Rastislav Jakuš

Peripheral modulation of pheromone response by inhibitory host compound in a beetle

SUMMARY We identified several compounds, by gas chromatographic–electroantennographic detection (GC–EAD), that were antennally active in the bark beetle Ips typographus and also abundant in beetle-attacked spruce trees. One of them, 1,8-cineole (Ci), strongly inhibited the attraction to pheromone in the field. Single-sensillum recordings (SSRs) previously showed olfactory receptor neurons (ORNs) on I. typographus antennae selectively responding to Ci. All Ci neurons were found within sensilla co-inhabited by a pheromone neuron responding to cis-verbenol (cV); however, in other sensilla, the cV neuron was paired with a neuron not responding to any test odorant. We hypothesized that the colocalization of ORNs had a functional and ecological relevance. We show by SSR that Ci inhibited spontaneous activity of the cV neuron only in sensilla in which the Ci neuron was also present. Using mixtures of cV and Ci, we further show that responses to low doses (1–10 ng) of cV were significantly reduced when the colocalized Ci neuron simultaneously responded to high doses (1–10 μg) of Ci. This indicated that the response of the Ci neuron, rather than ligand–receptor interactions in the cV neuron, caused the inhibition. Moreover, cV neurons paired with Ci neurons were more sensitive to cV alone than the ones paired with the non-responding ORN. Our observations question the traditional view that ORNs within a sensillum function as independent units. The colocalization of ORNs might sharpen adaptive responses to blends of semiochemicals with different ecological significance in the olfactory landscape.

Overview of development of an anti‐attractant based technology for spruce protection against Ips typographus: From past failures to future success

We describe a series of experiments on the protection of Norway spruce (Picea abies (L.) Karst.) logs and standing trees against Ips typographus L. (Coleoptera: Scolytidae). The tested protective materials were applied 1) at point sources and 2) area-covering using zeolite-based dispensers. We performed 11 terrestrial and one aerial application (using a helicopter) between 1992 and 2001. We gradually replaced polyethylene-bag verbenone dispensers by composite dispensers releasing verbenone and non-host volatiles (NHV: green leaf volatiles, trans-conophthorin, and C8-alcohols). Many experiments did not give statistically significant results, due to the low number of true replicates and the spatial heterogeneity of beetle attacks. The protection of horizontal objects like logs or lying trees with anti-attractants was not always successful. In the future, positive results can be expected with an area covering spray applied by zeolite dispensers containing mixtures of verbenone and NHV. A similar approach could probably be used in aerial spraying of anti-attractants.We have succeeded in our attempt to protect zones of standing spruce trees from I. typographus attacks. We used dispensers with a combination of verbenone and a full blend of NHV compounds. The protection is not complete, but in both 2000 and 2001, the treatment significantly decreased the probability of the insect attack on trees by 60–80 %.

A method for the protection of spruce stands againstIps typographus by the use of barriers of pheromone traps in north-eastern Slovakia

A method and layout of various systems of barriers of pheromone traps is described. The rationale of the method is based on the stress of spruce and the chemical communication ofIps typographus. Various systems of pheromone trap barriers were used according to the required level of protection. The method was used in two areas affected by air pollution and bark beetle outbreak. The first area was in an unnatural spruce monoculture region while the second was in a National Park.The described method led to a significant reduction of bark beetle attacks on trees along forest edges and subsequently to the stopping of movement of the zone of attacked trees due to bark beetle damage. In the case of forest stands with no reduced canopy closure, the reduction of attacks led to a dramatic decline of bark beetle outbreak. On the other hand, this method was not able to reduce the level of bark beetle attack in the interior of forest stands with a decreased level of canopy closure. This method, as part of differentiated pest management measures in one forest district, was successful without any other protection measures in the condition of natural reserves in a spruce vegetation zone.

Semiochemical diversity diverts bark beetle attacks from Norway spruce edges

The semiochemical diversity hypothesis (SDH) states that interference with host‐selection from non‐host volatiles (NHV) is an important mechanism for associational resistance. Inhibition of bark beetle attraction to point sources by non‐host volatiles (NHV) is well established and might be a signal serving in host‐selection also at the habitat scale. In forests dominated by Norway spruce in middle and northern Europe (N Slovakia 2006 & 2007, SE Sweden 2007), we applied a blend of NHV and verbenone, released from dispensers fixed at 2 and 6 m height at forest edges with high Ips typographus populations. In Slovakia, three different doses (0.2–0.7 dispensers/m forest edge) were tested in 20‐tree zones of spruce stand edges. The Swedish experiments used only the middle dose. In Slovakia, there was high tree mortality but dispensers with the anti‐attractants reduced killed trees in a dose‐dependent manner. The reduction in tree killing ranged from 35 to 76% compared to untreated zones. Regression analysis of relative tree kill on log dispenser density was highly significant ( = 0.34, corresponding effect size d≈ 0.98). In Sweden, with lower beetle populations, most attacks (99%) were found outside the experimental areas, with high attack rates (15 trees/ha) in a range of 15–30 m from treated groups, indicating an active inhibitory radius exceeding the previous estimates. The SDH as a functional aspect of biodiversity was tested by converting spruce monocultures into an artificial semiochemically mixed forests. The use of NHV provides the only non‐insecticidal method of direct protection of conifer forests. The demonstrated principle of protection is still too expensive for area‐wide use, but viable for high‐value areas (nature reserves). Further development of push–pull strategies or area‐wide applications may prove more cost efficient. In the long‐term, the only sustainable approach is a forest landscape of mixed habitats.

Types of bark beetle (Coleoptera: Scolytidae) infestation in spruce forest stands affected by air pollution, bark beetle outbreak and honey fungus (Armillaria mellea)

Abstract583 spruce stands in an area affected by air pollution and bark beetle outbreak in Eastern Slovakia were studied in 1996. According to bark beetle infestation of dominant and codominant trees, stands were classified into following types of spruce stand decline:Ips typographus-A,Ips typographus-B,Polygraphus poligraphus, I. typographus/P. poligraphus—A,I. typographus/P. poligraphus—B. The presence of attacked trees in forest edges, bark beetle spots and forest interior was the key important factor for the classification.Data from forest inventory and forest management evidence together with data on types of spruce stands decline were used in further analyses. Results shows that the distribution of forest stands classified into different types or uninfested stands is related mainly to host size and site quality. The percentage of spruce, exposition of stands and stand density showed significant effects. The mechanisms of spreading of studied bark beetle outbreak could be explained by direct effects of stress of trees caused by an abrupt increase of level of solar irradiation and by weakening of trees by the honey fungus.

Characteristics of Norway spruce trees (Picea abies) surviving a spruce bark beetle (Ips typographus L.) outbreak

The characteristics of spruce individuals, which survived a massive bark beetle outbreak, were compared with the characteristics of neighbouring attacked trees in Šumava National Park (Czech Republic). Selected parameters related to crown geometry, stand conditions and distances between trees were measured or estimated. Significant differences were found between the surviving trees and the neighbouring trees attacked by I. typographus. Trees with a higher level of stem shading (longer crown length) tended to survive. The attacked trees were usually located in areas with larger basal area, especially southwards from them. A shorter distance to a previously attacked tree increased the probability of additional attack. Spruce trees with more progressive crown structure transformation (primary structure defoliation) were significantly more frequently attacked by spruce bark beetle. Superior and taller trees had a clearly longer life expectancy than dominant ones. These results show that the attack of trees by bark beetle can be predicted to a certain degree, which can be used in management of endangered spruce forests.

Influence of the proportion of (−) α‐pinene in pheromone bait on Ips typographus (Col., Scolytidae) catch in pheromone trap barriers and in single traps

Abstract:  Further possibilities of increasing catch efficiency of Ips typographus L. in pheromone trap barriers with two different levels of release of (4S)‐cis‐verbenol (cV) and in single traps were tested. Pheromone mixtures containing cV, 2‐methyl‐3‐buten‐2‐ol (MB) and 0, 1, 10, 35, 60 and 90% (−) α‐pinene were prepared. Basically, MB was successively replaced by (−) α‐pinene in the mixtures. Tests were performed on the edge of a spruce forest when outbreaks of I. typographus occurred. The substitution of MB with (−) α‐pinene had different effects in different types of pheromone trap barriers and in single traps. It is not possible to increase the attractiveness of the bait having a higher percentage of cV by supplementing a certain percentage of MB with (−) α‐pinene. The catch of I. typographus decreased with an increase in the percentage of (−) α‐pinene in the bait. It is possible to increase the attractiveness of the bait having a lower percentage of cV by supplementing certain percentages of MB with (−) α‐pinene. A slight synergistic effect of (−) α‐pinene with MB/cV mixture is visible in this type of bait. In the case of single traps, supplementing MB with (−) α‐pinene possibly causes a slight synergistic effect. An increase in the percentage of (−) α‐pinene increases the percentage of males in the catch.

Bark beetle (Coleoptera, Scolytidae) outbreak and system of IPM measures in an area affected by intensive forest decline connected with honey fungus (Armillaria sp.)

The character of a bark beetle outbreak, planning system, and implementation of IPM measures in a forest affected by intensive decline connected with honey fungus (Armillaria sp.) is described. It is possible to distinguish two levels of outbreak in the study area. The first level is characterized by spruce mortality connected with yellowing of spruce and presence of plant pathogens, mostly Armillaria sp. Forest stands have disperse infestations of bark beetles. Identification of infested trees and salvage cutting in time are problematic. In case of late processing of infested trees, the second level of outbreak could become manifest. The populations of bark beetle increase. Beetles could attack relatively healthy trees. The classical outbreak with forest edges and spot infestations starts. The identification of infested trees is easier. The application of pheromone trap barriers is effective. Thus, the second level of outbreak could be easily managed. The planning of IPM measures is concerned with localization of various pheromone trap systems in particular forest stands. It is based on field survey, data acquisition, estimation of swarming bark beetle population, and information of pheromone system effectiveness.

Patterns of bark beetle occurrence in Norway spruce stands of national parks in Tatra Mts. in Poland and Slovakia

This paper presents the results of a research project concerning bark beetle populations in Norway spruce stands managed under a nature protection regime in national parks in Tatra Mountains in Poland and Slovakia. The preliminary data on bark beetle occurrence and related selected stand characteristics are presented. The dynamics of a bark beetle outbreak that occurred from 1994–97 in the stands localized on both sides of the state border and in various nature protection regimes is analyzed retrospectively. The data concerning the tree/stand characteristics, i.e. DBH and increment of trees are discussed in relation to its potential role in the mechanism for the spread of bark beetle populations. The patterns of the spatial distribution of trees attacked during the research period (1999–2001) are described.

Can Ips typographus (L.) (Col., Scolytidae) smell the carrion odours of the dead beetles in pheromone traps? Electrophysiological analysis

Abstract:  Head‐space samples of Ips typographus dead beetles taken from pheromone traps were tested by combined gas chromatography and electroantennographic detection (GC–EAD) on the antennae of the same species, and identified by GC‐mass spectroscopy (MS). The GC‐EAD analysis showed that antennae of both sexes responded to 1‐hexanol and verbenone from the aeration samples, while the typical and strong carrion odours from the dead beetles such as dimethyl disulphide, 3‐methyl‐1‐butenol (isoamyl alcohol), 2,5‐dimethyl pyrazine and isovaleric acid elicited no antennal responses. The EAD active compounds, 1‐hexanol and verbenone, have been shown in earlier studies to be inhibitory on attraction of I. typographus to pheromone traps. Thus, the decreased catching efficacy of pheromone traps with many dead beetles might be the result of the release of 1‐hexanol and verbenone from the dead beetles, but not to the typical carrion smells which are strongly detected by the human nose.