Differential toxicity of a phenolic glycoside from quaking aspen to Papilio glaucus butterfly subspecies,hybrids and backcrosses

Summary Papilio glaucus subspecies, hybrids and backcrosses exhibit greatly different abilities to use quaking aspen (Populus tremuloides) and other members of the Salicaceae as host plants. This study was conducted to test the hypotheses that phenolic glycosides account for the differences in larval performance, and that differential performance is correlated with differential larval esterase activities. To test the hypotheses we conducted first instar survival trials and fourth (penultimate) instar feeding trials with tremulacin, a phenolic glycoside. We also conducted assays of -glucosidase, esterase, and glutathione transferase activities, using midgut enzyme preparations from fifth instars. First instar survival on the tremulacin treated diet generally improved with a higher proportion of Papilio glaucus canadensis genes in the genotype, although survival in one backcross treatment was surprisingly low. Penultimate instars of P.g. glaucus and P.g. australis fed tremulacin treated black cherry leaves experienced a severe reduction in growth rate relative to larvae fed control leaves. This seriously suppressed growth was partially due to reduced consumption rates and reduced conversion efficiencies, however, approximate digestibility was not affected. In contrast, P. g. canadensis and hybrids showed no differences in growth rates between tremulacin treated and control leaves. Reciprocal backcrosses of hybrids with P. g. glaucus resulted in slightly suppressed growth on treated versus control leaves. The results suggest that after a certain threshold, increased proportions of P. g. glaucus genes resulted in poorer growth performance with tremulacin in the diet. Soluble esterase activities generally increased with the proportion of Papilio glaucus canadensis genes in the genotype, and paralleled overall trends in larval survival and feeding performance. We conclude that phenolic glycosides such as tremulacin are responsible for differential performance of Papilio glaucus subspecies, hybrids and backcrosses fed plants in the Salicaceae, and that detoxification of phenolic glycosides by midgut esterase explains why some Papilio glaucus genotypes can effectively utilize these plants.     

Origins of the regional feeding abilities in the tiger swallowtail butterfly: ecological monophagy and the Papilio glaucus australis subspecies in Florida

Summary The Highlands County, Florida populations of the eastern tiger swallowtail butterfly, Papilio glaucus L., represent a putative subspecies, P. g. australis Maynard. The only foodplant ever reported in southern half of Florida is sweetbay, Magnolia virginiana L. (Magnoliaceae). In fact, there are apparently no other reported foodplants for the polyphagous tiger swallowtail species which exist in southern Florida. These ecologically monophagous P. g. australis populations were studied in order to determine whether or not significant behavioral and biochemical adaptations in the larvae are detectable as a result of the ecological specialization on sweetbay. These results suggest an emphatic yes. On sweetbay, neonate larval survival of these Florida populations is the best of any other population tested from across North America. In addition, penultimate instar larval growth rates are significantly faster on sweet bay than any other foodplant tested. Furthermore, these biochemical adaptations to sweetbay by P. g. australis in Florida are accompanied by a significantly poorer neonate survival and growth performance on northern foodplants (most notably the Salicaceae and Betulaceae). Reciprocal inabilities of the northern subspecies, P. g. canadensis, and the western P. rutulus to utilize sweetbay are described, as well as the reciprocal inabilities of two congeneric and sympatric (Florida) species, Papilio palamedes and Papilio troilus, that have specialized on another bay (redbay, Persea borbonia; of the Lauraceae). It remains uncertain to what degree such apparent negative genetic correlations in foodplant adaptations represent a driving force in the formation of host races or species, especially since so little is known about oviposition preferences of Papilio glaucus across North America. In Florida we have an apparently strict ecological monophagy coupled with a degree of reproductive isolation that has resulted in its tentative classification as a subspecies, P. g. australis. The role of foodplants in the speciation process can be significant even for P. glaucus, the most polyphagous of all 563 species of world Papilionidae. It remains uncertain whether the significant differences in detoxication and biochemical processing abilities observed in this study were evolved in parapatry or allopatry. Disjunct populations of P. alexiares in Mexico retain some ability to detoxify sweetbay, as do some individuals in most P. glaucus glaucus populations tested from ten different states north of Florida.     

Differential responses of tiger swallowtail subspecies to secondary metabolites from tulip tree and quaking aspen

Summary Two subspecies of the eastern tiger swallowtail butterfly, Papilio glaucus, exhibit reciprocal inabilities to survive and grow on each other's preferred foodplant. P. g. canadensis R. & J. performs well on quaking aspen (Populus tremuloides Michx.) but not on tulip tree (Liriodendron tulipifera L.); P. g. glaucus L. performs well on tulip tree but not on quaking aspen. This study was designed to test the hypothesis that secondary metabolites in tulip tree and quaking aspen are responsible for these differential utilization abilities. We extracted and fractionated leaf constituents into different chemical classes, applied them to a mutually acceptable diet (black cherry, Prunus serotina, leaves), and bioassayed them against neonate larvae (survival) and penultimate instar larvae (survival, growth, digestibility and conversion efficiencies). For each plant species, one fraction in particular showed activity against the unadapted subspecies. One tulip tree fraction dramatically reduced survival of P. g. canadensis neonates, and reduced consumption rates, growth rates, and ECI's of fourth instar larvae. The tulip tree constituents most likely responsible for these effects are sesquiterpene lactones. One quaking aspen fraction greatly lowered survival of P. g. glaucus neonates, and decreased survival, consumption rates, growth rates and ECD's of fourth instar larvae. The compounds responsible for these results are probably simple phenols or phenolic glycosides. Surprisingly, P. g. glaucus and P. g. canadensis showed slightly poorer performance on the active tulip tree and quaking aspen fractions, respectively, indicating that even adapted insects incur a metabolic cost in the processing of their host's phytochemicals.     

Field bioassays show heterospecific mating preference asymmetry between hybridizing North American Papilio butterfly species (Lepidoptera: Papilionidae)

Few studies of interspecific mating preferences of naturally hybridizing species have been done in the field. Yet this is the only way potentially critical habitat-specific factors can be included in mating behavior evaluations. We conducted mate selection (male preference) studies using tethered pairs of heterospecific, size-matched virgin yellow females of Papilio glaucus and P. canadensis with free-flying male P. glaucus populations in south-central Florida, and with free-flying male P. canadensis populations in northern Michigan. Florida males clearly preferred the conspecific P. glaucus females of the pairs, as indicated by the touches/attempted matings and by the actual copulations observed. In 1997, 66.7% of the total attempts (n=168) and 94.2% of all copulations (n=69) were with the conspecific. In 1998 the same pattern of conspecific male preference was observed, with 67.3% of the touches/attempts (n=49) and 100% of the copulations (n=30) being with the P. glaucus females. In contrast, the natural populations of P. canadensis males did not show the expected conspecific preference. In fact, with wild P. canadensis males in 1997 a very strong preference was observed for the heterospecific P. glaucus females of the pairs, accounting for 75.8% of all touches/attempted matings (n=483) and 81.7% of all copulations (n=476). The strong asymmetry in interspecific mating preferences among these hybridizing Papilio species may reflect an ancestral trait in P. glaucus females that elicits strong preference in P. canadensis males, due perhaps to previously possessed male sensory bias or "good genes" indicators in the females. However, reinforcement of reproductive isolation in or near the hybrid zone of species overlap (sympatry) cannot be ruled out. Electronic Publication     

Biochemical detoxication: mechanism of differential tiger swallowtail tolerance to phenolic glycosides

Summary Phenolic glycosides, commonly occurring allelochemicals in the plant family Salicaceae, are differentially toxic to subspecies of the eastern tiger swallowtail and responsible for striking differences in the abilities of Papilio glaucus canadensis and P.g. glaucus to utilize the Salicaceae as food plants. This research was designed to test the hypothesis that particularly high esterase activity confers resistance to phenolic glycosides in P.g. canadensis. I conducted larval survival trials in which the phenolic glycosides salicortin and tremulacin were administered with and without inhibitors of the major detoxication enzymes. Results for P.g. canadensis showed that when esterases were inhibited, toxicity of the phenolic glycosides increased greatly. None of the inhibitors significantly increased toxicity of the compounds to P.g. glaucus. I also conducted in vitro assays of the major detoxication enzymes (polysubstrate monooxygenases, esterases, and glutathione transferases) in larval midguts. Soluble esterase activity was 3-fold higher in P.g. canadensis than in P.g. glaucus. Moreover, esterase activity was inducible by prior consumption of phenolic glycosides in P.g. canadensis but not in P.g. glaucus. Glutathione transferases may also be involved in the terminal metabolism of phenolic glycosides. This is the first verified case of detoxication of an allelochemical by esterase enzymes in herbivores. The biochemical adaptation has played an important role in the evolution of food plant preferences in P. glaucus subspecies.     

Salicaceae detoxification abilities in Florida tiger swallowtail butterflies (Papilio glaucus maynardi Gauthier): Novel ability or Pleistocene holdover?

Abstract Florida populations of the eastern tiger swallowtail butterfly, Papilio glaucus L., have unique morphological features and ecological adaptations that have contributed to their subspecies status (P. g. maynardi Gauthier). We describe geographically unique abilities for detoxification of Carolina willow, Salix caroliniana Michx. (Salicaceae), for several Florida populations of P. g. maynardi. Of all the approximately 570 worldwide species of the Papilionidae, such Salicaceae detoxification abilities exist only in the allopatric North American western and northernmost species (P. rutulus Lucas, P. eurymedon Lucas and P. canadensis Rothschild & Jordan). Females of P. glaucus collected from populations in southeastern USA were examined for oviposition preference in 5‐choice assays, and displayed a low preference for Salicaceae (<5%), but larvae from Florida populations exhibited a high survival (>60%) on these plants. Detoxification abilities have previously shown to be autosomally inherited, and can be transferred via natural or hand‐paired interspecific hybrid introgression. However, these Florida populations are at least 700–1 500 km from the nearest hybrids or the hybrid species, P. appalachiensis Pavulaan & Wright, which possess these detoxification abilities. In any case, the Z ( = X)‐linked oviposition preferences for Salicaceae are lacking in these Florida populations, illustrating genetic independence of oviposition preference determination and larval survival/performance abilities. The origins of detoxification abilities are unlikely to be due to recent climate‐driven introgression, and may represent ancestral trait carry‐overs from interglacial refugium populations of the Pleistocene epoch.     

GENETICS OF MIMICRY IN THE TIGER SWALLOWTAIL BUTTERFLIES,PAPILIO GLAUCUS AND P. CANADENSIS (LEPIDOPTERA: PAPILIONIDAE)

The tiger swallowtail butterfly, Papilio glaucus, exhibits a female-limited polymorphism for Batesian mimicry; the Canadian tiger swallowtail, Papilio canadensis, lacks the mimetic (dark) form entirely. The species hybridize to a limited extent where their ranges overlap. Field collections and censuses indicate that mimetic females occur throughout the range of P. glaucus but at lowest frequencies in populations at the latitudinal edges of its geographic range such as the southernmost part of Florida and along the entire northern edge of its distribution from Massachusetts to Minnesota. Frequencies of mimetic females have remained relatively stable over time. Inheritance of the mimetic form is controlled primarily by two interacting sex-linked loci. The typical matrilineal pattern of inheritance in P. glaucus can be explained by polymorphism at a Y-linked locus, b. Analysis of P. glaucus × P. canadensis crosses has also revealed an X-linked locus, s, which controls the expression of the mimetic phenotype. The P. canadensis allele, s^can, suppresses the mimetic phenotype in hybrid and backcross females. Results from more than 12 yr of rearing tiger swallowtails, including interspecies hybrids, indicate that the absence of mimetic P. canadensis females is due to both a high frequency of the “suppressing” allele s^can and low frequency of the black-pigment-determining b + allele. The frequency of s^can (or other suppressing alleles of s) in P. glaucus populations outside the hybrid zone is low. Some males heterozygous at the s locus and some suppressed mimetic females occur within the hybrid zone. A simple genetic model predicts the frequency of daughters that differ in phenotype from their mothers.     

Hybridization leads to host-use divergence in a polyphagous butterfly sibling species pair

Climate warming has lead to increased genetic introgression across a narrow hybrid zone separating the eastern and Canadian tiger swallowtails (Papilio glaucus and Papilio canadensis). This situation has led to the formation of an allochronically separated hybrid population with a delayed emerging phenotype or “late flight”. Here, we assess how the recombination of the parental genomes that lead to this phenotype may have facilitated another major ecological shift, host-use divergence. We first contrast the ovipositional profiles of the late flight population to that of the parental species P. glaucus and P. canadensis. Subsequently we contrast the larval survival and growth of the late flight, a P. canadensis and a P. glaucus population, and a population from the northern edge of the hybrid zone on five hosts. Our results indicate that the ovipositional preference of this hybrid swarm is identical to that of the introgressing parental species, P. glaucus. Due to the absence of the preferred hosts of P. glaucus (Liriodendron tulipifera L. and Ptelea trifoliata L.) where the late flight occurs, this ovipositional pattern implies a functional specialization onto a secondary host of both parental species, Fraxinus americana L. In contrast, the larval host-use abilities represent a mixture of P. glaucus and P. canadensis, indicating divergence in larval host-use abilities has not taken place. However, high genetic variability (genetic coefficient of variation) is present for growth on F. americana in the late flight hybrid swarm and tradeoffs for larval performance on the preferred hosts of the parental species are evident; indicating a strong potential for future specialization in larval host-use abilities. This current scenario represents an instance where a shift in a major ecological trait, host use, is likely occurring as a byproduct of a shift in an unrelated trait (delayed emergence) leading to partial reproductive isolation.     

MAINTENANCE OF ECOLOGICALLY SIGNIFICANT GENETIC VARIATION IN THE TIGER SWALLOWTAIL BUTTERFLY THROUGH DIFFERENTIAL SELECTION AND GENE FLOW

Differential selection in a heterogeneous environment is thought to promote the maintenance of ecologically significant genetic variation. Variation is maintained when selection is counterbalanced by the homogenizing effects of gene flow and random mating. In this study, we examine the relative importance of differential selection and gene flow in maintaining genetic variation in Papilio glaucus. Differential selection on traits contributing to successful use of host plants (oviposition preference and larval performance) was assessed by comparing the responses of southern Ohio, north central Georgia, and southern Florida populations of P. glaucus to three hosts: Liriodendron tulipifera, Magnolia virginiana, and Prunus serotina. Gene flow among populations was estimated using allozyme frequencies from nine polymorphic loci. Significant genetic differentiation was observed among populations for both oviposition preference and larval performance. This differentiation was interpreted to be the result of selection acting on Florida P. glaucus for enhanced use of Magnolia, the prevalent host in Florida. In contrast, no evidence of population differentiation was revealed by allozyme frequencies. F_ST-values were very small and Nm, an estimate of the relative strengths of gene flow and genetic drift, was large, indicating that genetic exchange among P. glaucus populations is relatively unrestricted. The contrasting patterns of spatial differentiation for host-use traits and lack of differentiation for electrophoretically detectable variation implies that differential selection among populations will be counterbalanced by gene flow, thereby maintaining genetic variation for host-use traits.     

Evolutionary conservatism of oviposition preference in a widespread polyphagous insect herbivore, Papilio zelicaon

We analyzed geographic differentiation in oviposition preference in the anise swallowtail butterfly, Papilio zelicaon Lucas, which is one of the most widely distributed and polyphagous butterflies in western North America. Among 13 populations that span 1200 km of the range of P. zelicaon in the Pacific Northwest of North America, the overall oviposition preference hierarchy has not diverged significantly, even though these populations differ in the plant species they use in the field. The results indicate that differences in host availability and use have not favored major reorganizations in the preference hierarchy of ovipositing females. Instead, this butterfly has a conserved preference hierarchy that varies within a narrow range among populations. All populations ranked the four test plant species in the same overall relative order, even though these populations differ in the plant species they use in the field. Received: 9 February 1996 / Accepted: 24 February 1997     

Effects of plant odor on oviposition by the black swallowtail butterfly,Papilio polyxenes (Lepidoptera: Papilionidae)

Black swallowtail females laid more eggs on plant models treated with contact stimulants and volatiles from carrot leaves than on models treated only with contact stimulants. The volatiles enhanced landing rates and females alighted more frequently on artificial leaves treated with host volatiles than on adjacent control leaves. Volatiles from cabbage, a nonhost, inhibited landing rates on artificial leaves treated with carrot contact stimulants. Examination of antennae revealed two major types of sensilla, believed to be olfactory in function. Electroantennogram preparations responded more strongly to carrot volatiles than to cabbage volatiles and several shared responses at particular retention times to carrot volatile components eluting from a gas chromatograph. Our results are consistent with a long-standing hypothesis that behavioral responses to essential oil components characteristic of the larval food plants have facilitated host shifts in the genus Papilio.     

Inconsistent use of host plants by the Alaskan swallowtail butterfly: adult preference experiments suggest labile oviposition strategy

Abstract 1. The Alaskan swallowtail butterfly (Papilio machaon aliaska) uses three unrelated plant species as hosts: Cnidium cnidiifolium (Apiaceae), Artemisia arctica (Asteraceae), and Petasites frigidus (Asteraceae). The research presented here investigated whether there are any consistent patterns in host choice by P. m. aliaska females. 2. The first two experiments were designed to test if P. m. aliaska host preference is constant or if it changes from day to day. If host preference is labile, the experiments were designed to also test whether a female’s diet breadth narrows or expands over time. 3. The third experiment tested the host preferences of female offspring from several wild-caught P. m. aliaska females. If P. m. aliaska individuals are specialised in their host use, then all of the offspring from a single female would likely prefer the same host-plant species. This experiment was also designed to test the Hopkins’ host selection principle; does the food plant on which a female is reared as a larva influence her future choices when she is searching for host plants for her own offspring? 4. The results from all of these experiments indicate that P. m. aliaska females vary greatly in their oviposition behaviour and in their preferences for the three host plants. Most populations appear to consist of generalists with labile oviposition behaviour. There is no evidence to support the Hopkins’ host selection principle. 5. It is suggested that the generalised selection of host plants by P. m. aliaska females may be a ‘bet-hedging’ strategy and that this strategy may maximise reproductive fitness in an unpredictable environment.     

The chemical world of crucivores: lures, treats and traps

Papilio polytes utilizes only a few plant species of Rutaceae as hosts in the field. We examined in detail the acceptability of Toddalia asiatica (a major host plant) and three other potential rutaceous hosts, Murraya paniculata, Melicope triphylla, and Phellodendron amurense, for ovipositing females of the butterfly. Female responses to the foliage, methanol extracts, and partitioned fractions from these plants were assayed for the presence of oviposition stimulants and/or deterrents. Larval survivorship on these plant species was also compared as an estimate of fitness. The foliage and a methanol extract of T. asiatica readily induced egg-laying, while females responded moderately to the foliage and a methanol extract of P. amurense. By contrast, ovipositing females only marginally accepted Me. triphylla and completely rejected Mu. paniculata. Further experiments to test the biological activity of fractions derived from the respective plant species revealed that T. asiatica contains potent oviposition stimulant(s) and that weak stimulant(s) are present in P. amurense. Poor or negative oviposition responses to both Me. triphylla and Mu. paniculata proved to be attributable to strong deterrent(s) present in these plants. Larvae performed very well on T. asiatica and P. amurense, whereas larval mortality was much higher on Mu. paniculata and Me. triphylla, suggesting the involvement of antifeedant(s) or toxic substance(s) in these plants.     

Within- and among-population variation in oviposition preference for urea-supplemented food inDrosophila melanogaster

Oviposition preference for ureasupplemented food was assayed by simultaneous choice trials on five pairs of closely related laboratory populations of Drosophila melanogaster.Each pair of populations had been derived from a separate ancestral population about 85 generations prior to this study. One population in each pair had been subjected to selection for larval tolerance to the toxic effects of urea; the other population served as a control. Considerable variation in oviposition preference was seen both within and among populations, with four of the ten populations showing a significant mean preference for ureasupplemented food. The degree of specificity shown by individual females was surprisingly high, leading to a bimodal distribution of oviposition preference in some populations. Overall, selection for larval tolerance to urea did not significantly affect oviposition preference. However, the data indicated that pairwise comparisons between randomly selected populations from the two larval selection regimes would lead to a range of possible outcomes, suggesting, in several cases, that selection for larval urea tolerance had led to significant differentiation of adult oviposition preference for urea in one or the other direction. The results, therefore, highlight the importance of population level replication and caution against the practice, common in ecological studies, of assaying oviposition preference in two populations that utilize different hosts in nature, and then drawing broad evolutionary inferences from the results.     

Effects of different leaf traits on growth rates of insect herbivores on willows

We examined relative effects of traits of leaf quality of ten willow species (Salix: Salicaceae) on growth rates of five species of insect herbivores found in interior Alaska (a willow sawfly, Nematus calais; the tiger swallowtail butterfly, Papilio canadensis; and three species of chrysomelid beetles, Gonioctena occidentalis, Calligrapha verrucosa, and Chrysomela falsa). Leaf traits examined were water content, toughness, total nitrogen contnet, pubescence, and presence or absence of phenolic glycosides. Of ten Salix species, four species contain phenolic glycosides in their leaves. We examined relative effects of water content, toughness, and nitrogen content of the Salix leaves on larval growth rates at three different levels, i.e., on a single host species, between different host species, and between herbivore species. The within-host analyses showed that effects of water content, toughness and/or nitrogen content on herbivore growth rates were generally significant in early-season herbivores but not in late-season herbivores. For each herbivore species, differences in growth rates between hosts were not explained by differences in water content, toughness, or nitrogen content. The between-herbivore analysis showed that the interspecific difference in larval growth rates were related to difference in water and nitrogen content of the hosts. Pubescence of Salix leaves had little effects on herbivore growth rates. Presence of phenolic glycosides had a positive effects on growth rates of a specialist, N. calais, but no effect on the other specialist, Ch. falsa. Presence of phenolic glycosides had, in general, negative effects on growth rates of nonspecialists, G. occidentalis, C. verrucosa, and P. canadensis.     

PREFERENCE HIERARCHIES AND THE ORIGIN OF GEOGRAPHIC SPECIALIZATION IN HOST USE IN SWALLOWTAIL BUTTERFLIES

Four allopatric populations of the widely distributed western anise swallowtail butterfly, Papilio zelicaon, use different plant genera as hosts, but simultaneous choice experiments showed that these populations have diverged only slightly in oviposition preference. Of the four populations—two from southeastern Washington State, one from coastal southwestern Washington State, and one from central California—three use hosts that are not available to any of the others. Although variation for the degree of preference for particular plant species occurred within and among populations, all four populations ranked hosts in the same overall order. Monophagy on a local, low-ranking host outside the range of high-ranking hosts did not necessarily lead to the loss of preference for those high-ranking hosts, thereby indicating that the high-ranking hosts would still be accepted, and in some cases even preferred, if a population encountered them again. Hence, the overall preference hierarchy among P. zelicaon populations appears to be evolutionarily conservative. Analyses of differences among families within the California population indicated that increased preference for some hosts is inversely correlated, whereas preference for other hosts may be uncorrelated. Positive correlations may also occur but were not observed among the plant species tested. Overall, the results indicate local monophagy on different plant species in P. zelicaon has not involved major reorganizations in the preference hierarchy of ovipositing females, even in populations that may have fed on a low-ranking host for many generations. Instead, small increases in preference for local hosts have occurred within an evolutionarily conservative preference hierarchy.     

Assessing ecological and physiological costs of melanism in North American Papilio glaucus females: two decades of dark morph frequency declines

Polymorphisms for melanic form of insects may provide various selective advantages. However, melanic alleles may have significant/subtle pleiotrophic “costs.” Several potential pleiotrophic effects of the W (=Y)‐linked melanism gene in Papilio glaucus L. (Lepidoptera) showed no costs for melanic versus yellow in adult size, oviposition preferences, fecundity, egg viability, larval survival/growth rates, cold stress tolerance, or postdiapause emergence times. Sexual selection (males choosing yellow rather than mimetic dark females) had been suggested to provide a balanced polymorphism in P. glaucus, but spermatophore counts in wild females and direct field tethering studies of size‐matched pairs of virgin females (dark and yellow), show that male preferences are random or frequency‐dependent from Florida to Michigan, providing no yellow counter‐advantages. Recent frequency declines of dark (melanic/mimetic) females in P. glaucus populations are shown in several major populations from Florida (27.3°N latitude) to Ohio (38.5° N). Summer temperatures have increased significantly at all these locations during this time (1999–2018), but whether dark morphs may be more vulnerable (in any stage) to such climate warming remains to be determined. Additional potential reasons for the frequency declines in mimetic females are discussed: (i) genetic introgression of Z‐linked melanism suppressor genes from P. canadensis (R & J) and the hybrid species, P. appalachiensis (Pavulaan & Wright), (ii) differential developmental incompatibilities, or Haldane effects, known to occur in hybrids, (iii) selection against intermediately melanic (“dusty”) females (with the W‐linked melanic gene, b+) which higher temperatures can cause.     

In situ predatory behavior ofMysis relicta in Lake Michigan

Selectivity coefficients (W) and predation rates on Lake Michigan zooplankton were determined forMysis relicta during spring through fall using anin situ method. W values indicated the following ranked order of prey preference: Cladocera > copepod copepodites and copepod nauplii > adult diaptomids and cyclopoids. With few exceptions, W values for different prey categories remained fairly constant despite greatly changing relative abundances of prey. Predation rates and prey selectivity were similar in most cases to those determined in laboratory studies. Ingestion rates (percent dry body weight · day^–1) were correlated to total prey biomass (r = 0.38) and to effective prey biomass (r = 0.85), where the weighting factors were overall mean selectivity coefficients for the different prey categories. This result suggested that seasonally varying composition of prey caused much of the variation in ingestion rates among experiments. Feeding trials performed at the same depth with daytime and nighttime assemblages of zooplankton indicated that Cladocera may escape heavyMysis predation at night by migrating from the metalimnetic-hypolimnetic interface into the epilimnion.Contribution 333 from the Great Lakes Research Division, The University of Michigan and contribution 287 from the Great Lakes Environmental Research Laboratory, National Oceanic and Atmospheric Administration.Contribution 333 from the Great Lakes Research Division, The University of Michigan and contribution 287 from the Great Lakes Environmental Research Laboratory, National Oceanic and Atmospheric Administration.     

共生菌通过氧气浓度调控果蝇产卵偏嗜性

【目的】研究东方醋杆菌(Acetobacter orientalis)和短小乳杆菌(Lactobacillus brevis)通过氧气浓度调控黑腹果蝇(Drosophila melanogaster)产卵的偏嗜性,并研究其对后代存活率和发育历期的生物学意义。【方法】利用双相选择装置解析果蝇产卵行为。利用分光光度仪和pH计测量细菌的OD值与pH值。通过改变果蝇的视觉、味觉和嗅觉感官以检测介导果蝇产卵行为的感觉系统。用后代的发育历期与存活率评价其生物学意义。【结果】果蝇产卵对东方醋杆菌具有趋避性,产卵指数为–0.76,对短小乳杆菌具有趋向性,产卵指数为0.5,并且二者接种比例决定了果蝇的产卵选择。氧气作为一个调节因子,可改变果蝇对东方醋杆菌和短小乳杆菌的产卵偏嗜性。氧气是东方醋杆菌生长必需的条件,有氧培养的OD值为1.3,而无氧培养的OD值为0.4。氧气是短小乳杆菌生长非必需的条件,有氧和无氧培养的OD值均为2.2。破坏嗅觉可严重地影响果蝇对细菌的产卵偏嗜性。东方醋杆菌和短小乳杆菌均促进了果蝇的生长,分别提高存活率1倍和1.5倍,分别缩短果蝇发育历期1 d和2 d。【结论】东方醋杆菌和短小乳杆菌可影响果蝇产卵偏嗜性,并且二者比例决定了果蝇的最终产卵选择。东方醋杆菌和短小乳杆菌通过感知氧气浓度的改变而进行不同的代谢方式,从而进一步调节果蝇产卵选择。嗅觉系统介导果蝇对东方醋杆菌和短小乳杆菌的产卵选择行为,从而提高后代存活率和缩短发育时间。     

Food utilization and antioxidant enzyme activities of black swallowtail in response to plant phototoxins

Phototoxic compounds are widely distributed among plant families; due to their ability to bind covalently to DNA or to react with oxygen and generate toxic oxyradicals, these compounds are toxic to a variety of herbivorous organisms. Black swallowtail (Papilio polyxenes) larvae feed exclusively on phototoxic host plants in the Apiaceae and Rutaceae. In this study, we examined the toxicity of four phototoxins—three furanocoumarins and one β-carboline alkaloid—to P. polyxenes, as well as the inducibility of antioxidant enzyme defenses in response to these phototoxins. Neither the furanocoumarins nor the β-carboline alkaloid demonstrated any toxic effect on digestive efficiencies of P. polyxenes in the presence of light; harmine, the alkaloid, did significantly reduce growth and consumption rates. None of the compounds had a significant effect on antioxidant enzyme levels. These findings contrast with those reported in earlier studies for Trichoplusia ni, a generalist noctuid sensitive to both furanocoumarins and β-carboline alkaloids. Greater detoxicative metabolic capabilities, coupled with substantially higher constitutive levels of antioxidant enzyme activity, likely explain at least in part the absence of induced antioxidant enzyme responses in the specialist feeder on phototoxic plants. © 1993 Wiley-Liss, Inc.