Chemoreception of host volatiles in the bark beetleIps typographus

Summary In the bark beetleIps typographus the stimulating effects of host odour constituents were studied by linking a gas Chromatograph (GC) with electrophysiological recordings from single olfactory receptor cells. Cells were exposed to the complete aroma of natural bark, extracts from host and non-host trees and fractions of these stimuli separated via a GC column. Four minor fractions belonging to two extracts were found to excite olfactory receptor cells. Responsive cells were divided into two main groups reacting exclusively to one fraction and suggesting that the cells were highly specialized to a particular host odour constituent. However, an effective extract or fraction could not be found for half of the olfactory units responding to natural bark, indicating that the relevant stimulatory compounds were lost during the extraction and/or GC procedure.Abbreviations EAG electroantennogram - GC gas-chromatograph     

Effects of bark beetle pheromones on the attraction of Monochamus alternatus to pine volatiles

Abstract We evaluated the attraction of Monochamus alternatus Hope (Coleoptera: Cerambycidae), Dryocoetes luteus Blandford and Orthotomicus erosus Wollaston (Coleoptera: Curculionidae: Scolytinae) to multiple-funnel traps baited with the pine volatiles, ethanol and (+)-α-pinene and the bark beetle pheromones, ipsenol and ipsdienol. M. alternatus were attracted to traps baited with ethanol and (+)-α-pinene but not those baited with ipsdienol and ipsenol. Ipsdienol and ipsenol decreased catches of M. alternatus in traps baited with ethanol and (+)-α-pinene. Traps baited with either binary combinations of ethanol and (+)-α-pinene or ipsdienol and ipsenol were attractive to D. luteus and O. erosus. The addition of ipsenol and ipsdienol to traps baited with ethanol and (+)-α-pinene synergized attraction of O. erosus but not D. luteus.     

Differences in photoperiod‐induced diapause plasticity among different populations of the bark beetle Ips typographus and its predator Thanasimus formicarius

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Reemergence and second brood in the bark beetle Ips typographus

The knowledge about reemergence of parent spruce bark beetles Ips typographus . their dispersal, and production of a second brood is reviewed. A majority of the beetles reemerge after their first brood. The process is mainly determined by temperature but high breeding density decreases the average time spent in the tree. The difference between males and females in reemergence seems to be small. In the field, a positive relationship between residence time and fat content at reemergence seems to exist, whereas the opposite tendency is found under laboratory conditions. In the laboratory, the survival of beetles reemerging late is on the average lower than that for early reemerging beetles. The time of reemergence and size of the first brood do not, however, seem to influence the production of a second brood. A large proportion of the reemerging beetles are able to establish a new brood in the laboratory, but calculations based on the number of attacked trees and estimated reemergence in forests in South Norway suggest that only about one third of the beetles actually produce a second brood. In northern Europe, successful colonization of new trees by the reemerging beetles seems to require a period of several warm days during the main reemergence period.     

Quantification of different yeasts associated with the bark beetle,Ips typographus,during its attack on a spruce tree

There were different amounts and types of yeasts associated with individuals ofIps typographus spruce bark beetles during different phases of their attack on a healthy spruce tree. The yeasts were isolated on Sabouraud agar medium in order to identify them and estimate their numbers.Hansenula holstii andCandida diddensii type yeasts were most frequently isolated. The increase in number of these two yeast types probably accounted for most of the total yeast increase found during the later attack phases of the bark beetles. Lesser amounts ofHansenula capsulata, Pichia pinus, Candida nitratophila, and twoCryptococcus type yeasts were also found.     

Host resistance elicited by methyl jasmonate reduces emission of aggregation pheromones by the spruce bark beetle, Ips typographus

We treated Norway spruce (Picea abies) stems with methyl jasmonate (MeJA) to determine possible quantitative and qualitative effects of induced tree defenses on pheromone emission by the spruce bark beetle Ips typographus. We measured the amounts of 2-methyl-3-buten-2-ol and (S)-cis-verbenol, the two main components of the beetle's aggregation pheromone, released from beetle entrance holes, along with phloem terpene content and beetle performance in MeJA-treated and untreated Norway spruce logs. As expected, phloem terpene levels were higher and beetle tunnel length was shorter (an indication of poor performance) in MeJA-treated logs relative to untreated logs. Parallel to the higher phloem terpene content and poorer beetle performance, beetles in MeJA-treated logs released significantly less 2-methyl-3-buten-2-ol and (S)-cis-verbenol, and the ratio between the two pheromone components was significantly altered. These results suggest that host resistance elicited by MeJA application reduces pheromone emission by I. typographus and alters the critical ratio between the two main pheromone components needed to elicit aggregation. The results also provide a mechanistic explanation for the reduced performance and attractivity observed in earlier studies when bark beetles colonize trees with elicited host defenses, and extend our understanding of the ecological functions of conifer resistance against bark beetles.     

Behavioural sequence in the attraction of the bark beetle Ips typographus to pheromone sources

ABSTRACT. The functions of the two synergistic pheromone components, (4S)- cis -verbenol (cV) and 2-methyl-3-buten-2-01 (MB), and the role of ipsdienol in the attraction of Ips typographus (L.) (Scolytidae) to pheromone sources were studied in the field. Absolute and relative beetle catches were compared between several traps placed at and nearby a central pheromone source: a pipe trap containing the source, a surrounding sticky trap, a nearby window trap, and four distant window traps. A higher catch in the outer down-wind distant traps indicated an up-wind anemotaxis to the source. Increased MB release, with cV constant, increased the proportion caught in the central pipe trap, indicating MB as a landing stimulus. Release of MB alone gave a very small catch. Ipsdienol could not substitute for cV in the synergism with MB. An increase of cV, with MB constant, increased the number of beetles caught, but not the proportion caught in the pipe trap. The sex ratio was equal in the window traps, but fell to 30% males in both sticky and pipe traps, showing that a large proportion of the males attracted to the source did not land. The proportion of males in the pipe trap was reduced at the highest cV dose. The results support the idea of each pheromone component having a different relative importance in releasing different steps in the behavioural chain.     

Redundancy, synergism, and active inhibitory range of non-host volatiles in reducing pheromone attraction in European spruce bark beetle Ips typographus

Brandt's voles ( Microtus brandti ) are small native mammals that inhabit the grasslands of Inner Mongolia, China. The species is considered a pest, particularly during population outbreaks, which have increased in frequency since 1970 from 1 every 7 years to 1 every 3 to 5 years. Using historical records taken between 1948 and 1998, we found that there was significant correlation between years for which the monthly averages of the Southern Oscillation Index (SOI) were consistently high, and years in which outbreaks occurred. Also there was a tendency for outbreaks not to occur in years with precipitation above or below average. For some years at some sites, annual or biannual estimates are available for the density of Brandt's vole populations. We found that the seasonal rate of increase over the non-breeding season from autumn to spring was negatively correlated with the density in autumn. The rate of increase over the breeding season from spring to autumn was independent of the density in spring but instead, reflects the species' preference for habitat with short, sparse grass; populations do not persist in highly degraded grasslands or in tall, dense grassland. The link between outbreaks and climatic indices, and the numerical response of Brandt's voles to particular habitat characteristics, suggest that current grazing intensity tends to maintain grass at low height in years with average precipitation. The substantial increase in livestock numbers over the last 50 years appears to have increased the number of years when the balance of grazing and plant growth favours high rates of increase in Brandt's vole populations, and hence, there has been an increase in the frequency of outbreaks.     

The role of beetle and host volatiles in host colonization in the European oak bark beetle, Scolytus intricatus (Ratzeburg) (Col., Scolytidae)

Abstract:  To understand the role of chemical signals involved in Scolytus intricatus (Ratzeburg) (Col., Scolytidae) host colonization, the attractiveness of intact and beetle colonized host material was investigated in the field and in the laboratory. In the field, chemical signals operating at long range were investigated by means of sticky traps. In the laboratory, close-range chemical interactions were investigated with an arena olfactometer. Field experiments showed no differences in the attractiveness between infested and non-infested host material. On the contrary, laboratory experiments revealed higher attractiveness of infested host and beetle-produced frass in comparison with intact host and mechanically produced sawdust respectively. Laboratory data also disclosed the attractiveness of beetle extracts of both sexes. Our data show that: (1) host kairomones play an important role during host colonization, and that (2) S. intricatus does not use a sex-specific secondary attractant system. Differences between results of field and laboratory trials are discussed.     

Behavioural and electrophysiological responses of the bark beetle, Ips typographus, to potential pheromone components

ABSTRACT. Field tests showed ( S )-(-)- cis -verbenol and ( R )-(+)- trans -verbenol in combination with a second pheromonal component, 2-methyl-3-buten-2-ol, more attractive than the combination with their optical antipodes. Inhibition of response to the attractant component, ( S )-(-)- cis -verbenol, by high concentrations of its optical antipode did not occur. No significant differences were noted for response to the attractant, ( S )-(-)- cis -verbenol and 2-methyl-3-buten-2-ol, with the addition of either ipsdienol enantiomer or a racemic mixture of ipsdienol enantiomers. Electroantennogram (EAG) studies correlated well with the behavioural studies. EAGs recorded from male and female beetles revealed both sexes to have a lower threshold for the pheromone, ( S )-(-)- cis -verbenol, than its host terpene precursor, (-)- alpha -pinene. EAGs showed a greater number of acceptors for (-)- alpha-pinene in males than in females. EAGs at acceptor saturation to the enantiomers of alpha -pinene and the verbenol isomers showed males more responsive to (-)- alpha -pinene, (±)- cis -verbenol, and ( R )-(+)- trans -verbenol. Significantly greater EAGs were elicited in females than in males to (-)- alpha -pinene, and (±)- and ( S )-(-)- cis -verbenol. No significant differences in EAGs of females to the enantiomers of trans -verbenol were noted. EAGs showed similar thresholds in males and females to the pheromone component, 2-methyl-3-buten-2-ol; however, female response at threshold was significantly greater than male response. The results are discussed with regard to olfactory acceptor evolution.     

Fungal associates of the tree-killing bark beetle,Ips typographus,vary in virulence,ability to degrade conifer phenolics and influence bark beetle tunneling behavior

The bark beetle Ips typographus carries numerous fungi that could be assisting the beetle in colonizing live Norway spruce (Picea abies) trees. Phenolic defenses in spruce phloem are degraded by the beetle's major tree-killing fungus Endoconidiophora polonica, but it is unknown if other beetle associates can also catabolize these compounds. We compared the ability of five fungi commonly associated with I. typographus to degrade phenolic compounds in Norway spruce phloem. Grosmannia penicillata and Grosmannia europhioides were able to degrade stilbenes and flavonoids faster than E. polonica and grow on minimal growth medium with spruce bark constituents as the only nutrients. Furthermore, beetles avoided medium amended with phenolics but marginally preferred medium colonized by fungi. Taken together our results show that different bark beetle-associated fungi have complementary roles in degrading host metabolites and thus might improve this insect's persistence in well defended host tissues.     

Attraction of the cerambycid beetles Tetropium gabrieli,T. castaneum and T. fuscum to pheromones and host tree volatiles

1. Tetropium gabrieli is native to the European Alps where it breeds in European larch and may be an important pest during drought periods by killing trees. It has spread to larch plantations in several European countries including Sweden.
2. In trapping studies conducted in Sweden, we tested whether T. gabrieli was attracted to E-fuscumol, E,Z-fuscumol, E,Z-fuscumol acetate and the host volatile combination of α-pinene plus ethanol. We also compared its response with those of the native congeners, T. fuscum and T. castaneum.
3. All three Tetropium species were caught in significantly higher numbers in traps baited with E-fuscumol plus host volatiles than in traps baited with host volatiles alone or in unbaited traps.
4. There was no difference in catch of any Tetropium species between traps baited with E-fuscumol plus host volatiles versus traps baited with E,Z-fuscumol plus host volatiles, but the latter treatment caught more T. fuscum and T. castaneum than did E,Z-fuscumol acetate plus host volatiles.
5. The addition of E-fuscumol to host volatile-baited traps significantly reduced mean catch of T. gabrieli, significantly increased mean catch of T. castaneum, and had no significant effect on catch of T. fuscum. Mean catch of all three species was significantly greater in traps baited with E-fuscumol alone than in traps baited with host volatiles alone.
6. All three species were caught from May to August. The median date of captures was about 2 weeks earlier for T. fuscum compared with the other two species.

     

Biological differences reflect host preference in two parasitoids attacking the bark beetle Ips typographus (Coleoptera: Scolytidae) in Belgium

The basic reproductive biology of two ectoparasitoids developing on the late larval instars of the scolytid Ips typographus Linnaeus, a pest of spruce forests in Eurasia, was studied with the purpose of explaining which biological features allow the two species to share the same host. The anautogenous braconid Coeloides bostrichorum Giraud had a longer pre-oviposition period (5.1 vs. 3.3 days), a greater egg load (8.1 vs. 6.1 eggs), survived longer and emerged later than the pteromalid Rhopalicus tutela (Walker). In contrast, R. tutela was autogenous and tended to be more fecund under constrained conditions (9.7 vs. 5.1 total offspring per female). The longer pre-oviposition period of the specialist C. bostrichorum, coupled with its greater longevity, afforded the opportunity of better synchronization of ovipositing females with late instar larvae of I. typographus. By contrast, the polyphagous R. tutela matured rapidly, allowing parasitism of both younger and older larval instars of I. typographus in addition to other species. These small differences favoured the co-occurrence of the two parasitoid species on the same attacked trees.     

Behavioural response of the fruit tree bark beetle, Scolytus mediterraneus, to host and non-host plants

The response of beetles to various host extracts obtained by different solvents of a wide range of polarity have been examined by a bioassay method. Ether as a low-polar solvent was more effective than ethanol. The effectiveness of the ethanol decreases with an increase in its polarity.Extracts from infested hosts elicited a greater response than those from uninfested hosts. This has been demonstrated by ether and ethanol extracts, with plum and apricot hosts. Ether extract from an infested host was about four times more active than that from an uninfested host. These results were in accordance with those obtained in situ in which beetles were attracted to infested twigs more strongly than to uninfested twigs. An extract from a non-host plant (orange) was ineffective and its response value resembled that of untreated control.There is a strong indication that after an initial attack, beetle secretions together with plant material are responsible for a mass attraction of the beetles in the field.

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Résumé La réaction de ce Scolyte envers des extraits de plantes-hôtes obtenus au moyen de solvants de polarité très différente a été examinée. L'éther a été plus efficace que l'éthanol dont l'efficacité décroît inversement à sa polarité.Les extraits d'hôtes attaqués par l'insecte provoquaient une réaction plus vive que ceux d'hôtes non infestés. Ceci fut démontré avec des extraits éthériques et éthanoliques de prunier et d'abricotier.Les extraits éthériques d'hôtes attaqués étaient environ quatre fois plus actifs que ceux d'hôtes non infestés. Les résultats s'accordent avec ceux obtenus in situ, l'insecte étant attiré plus fortement par des branches infestées que par celles non attaquées.D'apr`es nos résultats il semble que dans le verger la sécrétion du scolyte associée à la matière végétale provoquent ensemble l'attaque massive.

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Contribution from The Volcani Institute of Agricultural Research, Bet Dagan, Israel. 1971 series; No. 1898 E.     

Flight propensity and diapause incidence in five populations of the bark beetle Ips typographus in Scandinavia

Emergence from brood logs, take-off activity and the duration of tethered flight was recorded in the first laboratory generation of the bark beetle Ips typographus L. (Coleoptera: Scolytidae) from five locations (ca 300 km apart) in Sweden and Denmark. Beetles of northerly origin emerged later from brood logs. This pattern was associated with an increasing proportion on non-fliers towards the north. After overwintering, both the rate of development of flight activity and the proportion of non-fliers were the same among populations. Flight duration of fliers was similar among populations and appeared unaffected by outbreak conditions.     

The role of semiochemicals in tritrophic interactions between the spruce bark beetle Ips typographus, its predators and infested spruce

Abstract:  Semiochemical interactions between the spruce bark beetle Ips typographus , its predators Medetera setiventris , Thanasimus formicarius and Thanasimus femoralis , and the host Norway spruce, Picea abies , were studied in the field. The chemicals S - cis -verbenol, 2-methyl-3-buten-2-ol, ipsdienol, (+)- α -pinene, (−)- α -pinene, (±)- α -pinene, limonene, camphor and their naturally occuring mixtures were used as trap baits in a multiple-choice design that allowed for comparison of their attractivity for the focal species. Medetera was attracted to both the prey aggregation pheromone and its multifunctional component, ipsdienol. On the contrary, both Thanasimus species responded predominantly to ipsdienol and less to the prey aggregation pheromone. In the case of I. typographus , the attractivity of aggregation pheromone seems to be increased by the addition of a mixture of monoterpenic tree volatiles, and by addition of ipsdienol. Bark beetles and predators showed species-specific responses to volatile mixtures representing different stages of tree decay and different stages of bark beetle colony establishment. These responses correlates with the optimal foraging habitat of each species. None of the predator species responded to 2-methyl-3-buten-2-ol, a substantial component of I. typographus pheromonal bouquet, thus it is hypothesized that only substances of monoterpenic origin attract predators.     

Effects of photoperiod and temperature on the development and diapause of the bark beetle Ips typographus

Abstract:  Diapause was induced in a Central European population of Ips typographus grown at 20°C when the day length decreased below 16 h [50% diapause incidence occurred in the 14.7:9.3 h L:D (light:dark) regime]. The non-diapausing adults fed on days 2–6 and 10–14 after the ecdysis and swarmed after the second feeding bout with chorionated eggs in the ovaries and sperm in the spermiducts. Neither gonads nor the flight muscles matured and no swarming occurred in the diapausing adults. The development from egg to adult took about 34 days in both 18:6 h (no diapause) and 12:12 h L:D (diapause) regimes, but it was extended by up to 30% without diapause induction when only larvae or pupae were exposed to L:D 12:12 h. Diapause was induced in insects reared at L:D 12:12 h through the last larval and the pupal instars and/or in the adult stage. Temperature ≥ 23°C prevented diapause induction at L:D 12:12 h but diapause occurred at L:D 14:10 h associated with 26:6°C thermoperiod. The effect of thermoperiods on the developmental rate requires further research. Exposure of the non-diapausing adults to 5°C for several days blocked feeding and evoked a diapause-like state, whereas diapausing adults fed and their gonads slowly developed at this temperature. Diapausing adults exposed in forest to low night temperatures and transferred in October to 20°C readily reproduced at 18:6, but not 12:12 h L:D photoperiods. After 2-months at 5°C and darkness, they became insensitive to the photoperiod, matured and most of them also swarmed at 20°C in the 12:12 h L:D regime. In a Scandinavian population, diapause occurred at 18:6 h L:D and was terminated either by exposure to 5°C or by very long photoperiod (L:D 20:4 h) combined with high temperature (23°C).     

Multiple component system of sugars and polyols in the overwintering spruce bark beetle, Ips typographus

Overwintering adults of the spruce bark beetle, Ips typographus (L.) showed an unusually complex sugar/polyol cryoprotectant system. The major components of the multiple system were: glucose (177.6 mmolL(-1), March); trehalose (175.0 mmolL(-1), December); sorbitol (147.9 mmolL(-1), January); mannitol (81.2 mmolL(-1), March); and erythritol (40.7mmolL(-1), March) (in the parentheses, the maximum concentrations are shown and the month when they were reached). Other minor components were glycerol, fructose, threitol, myo-inositol, arabinitol and ribitol. Distinct seasonal patterns of accumulation/depletion in various components were found. Glycerol, trehalose and glucose started to accumulate first, during early autumn, when the air temperatures fluctuated between 20 and 0 degrees C, and diapause beetles continued in feeding. Glycerol was depleted, glucose remained stable and trehalose continued in accumulation during late autumn when the temperatures oscillated around 0 degrees C. During early winter severe frosts reaching -20 degrees C came, the beetles terminated their diapause and trehalose was partially depleted, while mannitol, sorbitol, fructose, threitol and erythritol started to accumulate. Cold weather continued also during late winter when the beetles remained quiescent. During this period, trehalose was re-accumulated, threitol and erythritol continued to increase, mannitol remained stable and sorbitol, fructose decreased. All cryoprotectans were finally cleared in the beetles which were spontaneously leaving bark during early spring. The seasonal maximum of total concentration of all cryoprotectants (578.2 mOsmol L(-1)) was reached in March. Such a concentration results in colligative depression of melting point of body fluids down by 1.08 degrees C only. It suggests that the potential cryoprotective effect of accumulated sugars and polyols was related rather to their non-colligative functions.     

Exogenous application of methyl jasmonate elicits defenses in Norway spruce (Picea abies) and reduces host colonization by the bark beetle Ips typographus

The terpenoid and phenolic constituents of conifers have been implicated in protecting trees from infestation by bark beetles and phytopathogenic fungi, but it has been difficult to prove these defensive roles under natural conditions. We used methyl jasmonate, a well-known inducer of plant defense responses, to manipulate the biochemistry and anatomy of mature Picea abies (Norway spruce) trees and to test their resistance to attack by Ips typographus (the spruce bark beetle). Bark sections of P. abies treated with methyl jasmonate had significantly less I. typographus colonization than bark sections in the controls and exhibited shorter parental galleries and fewer eggs had been deposited. The numbers of beetles that emerged and mean dry weight per beetle were also significantly lower in methyl jasmonate-treated bark. In addition, fewer beetles were attracted to conspecifics tunneling in methyl jasmonate-treated bark. Stem sections of P. abies treated with methyl jasmonate had an increased number of traumatic resin ducts and a higher concentration of terpenes than untreated sections, whereas the concentration of soluble phenolics did not differ between treatments. The increased amount of terpenoid resin present in methyl jasmonate-treated bark could be directly responsible for the observed decrease in I. typographus colonization and reproduction.