Analele ICAS 50:203-212, 2007

INHIBITORY EFFECT OF (-)ALPHA-PINENE HIGH

RELEASE RATES ON IPS TYPOGRAPHUS (L.) RESPONSE
TO ITS AGGREGATION PHEROMONE

NICOLAI OLENICI, MIHAI-LEONARD DUDUMAN,

VALENTINA OLENICI
1

Institutul de Cercetri i Amenajri Silvice, Staiunea Cmpulung Moldovenesc, Romnia

Abstract
The aim of the research presented in this paper was to prove that the effect of (-)alphapinene on the response of Ips typographus (L.) to its aggregation pheromone depends on the release
rates of both pheromone and monoterpene volatiles. Six treatments have been tested in four
experimental areas using six replicates for each treatment: V1 pheromone + (-)alpha-pinen 200
mg/day, V2 pheromone + (-)alpha-pinen 500 mg/day, V3 pheromone + (-)alpha-pinen 2000
mg/day, V4 pheromone, V5 control (blank) and V6 1/3 pheromone dispenser with 1/3 of normal
release rate. In all experimental areas, most captures have been recorded at V4 (pheromone alone),
while alpha-pinene at cca. 200mg/day, 500 mg/day and 2000 mg/day caused the decrease of the
captures by 13.2-43.9%, 22-55.2% and 53.1-70.2% respectively. The traps baited with pheromone
dispensers which were conditioned to have a release rate equal to only 1/3 from the release rate of the
commercial dispensers, caught insects representing about 1/3 of those recorded at the traps with
normal dispensers. The results confirm the hypothesis that the response of Ips typographus beetles to
the combination of synthetic pheromone with (-)-alpha-pinene depends on the concentration of
volatile substances that are released around the source, respectively on the release rates of the
pheromone and monoterpene. They also demonstrate, that at high release rates of the pheromone (cca.
30 mg/day), (-)alpha-pinene released at a rate of 200-2000 mg/day has an inhibitory effect,
determining the decrease of the number of beetles that are caught in the pheromone traps.
Key words: Ips typographus, aggregation pheromone, (-)alpha-pinene, inhibitory effect

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Rezumat
EFECTUL INHIBITOR AL (-)ALFA-PINENULUI ASUPRA RSPUNSULUI
GNDACILOR DE IPS TYPOGRAPHUS (L.) LA FEROMONUL LOR
AGREGATIV N CAZUL RATELOR DE ELIBERARE MARI
Scopul cercetrilor prezentate n aceast lucrare a fost de a demonstra c efectul (-) alfapinenului asupra rspunsului gndacilor de Ips typographus (L.) la feromonul lor agregativ depinde
de intensitatea ambelor semnale chimice, cel feromonal i cel kairomonal, respectiv de ratele de eliberare a feromonului i a monoterpenei. S-au testat ase variante experimentale: V1 feromon +
(-)alfa-pinen 200 mg/zi, V2 feromon + (-)alfa-pinen 500 mg/zi, V3 feromon + (-)alfa-pinen 2000
mg/zi, V4 feromon, V5 martor (neamorsat) i V6 feromon cu rata de eliberare egal cu 1/3 din
rata de eliberare a unui dispenser normal. Experimentul s-a repetat n patru suprafee experimentale i
- n fiecare loc - variantele au fost repetate de ase ori. n toate suprafeele experimentale, majoritatea
capturilor s-au nregistrat la varianta V4 (doar feromon), n timp ce adugarea dispenserilor de alfapinen cu rate de eliberare de cca. 200 mg/zi, 500 mg/zi i 2000 mg/zi a provocat o reducere a
capturilor cu 13,2-43,9%, 22-55,2% i respectiv 53,1-70,2%. Capcanele amorsate cu dispenseri care
au fost condiionai astfel nct s aib o rat de eliberare egal cu 1/3 din cea a dispenserilor
comerciali au avut capturi ce au reprezentat aproximativ o treime din cele obinute la capcanele
amorsate cu dispenseri feromonali normali. Rezultatele experimentului demonstreaz c rspunsul
gndacilor de Ips typographus la combinaia de feromon sintetic cu (-)-alfa-pinen depinde de
concentraia n jurul capcanei a substanelor volatile ce sunt eliberate, respectiv de ratele de eliberare
a feromonului i a monoterpenei. n cazul unor rate mari de eliberare a feromonului (cca. 30 mg/zi),
(-)alfa-pinenul eliberat cu o rat de 200-2000 mg/zi are un efect de inhibare a rspunsului gndacilor,
ceea ce determin o reducere a numrului de capturi n cursele feromonale.
Cuvinte cheie: Ips typographus, feromon agregativ, (-)alfa-pinen, efect inhibitor

1. INTRODUCTION
Research conducted in 1970 revealed the existence of an aggregation
pheromone in Ips typographus (Bakke, 1970; Rudinsky et al., 1971a,b) and the main
components of the pheromone have been identified until 1977 (Vit et al., 1972; Bakke
1976; Bakke et al., 1977). In 1977, Krawielitzki et al. published data which proved that
of the two isomers of verbenol found in spruce bark beetle males, only (-)-(4S) cisverbenol is behaviorally active, while Hackstein & Vit (1978) demonstrated that Ips
typographus beetles produce cis-verbenol from (-)alpha-pinen and trans-verbenol from
(+)alpha-pinene, like Ips paraconfusus Lanier (Renwick et al., 1976). In this way it was
built the knowledge basis for the synthetic pheromone production. Many experiments
have been conducted thereafter in order to ascertain the effect of different
monoterpenes on the response of beetles to the synthetic pheromone, and to find a way
to enhance pheromone attraction. This demarche was justified by previous laboratory
and field results concerning positive reactions of beetles to alpha-pinene, beta-pinene
and limonene (Chararas, 1959; Rudinsky et al., 1971a, 1971b), as well as by the
discovery of the essential part played by alpha-pinene in the biogenesis of cis-verbenol
(Vit & Francke, 1976), one of the main pheromones components.
Bakke et al. (1977) reported that the addition of alpha-pinene, beta-pinene or
myrcene to the mixture of pheromone components had no positive influence on the
response of beetles in the field, but Sauerwein & Vit (1978), adding alpha-pinene to
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TYPOLUR I (a mixture of (S)-cis-verbenol (cV) and methylbutenol (MB) with a ratio

of 1:10), caught more beetles than using traps baited only with pheromone. Tomescu et
al. (1979) also proved, under laboratory conditions, that (-)-alpha-pinene has a synergic
effect in mixture with aggregation pheromone, but a significant increasing of attracted
beetles proportion was obtained only when the pheromone blend contained transverbenol (tV) too, not only cV, ipsdienol (Id) and MB, and the proportions of
substances were 1cV:1tV:1Id:20MB:1aP.
Subsequent experiments involving the synthetic pheromone of I. typographus
and alpha-pinene or host tree volatiles also yielded contradictory results. Vaupel et al.
(1981) found no positive impact of alpha-pinene or spruce resin on captures from the
traps baited with synthetic pheromone, while volatiles from spruce logs increased very
much the number of beetles attracted by Pheroprax (Bombosch, 1983; Bombosch &
Johann, 1985; Johann, 1986). Significant positive effect of volatiles from fresh spruce
slash on Ips typographus catches in pheromone traps have also been noted by Austara
et al. (1986), and the results of Bakke (1985) from spruce clear cuttings of different age
may be interpreted as a differentiated response of bark beetles to host volatiles
abundance too. However, spruce logs did not affect the results when pheromone blend
contained only MB and cV with release rates of 5.8 and 0.05 mg/day respectively
(Schlyter et al., 1987a).
Baiting the pheromone traps with Pheroprax and (-)-alpha-pinene, Reddemann
& Schopf (1996) observed no significant change of captures comparing with traps
having only pheromone dispenser, but the addition of limonene enhanced them by 81%.
Host volatiles from spruce bark had a similar positive effect. Gradual replacing of the
methylbutenol with (-)-alpha-pinene gave different results according to the proportion
of (4S)-cis-verbenol in the pheromone composition, as well as according to the trap
placement (Jaku & Blaenek, 2003).
Recently, Erbilgin et al. (2007), using pheromone dispensers containing only
MB and cV, with a ratio of 6:1, and a very low release rate (0.95 mg/day), obtained an
increase of captures by 100 % when added alpha-pinene so that the ratio between the
release rates of (-)-alpha-pinene and of pheromone was higher than 526:1, but the
captures remained unchanged at a ratio lower than 274:1.
It is obvious that the above mentioned results are so variable mainly due to
experimentation conditions. In some experiments, host volatiles from natural sources
(branches, bark, logs) have been used, and chemical composition as well as release
rates remained unknown. Even the volatile sources made artificially were very diverse
regarding the chemical composition, relative proportion of compounds and the release
rates. In some experiments the pheromone and the monoterpene were released from
separate dispensers, but in other they were mixed and the dispensers have been adjusted to have a certain release rate for the entire bouquet of volatile substances. The
differences between experiments, concerning trap type as well as the distance between
the traps, are other elements that explain the variability of results. If we take into
account the phases of host tree colonization process and the chemical signals used by
insects in each phase (Vit, 1980; Wood, 1982), as well as the permanent change of host
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volatile situation and pheromone production during the colonization process (Birgersson et al., 1984; Schlyter & Lfqvist, 1986; Birgersson & Bergstrm, 1989; Birgersson,
1989; Bohlander & Schopf, 2000), it becomes quite clear that variable concentration of
volatiles around the source induce different beetle responses (Schlyter et al., 1987b;
Reddemann & Schopf, 1996; Jaku & imko, 2000; Erbilgin et al., 2006, 2007;
Franklin & Grgoire, 2001).
Maintaining constant the ratios between pheromone components, the aim of
our research was to prove that: i) the effect of (-)alpha-pinene on the response of Ips
typographus to its aggregation pheromone depends on the release rates of both
pheromone and monoterpene volatiles; ii) at high release rates, alpha-pinene has an
inhibitory effect if pheromone release rate is high, but a synergic effect if pheromone
release rate is low. This paper presents only the results from the experiment with high
release rates of both synthetic pheromone and (-)alpha-pinene.

2. MATERIAL AND METHODS

E x p e r i m e n t l o c a t i o n . The experiment was conducted in three forest
districts (Pojorta, Moldovia and Crlibaba) located in the north-eastern part of Eastern
Carpathians, where the populations of Ips typographus underwent an important
increase during the years 2002-2003 as a consequence of the huge wind throw damages
of March 2002. However, in 2007, when the experiment took place, the population level
was quite low, mainly due to the control measures, but also due to unfavorable weather
conditions prevailing in 2005-2006.
Experimental plots Valea Putnei (forest district Pojorta) and Moldovia were
chosen in old clear cuttings without fresh slash, but with some wind thrown trees from
the last spring in the surrounding tree stands. Instead, the experimental plots Crlibaba
I and Crlibaba II were chosen in fresh clear cuttings, from winter 2006/2007, with high
quantities of fresh slash. In the tree stands bordering the plot Crlibaba I on two parallel
sides, there were also some fresh fallen trees.
E x p e r i m e n t d e s i g n . Six treatments have been tested in each
experimental plot: V1 pheromone + (-)alpha-pinen 200 mg/day; V2 pheromone +
(-)alpha-pinen 500 mg/day; V3 pheromone + (-)alpha-pinen 2000 mg/day; V4
pheromone alone; V5 control (blank); V6 pheromone dispenser having 1/3 of normal
release rate. Six replicates of each treatment were set up.
In the experimental plots Valea Putnei, Moldovia and Crlibaba I, the traps
have been emplaced at about 12-14 m of the forest stand edge, with 15 m distance
between them. The treatments were grouped into blocs, each bloc being a replication.
The distance between the blocs was at least 45 m excepting the experimental plot Crlibaba I, where the distance was of only 15 m due to the scanty space. In the
experimental plot Crlibaba II, the traps were disposed in a grid of squares having the
side of 15 m (4 rows x 9 traps/row).
In order to diminish the influence of trap position on captures, at each
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inspection (every 3 days) the insects were collected and the dispensers have been
moved to the next trap, but the rotation wasnt complete because only three inspections
took place.
D i s p e n s e r s . We used dispensers with synthetic pheromone of Ips
typographus (ATRATYP PLUS), produced by the Chemistry Institute in Cluj-Napoca,
which contain methylbutenol 94.8 %, cis-verbenol 4.7 % and ipsdienol 0.5 %
(M.A.P.D.R.-C.I.O.P.U.F., 2004). Under laboratory conditions (20.5 0.08oC, 58.9
0.3 % HR, without air stream), the pheromone dispensers had a mean release rate of
31.37 0.15 mg/day (mean standard error of mean).
The dispensers with (-)-alpha-pinene have been made of polyethylene foil
(0.100 mm thickness) and cellulose cloth of 5 mm thickness. Those which have been
o
planned to release 200 mg/day (at 20 C) had the size of 38 x 40 mm and were filled with
2.5 ml (-) alpha-pinene, and those planned to release 500 mg/day had 85 x 40 mm and
were filled with 6.5 ml (-) alpha-pinene. In order to obtain a rate of 2000 mg/day, 4
dispensers of 500 mg/day were used together. Before and after their use in the field, the
alpha-pinene dispensers were weighted to ascertain the real release rates of the alphapinene. These have generally been close to those planned (Table 1).
A n a l y s i s o f d a t a . In the experimental areas Valea Putnei, Moldovia and
Crlibaba I, the captures were very variable from one bloc to another (Table 2), while
at Crlibaba II were relatively homogeneous. Accordingly, in the case of the first three
experimental areas, the calculations were made using relative values of the captures
(the percentage represented by the captures of the each variant from the total number
of captures in the bloc) instead of the absolute values. For the statistically analyses, the
data have been transformed in arcsine of square root of percentage. Because not all the
distributions were normal, the significance of differences was evaluated using a oneway analysis of variance (Kruskal-Wallis test) followed by Bonferroni's test for
multiple comparisons.
Table 1. The release rates of the (-)-alfa-pinen in the field conditions
Ratele de eliberare a (-)alfa-pinenului n condiii de teren
The experimental area
Valea Putnei
Moldovi
a
Crlibaba I
Crlibaba II

The release rate (mean SD) at the dispensers of:

200 mg/day
500 mg/day
2000 mg/day
222.5 18.1
519.8 35.2
1865.4 82.7
244.4 2.8
585.7 34.3
2026.2 106.9
155.7 23.9
382.9 58.3
1419.0 148.0
196.7 3.4
464.4 16.2
1864.8 98.2

Table 2. Total number of the captures in the experimental areas V. Putnei, Moldovia and Crlibaba I
Numrul total de capturi n suprafeele experimentale V. Putnei, Moldovia i Crlibaba I
The experimental area
Valea Putnei
Moldovi
a
Crlibaba I

I
924
923
266

The total number of the captures in the block:

II
III
IV
V
VI
524
840
333
296
252
1887
1757
1456
972
1224
452
1581
1133
914
1213

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3. RESULTS
Overall, the most captures were at the 4th variant (the pheromone alone), while
at the treatments V1-V3 (pheromone + alpha-pinene) the number of captures was
smaller and the decreasing of the captures number was progressive from V1 to V3,
respectively along with the increasing of the release rate of the alpha-pinene (Table 3).
A release rate of about 200 mg/day of alpha-pinene caused the decreasing of captures
by 13.2-43.9%, that of 500 mg/day diminished the captures by 22-55.2% and that of
2000 mg/day reduced them by 53.1-70.2%. Because of captures variability, the
differences between V1 and V2, on one side, respectively V4 on the other side, are not
statistically assured in three of the experimental areas, but the differences between V3
and V4 are statistically assured in tree of the four areas. Still, it is remarkable the fact
that - when the traps were concentrated on a relatively small area, with relatively
homogeneous conditions (Carlibaba II) - the difference between the mean values were
statistically assured also in the case of the V1 and V2 variants.
In all experimental areas, the traps baited with Atratyp Plus dispensers which
were conditioned to have a release rate equal to only 1/3 from the release rate of the
commercial dispensers had captures representing about 1/3 from those recorded at the
traps with normal dispensers.

4. DISCUSSION
Because the release rates of the alpha-pinene in the field were close to those
planned for a mean temperature of 20 C, one can assume that the release rate of the
pheromone was also close to that observed in the laboratory (cca. 31 mg/day).
Consequently, the maximum ratio between the release rates of alpha-pinene and
pheromone was about 64:1. At this ratio, the captures of Ips typographus suffered an
evident decrease, statistically assured in three from the four experimental areas. There
are still enough reasons to consider that the beetle response to the pheromone is
o

Table 3. Distribution of Ips typographus captures on treatments

Distribuia capturilor de Ips typographus pe variante experimentale
Experimental
treatment
V1
V2
V3
V4
V5
V6

Valea Putnei I
22,1 5,4ab
17,8 1,9ab
11,7 3,5bc
39,3 11,7a
0,2 0,4c
9,0 2,5bc

Experimental plot1
Argel
Crlibaba I
23,3 5,7ab
24,8 5,4ab
16,1 4,0ab
22,3 5,7ab
11,9 3,9b
13,3 4,7ab
a
36,0 7,2
28,6 5,3a
0,1 0,1c
0,2 0,1c
b
12,6 3,7
10,9 4,2bc

Crlibaba II
179,0 16,3b
170,2 45,7b
107,0 27,7c
255,2 31,9a
3,5 1,2d
93,3 28,4c

Notes: 1) At Valea Putnei I, Argel and Crlibaba I, the values are expressed as percentage, but at Crlibaba II in
numbers of beetles. 2) The means in the same collum followed by the same letter are not different at P = 0.05.

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inhibited also at a lower ratio, respectively at 7:1, if the pheromone concentration is

high. This could mean that usually the beetles do not encounter such situation in the
nature. Responding to the pheromone of pioneer beetles, that are already boring in the
phloem of the host tree, flying beetles attack the tree and begin new galleries. If the
release of pheromone by new attacking beetles increases as a logistic function (Byers,
1989), the highest concentration of the pheromone around the tree will be during the
mass attack phase, when the most beetles are excavating the entrance into phloem or
the nuptial chamber, because in that time (before mating) they have the highest
production of methylbuthenol and cis-verbenol (Birgerson et al., 1984). For an attack
to be successful, beetle concentration and attack of the tree should be faster than tree
defence reactions. Schwerdtfeger (1955) noted that the resin flow decreases very much
in just three days after the beginning of the attack, and in that period a successful attack
is almost complete (Anderbrant et al., 1988). That means that increasing rate of
pheromone release is higher than the increasing of resin exudation rate, until the bark
chemical defences based on resin is overwhelmed. Accordingly, during the mass attack
phase, the ratio between concentrations of monoterpene and pheromone around an
attacked tree is steadily decreasing and presumably much lower than 7:1. The data
-concerning the release rates of volatiles from individual spruce bark beetle entrance
holes (Birgerson & Bergstrm, 1989) confirm this scenario.
The situation is totally different at the beginning of the attack. The trees
naturally release many volatile organic substances, including monoterpenes, but at very
low rates (see, for example, Kesselmeier & Staudt, 1999). However, the emission rates
increase under stress condition (drought, air pollution, diseases, mechanical injuries
etc.). As the pioneer beetles begin the excavation of galleries in the phloem, an
enhanced monoterpene release takes place due to resin exudation. When the entrances
number is still low and the resin flow high, the resin quantity per wound is high.
Consequently, at the beginning of the attack, the ratio between emissions of monoterpene and aggregation pheromone is very high. Data published by Birgerson &
Bergstrm (1989) denote a ratio of about 200:5, and it could be higher, even though not
as high as in the case of Ips pini (Say) when attacking Pinus resinosa (Aitman) trees,
for which Erbilgin & Raffa (2000) and Erbilgin et al. (2003) estimated a value of 2581.
Simulating the situation from the beginning of a beetle attack, Erbilgin et al.,
(2007) obtained an increasing of Ips typographus catches along with the increasing of
(-)-alpha-pinene release rate from about 53 mg/day to about 2465 mg/day, while the
pheromone rate was about 0.95 mg/day. However, it is expected that the level of the
captures will increase still in this situation only as long as the release rate of alphapinene is under a certain threshold, because the monoterpene is an element of the
constitutive and induced defensive systems of the tree against bark beetles and other
organisms (Baier et al., 1999; Franceschi et al., 2005), and at concentration higher than
3 % it is repellent for Ips typographus (Smelyanets & Vasechko, 1973), becoming lethal
for the beetles even at concentration of 0.04 %, if they are forced to stay in an
atmosphere with such a high concentration of (-)-alpha-pinene for 48 hours (Bohlander
& Schopf, 2000). Recent laboratory researches (Faccoli et al., 2005) demonstrated that
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high concentration of alpha-pinene has an obvious antifeedant effect, especially on

male beetles. Conclusively, we can say that the effect of (-)-alpha-pinene on the
response of the beetles to their aggregative pheromone differs according to the release
rate of monoterpene, but also of the pheromone, and this could explain why Niemeyer
& Watzek (1996) failed to reproduce the results of Reddemann & Schopf (1996) when
they used pheromone dispensers with a release rate threefold higher and also
monoterpene dispenser with an enhanced release rate.
On the other hand, our results confirm the fact that the number of Ips
typographus catches increases proportionally to the release rate of the pheromone (Schlyter et al., 1987; Franklin & Grgoire, 2001), as long as the release rate is lower than
50 mg/day, although the proportions between pheromone components were much
different from those used by the above mentioned authors. However, this trend did not
appear in the experiment conducted by Jaku & imko (2000), when IT-ECOLURE
dispensers were used, but the cause is unknown.

4. CONCLUSIONS
The response of the Ips typographus beetles to the combination of synthetic
pheromone with (-)alpha-pinene depends on the concentration of the volatile
substances that are released around the source, respectively on both release rates of the
pheromone and monoterpene. At high release rates of the pheromone (30 mg/day), (-)
alpha-pinene released with at rate of 200-2000 mg/day has an inhibitory effect, causing
the decreasing of captures in the pheromone traps.

ACKNOWLEDGEMENTS
The work was done within the frame of the project: Research concerning the
response of Ips typographus and Hylobius abietis adults to different release rates of
alpha-pinene in combination with the aggregation pheromone, and with ethanol
respectively supported by National University Research Council (CNCSIS) (Contract 56GR/25.05.2007). We thank C. Rotariu, C. urcanu, C. Jolobai and G. Zlei for
allowing us to carry out our experiment in the Crlibaba, Pojorta and Moldovia forest
districts, for their help with pheromone dispensers and pheromone traps, as well as with
labour force for the experiment set up in the field.

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Correspondence to / Coresponden: Nicolai Olenici: olenici.nicolae@icassv.ro

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