case study of two myrmecophyte lineages

Summary. We examined the role of plant phenology in the evolution of anti-herbivore defence in symbiotic ant-plant protection mutualisms. Phenology of the host-plant affects traits of its herbivores, including size, growth rate, development time, and gregariousness. Traits of herbivores in turn determine what traits ants must have to protect their host. Diversity in plant phenological traits could thus help explain the great ecological diversity of coevolved ant-plant mutualisms. We explored the postulated causal chain linking phenology of the plant, herbivore adaptations to phenology, and ant adaptations for protection, by comparing two myrmecophytes presenting strong contrasts in phenology. In Leonardoxa africana, a slow-growing understory tree, growth at each twig terminal is intermittent, the rapid flushing of a single leaf-bearing internode being followed by a pause of several months. In contrast, axes of Barteria nigritana, a tree of open areas, grow continuously. Analysis of the phenology (kinetics of expansion) and chemistry of leaf development (contents of chlorophylls, lignin, and nitrogen during leaf growth) showed that these two species exhibit strongly contrasting strategies. Leonardoxa exhibited a delayed greening strategy, with rapid expansion of leaves during a short period, followed by synthesis of chlorophylls and lignins only after final leaf size has been reached. In contrast, leaves of Barteria expanded more slowly, with chlorophylls and lignin gradually synthesised throughout development. Differences in the phenology of leaf development are reflected in differences in the duration of larval development, and thereby in size, of the principal lepidopteran herbivores observed on these two plants. This difference may in turn have led to different requirements for effective defence by ants. The strategy of phenological defence may thus affect the evolution of biotic defence.     

Growth and utilization of internal nitrogen reserves by the giant kelp Macrocystis pyrifera in a low-nitrogen environment

An adult giant kelp plant (Macrocystis pyrifera), moved from an inshore kelp forest to an offshore, low-nitrogen environment near Santa Catalina Island, California (USA), maintained growth for 2 wk on internal nitrogen reserves. Frond elongation rates decreased significantly during the third week, and plant growth rate (wet wt) dropped from an initial inshore rate of 3.6 to 0.9% d^-1. During this 3 wk period, nitrogen contents and free amino acid concentrations decreased, while mannitol and dry contents increased in frond tissues. After depletion of internal nitrogen reserves, the nitrogen content of lamina and stipe tissues averaged 1.1 and 0.7% dry wt, respectively. The experimental plant was exposed to higher ambient nitrogen concentrations during the last 2 wk. Rates of frond elongation and plant growth increased, but nitrogen content and amino acids in frond tissues remained low. Of the total nitrogen contained in frond tissue of the plant before transplantation, 58% was used to support growth in the absence of significant external nitrogen supply. Amino acids constituted a small proportion of these internal nitrogen reserves. Net movement of nitrogen occurred within large fronds, but not between different frond size classes. The nitrogen content of holdfast tissue remained relatively constant at 2.4% dry wt and accounted for 18 to 29% of the total nitrogen. Holdfast nitrogen was not used to support growth of nitrogen-depleted fronds. In comparison to Laminaria longicruris, which is adapted to long seasonal periods of low nitrogen availability, M. pyrifera has small nitrogen-storage capacity. However, internal reserves of M. pyrifera appear adequate to make nitrogen starvation uncommon in southern California kelp forests.     

The Mediterranean intertidal habitat as a natural laboratory to study climate change drivers of geographic patterns in marine biodiversity

Nylon bags containing yellow leaves of Rhizophora apiculata and Avicennia marina, were immersed for 80 days from August to October, 1996. the decomposing leaves were collected every 10 days and analysed for dry weight loss and six biochemical parameters: tannins, total amino acids, total sugars, total nitrogen, total lipids and fatty acid profile. the leaf weight initially decreased very rapidly by about 50% of the start in two species of mangroves within 10 days. Similar changes were observed with tannins, total amino acids and sugars. However, the concentration of nitrogen increased significantly with decomposition. There was no significant change in total lipid and fatty acid profile. the highest concentration of fatty acid in the decomposing leaves was palmitic acid (16:0). Unsaturated fatty acids such as, 18:1 w7c and 18:1 w9c were found to be present in decomposing leaves of both species.     

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.     

Activity of insect juvenile hormone III: seed germination and seedling growth studies

Summary. Juvenile hormones are sesquiterpenoids that regulate developmental processses such as metamorphosis and reproduction in insects. Insect juvenile hormone III (JH III), methyl-10R,11-epoxy-3,7,11-trimethyl-2E,6E-dodecadienoate, has also been identified in two sedge species, Cyperus iria and C. aromaticus (Toong et al. 1988). Potential allelopathic activity of this compound and the structurally related sesquiterpenoid farnesol was investigated using seed germination and seedling growth assays with radish, lettuce and rice. Treatment of seeds with JH III delayed lettuce seed germination and potently inhibited rice shoot growth. Both farnesol and JH III inhibited the growth of C. iria seedlings. The antimicrobial activity of JH III was also tested on a taxonomic and ecologically diverse range of fungi. Using the classic cytotoxic disk assay, JH III did not effect growth of the fungal species tested. We believe that JH III may contribute to the aggressive nature of this invasive weed species. Received 11 October 1999; accepted 26 February 2000     

Rhythmic release of dissolved free amino acids from partly synchronized Thalassiosira rotula under nearly natural conditions

In outdoor tanks enclosed monocultures of Thalassiosira rotula showed partly synchronized cell divisions during exponential growth. The changes in division activity showed a characteristic frequency, which also could be found for cell length (pervalvar axis) and assimilation rate as well as for concentration changes of dissolved free amino acids (DFAA) and ammonia. Excretion of DFAA occurred preferentially during periods of reduced cell division activity, when assimilation was at a higher level. The concentration of ammonia decreased during these periods. The rates per hour of these parameters showed significant multiple correlations. The release of six individual amino acids (ala, -ala, gly, lys, orn and ser) seemed to be controlled by the physiological status of the diatoms. These amino acids amounted to a portion of about 70–80% of the total DFAA. Uptake and release rates of DFAA nitrogen and ammonia were in the same range with maxima of about 350 ng at nitrogen dm^-3 h^-1 and litre. These dynamics pointed to strong fluxes of organic bound nitrogen in marine micro-heterotrophs.     

Lignoid chemical defenses in the freshwater macrophyte Saururus cernuus

Summary. Chemical defense against herbivores has rarely been investigated for freshwater plants, possibly due to the common misconception that herbivory on aquatic macrophytes is low and would not select for chemical defenses. In previous work, the freshwater angiosperm Saururus cernuus was shown to be a low preference food for omnivorous crayfish despite its high nutrient value and relatively soft texture. We used feeding by the crayfish Procambarus clarkii to guide fractionation of the deterrent lipid-soluble extract of this plant, leading to the identification of seven deterrent lignoid metabolites, (–)-licarin A, (+)-saucernetin, (–)-dihydroguaiaretic acid, (–)-sauriols A and B, (–)-saucerneol, and (–)-saucerneol methyl ether. Lignans have been implicated in terrestrial plant chemical defenses as insect growth inhibitors, insect toxins, nematocides, antibacterial, and antifungal agents. However, these activities have rarely been demonstrated using ecologically relevant methodologies in terrestrial systems, and never before in freshwater systems. The widespread nature of lignans amongst very distantly related plants, along with their rich diversity of molecular structure, suggests that they could play a large role in mediating plant-herbivore interactions. In addition to the lignoid compounds we identified, there were other compounds present in low concentration or unstable compounds that were deterrent, that did not appear to be lignans, but that we were unable to identify. This plant thus appears to be defended by a complex mixture of natural products. Received 6 June 2000; revised 23 August 2000; accepted 2 September 2000     

Nutritional quality of phloem sap in relation to host plant-alternation in the bird cherry-oat aphid

Summary. Host alternation in aphids has been attributed to complementary growth of host plants, or more specifically to seasonal changes in the nitrogen quality of the phloem sap. In this report, seasonal fluctuation of free amino acids in phloem of the winter and summer host plants (Prunus padus, bird cherry and Hordeum vulgare, barley) of Rhopalosiphum padi (the bird cherry-oat aphid) were investigated in the context of aphid growth and behaviour. Phloem was collected from the cut stylets of aphids taken from plants that were grown outdoors. The total concentration of amino acids in P. padus phloem increased between bud break and late flush (spring), decreased in mature leaves (summer), and increased again in early senescent leaves (autumn). In H. vulgare, however, amino acid concentration fluctuated less from seedlings to flowering. Spring aphids from P. padus grew rapidly on this host from bud break to late flush, but died on mature and early senescent leaves. Summer aphids from H. vulgare grew as fast on this host as spring aphids did on flush leaves of P. padus. Sexual females grew more slowly than other generations and nearly as well on mature as on early senescent P. padus leaves. As judged by aphid growth and phloem nitrogen quality, P. padus during spring equals the summer host H. vulgare. However, the lower growth rates of R. padi on mature and senescent leaves of P. padus appear only loosely correlated with phloem amino acid concentrations. Therefore, factors influencing aphid nutrition, or ecology, other than seasonal changes in phloem sap amino acid concentration may explain the existence of host alternation in R. padi.