Responses of two hymenopteran predators to surface Chemistry of their prey: Significance for an alkaloid-sequestering caterpillar

Larvae ofUresiphita reversalis (Guenée) (Lepidoptera: Pyralidae) sequester quinolizidine alkaloids from their leguminous hosts and store them primarily in the cuticle. Stored alkaloids are lost with the last larval molt. Extracts of late-instar larvae and of pupae were applied to larvae of the potato tuber moth,Phthorimaea operculella (Zeller) (Gelichiidae), which are normally palatable to two hymenopteran predators, the Argentine ant,Iridomyrmex humilis (Mayr) (Formicidae), and the paper wasp,Mischocyttarus flavitarsus (Sauss.) (Vespidae). Larvae ofP. operculella treated with alkaloid extracts ofU. reversalis larval exuviae, or with surface extracts of whole larvae, were deterrent to both predators, compared to untreated prey. Extracts of pupal exuviae added toP. operculella, however, were not deterrent.P. operculella larvae treated with the authentic alkaloids sparteine and cytisine were also deterrent to these hymenopteran predators. Storage of small but concentrated amounts of plant secondary compounds in the cuticle appears to be an efficacious means of defense against at least two common predators of lepidopteran larvae.     

Detrimental effects ofCinchona leaf alkaloids on larvae of the polyphagous insectSpodoptera exigua

YoungCinchona ledgeriana plants contain two types of alkaloid: indole alkaloids in the leaves and quinoline alkaloids in the root. FromCinchona leaves, a crude alkaloid extract was made, containing the cinchophylline type of indole alkaloids and a small amount of 5-methoxytryptamine. The leaf alkaloid extract exerted a strong detrimental effect on the growth of larvae of the polyphagous beet armyworm,Spodoptera exigua (Lepidoptera). Feeding of larvae on an artificial diet containing the leaf alkaloids at the same concentrations as those found in the plant resulted in significant growth reduction, retardation in development, and mortality of the larvae. Cinchophyllines are composed of 5-methoxytryptamine coupled to a corynantheal unit. When incorporated into the artificial diet, 5-methoxytryptamine alone had no effect on the 5.exigua larvae. Corynantheal, however, had a strong detrimental effect on growth of the larvae, its effect being comparable to that of the leaf alkaloid extract. In contrast to the indole-type leaf alkaloids, the quinolinetypeCinchona root alkaloids did not affect growth and development of the larvae. These results suggest that the indole-type alkaloids, which inCinchona plants are present at the highest concentrations in the young, vulnerable leaflets, are involved in the chemical defense of the plant against herbivorous insects.     

Sequestration ofVeratrum alkaloids by specialistRhadinoceraea nodicornis konow (Hymenoptera,Tenthredinidae) and its ecoethological implications

The larvae of the specialist sawflyRhadinoceraea nodicornis Konow (Hymenoptera, Tenthredinidae) store in their hemolymph ceveratrum alkaloids originating from the host plantVeratrum album L. (Liliales, Melanthiaceae). The major alkaloid found in the hemolymph is 3-acetyl-zygadenine. Qualitative and quantitative data showed that the plant alkaloid 3-angeloylzygadenine is most probably metabolized in the larval gut to zygadenine and then acetylated. A still unidentified alkaloid with a molecular weight of 591 Da was detected in plant leaves as well as in the gut, hemolymph, and excrement of larvae. Protoveratrine A and B, on the other hand, seem to be degraded by the larvae. These findings indicate that the pathway of ceveratrum alkaloids inR. nodicornis larvae is fourfold: direct sequestration, metabolism followed by sequestration, excretion of intact alkaloids, and degradation. In contrast, no ceveratrum alkaloids were detected in the hemolymph and excrement of larvae of the generalist sawflyAglaostigma sp. fed withV. album leaves. Bioassays with the antMyrmica rubra L. proved that the hemolymph ofR. nodicornis larvae is highly deterrent and toxic. In bioassays evaluating defensive efficiency against predators (ants, spiders, and bushcrickets), no larvae were eaten. Ceveratrum alkaloids were also detected in the hibernating prepupae ofR. nodicornis. In feeding bioassays, the shrewCrocidura russula Hermann rarely fed upon prepupae, suggesting that this stage is also protected from predation to some degree. In field surveys, the only parasitoids recorded were two ichneumonid species that are believed to be specialized onR. nodicornis. Bioassays and field observations enable us to suppose thatR. nodicornis and its enemies produce a food web of ion connectance.     

Toxicity of allelochemicals from wild insect-resistant tomatoLycopersicon hirsutum f.glabratum toCampoletis sonorensis,a parasitoid ofHeliothis zea

Greenhouse-grown plants of five tomato lines varying in their level of 2-tridecanone-mediated resistance toManduca sexta (L.) andLeptinotarsa decemlineata (Say) were found to adversely affect larvae ofCampoletis sonorensis (Cameron), a larval endoparasitoid ofHeliothis zea (Boddie), in a manner directly related to their level of resistance. The parasitoid larvae, which emerge as fifth instars from their host and construct a cocoon on the foliage of their hosts' host plant, suffered extensive mortality during cocoon spinning on highly resistant foliage. Mortality was greatest (82%) on the highly resistant plants ofLycopersicon hirsutum f.glabratum (accession PI 134417) and an F₁ backcross [(L. esculentum × PI 134417) × PI 134417] selection. Mortality was intermediate (40 and 28%, respectively) on backcross selections with moderate and low levels of resistance and least (8%) on susceptibleL. esculentum. Removal of the glandular trichomes, which contain 2-tridecanone in their tips, from the foliage eliminated differences in parasitoid mortality among plant lines.Bioassays of 2-tridecanone indicated that it is acutely toxic to fifth instarC. sonorensis larvae at the quantities associated with highly resistant foliage and produces symptoms identical to those observed on resistant foliage. 2-Undecanone, a second methyl ketone present in the glandular trichomes of resistant foliage, was also toxic toC. sonorensis larvae, but significantly less so than 2-tridecanone. The results support the hypothesis that 2-tridecanone is responsible for the observed mortality ofC. sonorensis larvae during cocoon construction on resistant foliage.     

Role of Alkaloids in Resistance of Yellow Lupin to Red-Legged Earth Mite Halotydeus destructor

Eleven varieties of Lupinus luteus were tested in choice and no-choice experiments for their resistance to feeding by the red-legged earth mite (Halotydeus destructor). Three were found to show resistance, and the alkaloid fraction from these varieties strongly deterred mites from feeding. The alkaloid components of the extracts, lupinine, a number of acyl derivatives of lupinine, and sparteine were tested for deterrent activity. Sparteine was the most potent, with significant activity at 0.001% concentration. Quantitation of the levels of nonpolar alkaloids in the cotyledons and leaves of the 11 varieties was by gas chromatography–mass spectrometry. An inverse correlation was found between the concentrations of these alkaloids and the damage caused by the mites; varieties with levels <100 g/g (fresh wt) showed a high damage index. The involvement of sparteine and the bipiperidine alkaloid ammodendrine in the deterrence to mite feeding is discussed.     

Visualizing a Plant Defense and Insect Counterploy: Alkaloid Distribution in <Emphasis Type="Italic">Lobelia</Emphasis> Leaves Trenched by a Plusiine Caterpillar

Insects that feed on plants protected by latex canals often sever leaf veins or cut trenches across leaves before feeding distal to the cuts. The insects thereby depressurize the canals and reduce latex exudation at their prospective feeding site. How the cuts affect the distribution and concentration of latex chemicals was not known. We modified a microwave-assisted extraction technique to analyze the spatial distribution of alkaloids in leaves of Lobelia cardinalis (Campanulaceae) that have been trenched by a plusiine caterpillar, Enigmogramma basigera (Lepidoptera: Noctuidae). We produced sharp two dimensional maps of alkaloid distribution by microwaving leaves to transfer alkaloids to TLC plates that were then sprayed with Dragendorff’s reagent to visualize the alkaloids. The leaf prints were photographed and analyzed with image processing software for quantifying alkaloid levels. A comparison of control and trenched leaves documented that trenching reduces alkaloid levels by approximately 50% both distal and proximal to the trench. The trench becomes greatly enriched in alkaloids due to latex draining from surrounding areas. Measurements of exudation from trenched leaves demonstrate that latex pressures are rapidly restored proximal, but not distal to the trench. Thus, the trench serves not only to drain latex with alkaloids from the caterpillar’s prospective feeding site, but also to isolate this section, thereby preventing an influx of latex from an extensive area that likely extends beyond the leaf. Microwave-assisted extraction of leaves has potential for diverse applications that include visualizing the impact of pathogens, leaf miners, sap-sucking insects, and other herbivores on the distribution and abundance of alkaloids and other important defensive compounds.     

Defensive Role of Tomato Polyphenol Oxidases against Cotton Bollworm (<Emphasis Type="Italic">Helicoverpa armigera</Emphasis>) and Beet Armyworm (<Emphasis Type="Italic">Spodoptera exigua</Emphasis>)

Tomato (Solanum lycopersicum) polyphenol oxidases (PPOs), enzymes that oxidize phenolics to quinones, have been implicated in plant resistance to insects. The role of PPO in resistance to cotton bollworm [Helicoverpa armigera (Hübner)] and beet armyworm [Spodoptera exigua (Hübner)] (Lepidoptera: Noctuidae) was evaluated. Consumption, weight gains, and mortality of larvae feeding on foliage of transgenic tomato lines overexpressing PPO (OP lines) and of larvae feeding on foliage of transgenic tomato lines with suppressed PPO (SP lines) were compared with consumption, weight gains, and mortality of larvae feeding on non-transformed (NT) plants. Increases in foliage consumption and weight gains were observed for cotton bollworms feeding on leaves of SP plants compared to NT and OP plants. PPO activity was negatively correlated with both weight gains and foliar consumption of cotton bollworm, substantiating the defensive role of PPO against this insect. Similarly, beet armyworm consumed less foliage (both young and old leaves) from OP plants than SP plants. Larvae feeding on OP leaves generally exhibited lower weight gains than those feeding on SP leaves. These results indicate that tomato PPO plays a role in resistance to both cotton bollworm and beet armyworm.     

Chemical and experiential basis for rejection ofTropaeolum majus byPieris rapae larvae

Rejection of nasturtium,Tropaeolum majus, by cabbage-reared larvae ofPieris rapae has been explained by the presence of feeding deterrents in the nastrutium foliage. Sensitivity to the deterrents develops as neonate larvae feed on cabbage. The most prominent deterrent compound, which is present in nasturtium at a concentration of 40 mg/100 g fresh leaves, was identified as chlorogenic acid. When neonate larvae were fed on a cabbage leaf treated with high concentrations of deterrent-containing extracts of nasturtium foliage, they remained insensitive to the deterrents, so they accepted nasturtium when transferred as second instars. When neonate larvae were reared on a cabbage leaf treated with 0.1 mg chlorogenic acid, ca. 35% of the second instars accepted nasturtium. Similar dietary exposure of neonates to the subunits of chlorogenic acid, caffeic acid and quinic, acid resulted in much less or no effect on the rejection behavior of second instars. The results suggest that the combined effects of specific chemical constituents of nasturtium can explain the rejection of this plant by larvae ofP. rapae, but if larvae are continuously exposed to these compounds immediately after hatching, they apparently become habituated to the feeding deterrents. The lack of activity of the subunits of chlorogenic acid suggests that specific structural features are necessary for a dietary constituent to cause such habituation or suppression of sensitivity development.     

Effects of dietary protein and lupine alkaloids on growth and survivorship ofSpodoptera eridania

Toxic chemicals and nutrients are often positively correlated within and among plants. We studied how such correlations affect the suitability of plants as food for herbivores by assessing the growth and survivorship ofSpodoptera eridania (army worm) on artificial diets containing lupine alkaloids and casein. We found that (1) the effects of casein were determined by other dietary components: increased dietary casein led to increased larval growth only when the diet was also high in wheat germ. (2) Dietary alkaloids were effective at very low concentrations, reducing both growth and survivorship. The alkaloids lupanine and sparteine were not synergistic in their effects, and the interaction between alkaloids and casein was significant only in the low-wheat-germ diets. (3) The effects of casein and alkaloids were generally apparent only in the first instar, and the growth of fifth-instar larvae was unaffected by diet. (4) Using these artificial diet experiments, we can make simple predictions about the food quality of plants grown under various nutrient regimes. These predictions are consistant with recent ideas about optimal plant defenses.     

The “Raison D'être” of pyrrolizidine alkaloids inCynoglossum officinale: Deterrent effects against generalist herbivores

In this study we tested whether pyrrolizidine alkaloids (PAs) ofCynoglossum officinale serve as antifeedants against herbivores. Total PA N-oxide extracts of the leaves significantly deterred feeding by generalist herbivores. Specialist herbivores did not discriminate between food with high and low PA levels. Three PAs fromC. officinale, heliosupine, echinatine, and 3-acetylechinatine, equally deterred feeding by the polyphagous larvae ofSpodoptera exigua. Although the plants mainly contain PAs in their N-oxide form, reduced PAs deterred feeding byS. exigua more efficiently than PA N-oxides. On rosette plants, the monophagous weevilMogulones cruciger significantly consumed more of the youngest leaves, which had the highest PA level and the highest nitrogen percentage. Larvae ofEthmia bipunctella, which are oligophagous within the Boraginaceae, did not discriminate between leaves. All generalist herbivores tested significantly avoided the youngest leaves with the highest PA levels. In the field, the oldest leaves also were relatively more damaged by herbivores than the youngest leaves. It is hypothesized that the skewed distribution of PAs over the leaves of rosette plants reflects optimal defense distribution within the plant.     

Sequestration of pyrrolizidine alkaloids in several arctiid moths (Lepidoptera: Arctiidae)

Sequestration of dietary pyrrolizidine alkaloids (PA) by larvae and adults of six European arctiid moth species (Spilosoma lubricipeda, Arctia caja, Phragmatobia fuliginosa, Callimorpha dominula, Diacrisia sannio, andTyria jacobaeae) was investigated for comparison with the well-studied Asian arctiidCreatonotos transiens. Larvae of all species metabolized free PA bases into the respectiveN-oxides. Only adults ofA. caja, P. fuliginosa, andS. lubricipeda, but not their larvae, converted dietary 7(S)-heliotrine to 7(R)-heliotrine, a direct precursor of a male pheromone in some arctiids, 7(R)-hydroxydanaidal. The larval integument figures as the main storage site for resorbed alkaloids; only minor amounts were found in other tissues. In addition, a significant amount of alkaloid is deposited in the cocoon ofArctia caja; only traces of alkaloids could be found in the meconium and the exuviae of this species. A substantial part of the alkaloids fed was degraded to unknown, nonalkaloidal products.     

Dihydropyrrolizine attractants for arctiid moths that visit plants containing pyrrolizidine alkaloids

Adults of three species of arctiid moths (Cisseps fulvicollis, Ctenucha virginia, andHalysidota tessellaris) are attracted to plants that contain pyrrolizidine alkaloids (PAs). The moths use olfactory cues to locate these plants, then feed on leaves, flowers, and roots with the proboscis. To investigate the chemical basis of attraction, sticky traps were baited with roots of a PA-containing plant,Eupatorium maculatum, alkaloids ofE. maculatum, and several derivatives of these alkaloids. Volatile derivatives of the bicyclic pyrrolizidine skeleton attracted all three arctiid species. The dihydropyrrolizines, (S)-(+)-hydroxydanaidal and (R)-(–)-hydroxydanaidal, proved to be the most attractive compounds tested, accounting for over 70% of the moths captured. Different alkaloid derivatives attracted different proportions of male and femaleCisseps. Both (S)-(+)-hydroxydanaidal (52% male) and (R)-(–)-hydroxydanaidal (71% male) attracted a significantly lower percentage ofCisseps males thanE. maculatum roots (87% male).Cisseps males possess eversible scent organs (coremata) that are displayed during courtship. Analysis of corematal extracts revealed the presence of hydroxydanaidal.Cisseps moths thus resemble danaine and ithomiine butterflies, both in their attraction to PA sources and in the presence of PA derivatives in the male scent organs.     

Systemic induction of feeding deterrents in cotton plants by feeding ofSpodoptera SPP. Larvae

Cotton,Gossypium hirsutum L., has been shown to exhibit systemic induced resistance to arthropods under certain conditions. We conducted experiments to determine the effects of previous feeding ofSpodoptera exigua Hübner andSpodoptera littoralis (Boisd.) larvae on feeding behavior, growth, and survival of larvae subsequently feeding on cotton. In one feeding choice test,S. exigua larvae preferred young leaves from undamaged control plants to undamaged young leaves from a previously damaged plant. Feeding deterrence was noticeable after only 6 hr of initial feeding damage by larvae, and there was almost complete deterrence after 30 and 54 hr of continuous feeding. In a second feeding choice test,S. littoralis larvae fed more on mature leaves from undamaged control plants than on undamaged mature leaves from previously damaged plants. In no-choice tests, third instars ofS. littoralis fed undamaged young leaves from damaged plants did not gain weight and died by the seventh day, whereas larvae fed young leaves from undamaged control plants gained weight and pupated within 11 days. Sixth instars ofS. littoralis fed either old damaged leaves, old undamaged leaves, or young undamaged leaves all from previously damaged plants gained weight slowly and took more than 12 days to pupate, whereas larvae fed young leaves from undamaged plants gained weight rapidly and pupated within five days of the beginning of the experiment.     

Systemic Induction of Allelochemicals in Glanded and Glandless Isogenic Cotton by Spodoptera exigua Feeding

Induction of systemic resistance to feeding of beet armyworm, Spodoptera exigua, was investigated in two isogenic lines of Stoneville 213 cotton, Gossypium hirsutum, that differed in the presence of pigment glands. In laboratory bioassays, larvae strongly preferred to feed on glandless cotton plants when presented a choice between undamaged terminal leaves of undamaged glanded and glandless plants. Feeding damage inflicted by S. exigua larvae on the two oldest leaves of glanded plants seven days prior to feeding bioassays caused larvae to prefer by 33-fold the undamaged terminal foliage from undamaged plants compared to that from damaged plants. Feeding damage on glandless plants caused only a 2.6-fold greater preference for terminal foliage from undamaged plants over foliage from previously damaged plants. Extracts of terminal foliage from glanded cotton damaged seven days earlier had significantly greater quantities of terpenoid aldehydes (hemigossypolone, gossypol, and heliocides) than did foliage from undamaged glanded plants. Terpenoid aldehydes were undetectable in extracts of both undamaged and previously damaged glandless plants. The profile of volatile compounds collected from the headspace of mechanically damaged terminal leaves of undamaged glanded and glandless plants differed. Both cotton isolines released large quantities of lipoxygenase products (hexenyl alcohols, acetates, and butyrates), but glandless plants released only small amounts of mono- and sesquiterpenes compared to glanded plants. Glandless plants damaged seven days prior to volatile collection released significantly greater quantities of lipoxygenase products, -ocimene, and - and -farnesene than did undamaged glandless plants. Previously damaged glanded plants released significantly greater quantities of all mono- and sesquiterpenes and hexenyl acetates and butyrates, but not alcohols. The relative importance of volatile compounds versus terpenoid aldehydes in induced feeding deterrence in cotton to S. exigua larvae is still unclear.     

Diterpenoid Alkaloid Concentration in Tall Larkspur Plants Damaged by Larkspur Mirid

Tall larkspur (Delphinium barbeyi) is a serious poisonous plant threat to cattle on mountain rangelands. The larkspur mirid [Hopplomachus affiguratus] has been proposed as a biological tool to damage tall larkspur in an effort to deter grazing by cattle and thus prevent poisoning. Preliminary data suggested that it may also reduce toxic alkaloid levels. The objective of this study was to determine if damage caused by the larkspur mirid would reduce toxic alkaloid concentration. Larkspur mirids were collected in the field in 1992 and placed on potted plants in the greenhouse. The resulting mirid-damaged leaves were lower in toxic alkaloids than leaves from uninfested plants. In the 1995 field study, toxic and total norditerpenoid alkaloid concentrations were measured in two larkspur populations having established mirid populations and in two newly infested larkspur populations. In the 1996 field study, three widely separated larkspur populations infested with mirids were sampled. Mirid-damaged leaves were lower in toxic alkaloids in both years, but there were no differences in flowering heads. However, only at Yampa, Colorado, did mirids reduce toxic alkaloids to levels that would not pose a threat to cattle. There was no difference in toxic or total alkaloid concentration between larkspur populations with long-term mirid infestations compared to newly infested plants. The plant-to-plant variability in alkaloid concentration was greater than differences due to mirids.     

Quinolizidine alkaloids obtained byPedicularis semibarbata (Scrophulariaceae) fromLupinus fulcratus (Leguminosae) fail to influence the specialist herbivoreEuphydryas editha (Lepidoptera)

Pedicularis semibarbata is apparently an obligate hemiparasite of coniferous trees. It is also a facultative parasite ofLupinus fulcratus from which we find that it obtains quinolizidine alkaloids, principally -isolupanine. As a result, a single population ofP. semibarbata contains both alkaloidrich and alkaloid-free plants. The butterflyEuphydryas editha naturally oviposits on both plant types. This butterfly population, which is the principal herbivore attackingP. semibarbata at this site, is known to contain two morphs. Individuals of a specialist morph discriminate when ovipositing among individualP. semibarbata plants and produce offspring that survive better on accepted than on rejected plants. Those of a generalist morph accept allP. semibarbata plants and produce offspring that survive equally well on plants accepted or rejected by the discriminating morph. Because of the existence of this complex variation among the butterflies, the presence of naturally laid eggs on alkaloid-containing plants still leaves the possibility that the alkaloids may defend the plants against the specialist morph. In experiments on both oviposition preference and larval performance in early instars, we failed to detect any correlation between alkaloid content of a plant and either its acceptability to or suitability for the discriminating morph of the insect. Alkaloid presence in the host-plant population, achieved through root parasitism, is currently neither subject to strong insect-mediated selection nor a major cause of selection on the insects.     

Chemical and ultrastructural analysis of corn cuticular lipids and their effect on feeding by fall armyworm larvae

The cuticular lipid composition of lower and upper leaves of five genotypes of field-grown corn,Zea mays L., was determined by combined gas chromatography-mass spectrometry. Surface lipids of the upper leaves had a higher proportion ofn-alkanes (45–52%) than the lower leaves, while the lower leaves had higher percentages of fatty alcohols (12–18%) than the upper leaves. Scanning electron microscopy showed that the upper leaves of two corn genotypes, MpSWCB-4 and Cacahuacintle X's, had a smooth amorphous appearance, while the lower leaves had a dense array of wax crystals.Spodoptera frugiperda (J.E. Smith) larvae weighed more and developed more rapidly when they were reared on diet containing corn foliage from which the cuticular lipids had been removed than when they were fed untreated foliage. However, growth was not inhibited when larvae were fed diet containing the cuticular lipid extracts or individual cuticular lipid components.     

Colorado Potato Beetle Toxins Revisited: Evidence the Beetle Does Not Sequester Host Plant Glycoalkaloids

The Colorado potato beetle feeds only on glycoalkaloid-laden solanaceous plants, appears to be toxic to predators, and has aposematic coloration, suggesting the beetle may sequester alkaloids from its host plants. This study tested 4th instars and adults, as well as isolated hemolymph and excrement, to determine if the beetles sequester, metabolize, or excrete alkaloids ingested from their host plants. HPLC analysis showed: that neither the larvae nor the adults sequestered either solanine or chaconine from potato foliage; that any alkaloids in the beetles were at concentrations well below 1 ppm; and that alkaloids were found in the excrement of larvae at approximately the same concentrations as in foliage. Analysis of alkaloids in the remains of fed-upon leaflet halves plus excreta during 24 hr feeding by 4th instars, as compared to alkaloids in the uneaten halves of the leaflets, showed that equal amounts of alkaloids were excreted as were ingested. The aposematic coloration probably warns of a previously-identified toxic dipeptide instead of a plant-derived alkaloid, as the Colorado potato beetle appears to excrete, rather than sequester or metabolize, the alkaloids from its host plants.     

Short-term induction of alkaloid production in lupines Differences between N2-fixing and nitrogen-limited plants

We used N₂-fixing and nonfixing lupines to examine the effects of plant nutrition on short-term alkaloid production in damaged leaves. Three different treatments were used: damaged leaves from N₂-fixing plants; undamaged leaves from these damaged, N₂-fixing plants; and damaged leaves on nitrogen-limited, nonfixing plants. Relative to controls, alkaloids increased in concentration more quickly in the N₂-fixing than in the nitrogen-limited plants. The magnitude of this increase in alkaloids was correlated with the initial alkaloid concentration. These results suggest that nitrogen-rich plants may benefit from faster and higher alkaloid induction than nitrogen-limited plants. In addition, the detailed dynamics of individual alkaloids are consistent with earlier proposals for the mechanism of lupine alkaloid induction.     

Variation of Piperidine Alkaloids in Ponderosa (Pinus poderosa) and Lodgepole Pine (P. contorta) Foliage from Central Oregon

We quantified 2,6-disubstituted piperidine alkaloids in Pinus ponderosa and P. contorta needles from three forest sites in April, June, August, and December. Alkaloids were detected on at least one date in 71% of the ponderosa pine and in 29% of the lodgepole pine trees sampled. Pinidine was the major alkaloid constituent of ponderosa pine, while euphococcinine was the predominant compound in lodgepole pine. For ponderosa pine, total alkaloid concentrations were very low at two sites on all dates. At the third site, concentrations were variable but significantly higher on all dates. Total alkaloid concentrations in previous-year foliage from this site were highest in April, then significantly lower from June through December. Current-year foliage collected in August and December had significantly higher alkaloid concentrations than previous-year foliage on the same dates. Variation in foliar nitrogen concentrations accounted for some of the alkaloid variation in current-year foliage sampled in August.