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Biosysteras Diversity ; , , • Biosystems
B ^ ■ fi H Diversity
ISSN 2519-8513 (Print) ISSN 2520-2529 (Online) Biosyst. Divers., 2019, 27(2), 177-185 doi: 10.15421/011924
Influence of essential oils of plants on the migration activity of Tribolium confusum (Coleoptera, Tenebrionidae)
V. O. Martynov, O. G. Titov, T. M. Kolombar, V. V. Brygadyrenko
Oles Honchar Dnipro National University, Dnipro, Ukraine
Article info
Received 04.04.2019 Received in revised form
03.05.2019 Accepted 06.05.2019
Oles Honchar Dnipro National University, Gagarin ave., 72, Dnipro, 49010, Ukraine. Tel.: +38-050-93-90-788. E-mail: [email protected]
Martynov, V. O., Titov, O. G., Kolombar, T. M., & Brygadyrenko, V. V. (2019). Influence of essential oils of plants on the migration activity of Tribolium confusum (Coleoptera, Tenebrionidae). Biosystems Diversity, 27(2), 177-185. doi:10.15421/011924
Pest control should be ecologically-based, therefore use of ecologically safe approaches is the best variant. Essential oils of plants can affect the main metabolic, biochemical, physiological and behavioural functions of insects. In the experiment, we evaluated the influence of 20 essential oils on migration activity of imagoes of Tribolium confusum Jacquelin du Val, 1863 in the conditions of a laboratory experiment. Notable repellent activity against T. confusum was exhibited by essential oils of Jasminum officinale and Thuja occidentalis. Essential oils of Zingiber officinale and Cedrus atlantica had an attractant effect on imagoes of T. confusum. Essential oils of Rosmarinus officinalis, Melaleuca alternifolia, Lavandula angustifolia and Cinnamomum verum exhibited repellent properties while essential oils of Juniperus communis and Citrus sinensis had an attractant effect on the pests. Therefore, out of 20 studied essential oils, only four samples had notable biological effect on migration activity of T. confusum imagoes. These data indicate the possibility of using essential oils or their main components as ecologically safe natural repellents against pests of stored wheat and products of its processing.
Keywords: pest control; biopesticides; plant protection; repellent; attractant.
Introduction
Taking measures against pests should be ecologically-based (EBIPM) and be undertaken in the context of integrated management of agricultural crops and complex control of pests, which means that use of ecologically safe methods is the best variant (Koul & Walia, 2009). Over the past 50 years, pest control in the agriculture has been based on using synthetic chemical insecticides in field agrocenoses and in conditions of greenhouse cultivation.
However, synthetic insecticides are toxic, they have unfavourable effects on the environment, polluting soil, water and air, and also when broadly used provoke development of resistance of target species and significant damage to populations of non-target species of invertebrates (Benhalima et al., 2004; Pimentel et al., 2009; Brygadyrenko & Ivany-shyn, 2015; Martynov & Brygadyrenko, 2017, 2018; Shulman et al., 2017). Furthermore, synthetic insecticides negatively affect human health; strict ecological regulation of using pesticides has led to increase in the number of studies on use of natural plant extracts as alternative synthetic preparations (Isman, 2004; Pérez et al., 2010).
There are 17,500 species of aromatic plants and around 300 essential oils that are commercially valuable for cosmetics, the pharmaceutical and food industries (Bakkali et al., 2008; Pushpanathan et al., 2008; Ebadollahi et al., 2015). Over 2,000 species of plants have insecticide activity (Klocke, 1989). Many commercial essential oils are included in the Generally Recognized as Safe List, which is fully recognized by Environmental Protection Agency and Food and Drug Administration in the USA (Burt, 2004).
Essential oils are secondary metabolites and are present in all parts of plants. They are complex compounds which contain many components which determine the properties of the essential oils. Among the components, there are terpenes, aromatic and aliphatic compounds. The main terpenes are monoterpenes and sesquiterpenes (Bakkali et al., 2008; Koul et al., 2008). Monoterpenes make up to 90% of essential oils and are represented by compounds different in structure: acyclic
(geraniol) and cyclic (terpineol) spirits, phenols (thymol), ketones (thu-jone), aldehydes (citronellal), acids (chrysanthemic acid) and oxides (1,8-cineole). Aromatic compounds, such as cinnamaldehyde, chavicol, anethole, safrole and apiole, are derivatives of phenylpropane and are present in lower content (Isman, 2006; Tripathi et al., 2009).
Essential oils affect the main metabolic, biochemical, physiological and behavioural functions of insects (Mann & Kaufman, 2012), and can also block respiratory tracts and lead to asphyxiation and death of pests (Kaufmann & Briegel, 2004; Rotimi et al., 2011). They can have toxic, fumigative, repellent, antifeedant, ovicidal, attractant and other effects (Werdin-Gonzalez et al., 2008). A number of scientists (Isman, 2000; Gutierrez et al., 2009) have reported neurotoxic, cytotoxic, phototoxic and mutagenic activity of essential oils on insects.
Botanical insecticides have a number of advantages: they do not persist in the environment, pose relatively low risk for non-target organisms (useful predators and parasites) and are relatively non-toxic for mammals (Weinzierl, 1998; Scott et al., 2003). They usually quickly decompose in the environment and are easily digested by the animals, which receive sub-lethal doses (Grdisa & Grsic, 2013).
Reasons for the limited commercial development of botanical insecticides are their relatively slow impact, variable efficiency, absence of stability and non-constant availability (Isman, 2008) compared to synthetic analogues. Other obstacles for commercializing botanical insecticides are deficiency of natural resources, difficulties of standardisation, control of quality and registration (Isman, 1997).
Tribolium confusum Jacquelin du Val, 1863 is one of the commonest insect pests of storages, damage from which is 5-30% of global agricultural production. The reasons for broad distribution of T. confu-sum are the morphological, physiological and behavioural features of the insect, and also the favourable conditions created for it by humans (Hana & Mohammed, 2013).
The objective of this article was to evaluate the impact of different essential oils on the migration activity of T. confusum in the conditions of a laboratory experiment.
Materials and methods
In the experiment, we used imagoes of T. confusum. Before the experiment, the animals were kept in a common container with wheat flour. The beetles were selected randomly. The experiment included three stages. At the first stage, we planned to determine the essential oils which can frighten off or attract imagoes of T. confusum more efficiently than others. In the container (50 x 33 x 19 cm), we put wheat flour of higher sort (400 g) in a layer of 1 cm thickness. Then, in the container, we placed 44 plastic cups with the bottoms removed (0.1 L capacity) at a distance of 0.5 cm one from another with 80 g of flour and 30 imagoes of T. confusum in each one. In 40 cups, in flour on depth of 3 cm, we put a cotton pad of 0.4 cm diameter, saturated with 0.06 mL of essential oil (0.48 mL/cm2). In one experiment, for each of 20 types of essential oil (Table 1), we used two cups. The other four cups were used as control (in them, we put cotton pads of 0.4 cm diameter without processing with any essential oil).
Table 1
Essential oils used in the experiment
for determining migration activity of T. confusum
Plant compounds concentration, % ISO References
stance
Jasmine Jasminum linalool 6.4 - Wei &
oil officinale benzyl acetate 22.9 Shibamoto,
Linnaeus, benzyl alcohol 6.5 2007
1753 cis-jasmone 2.9
p-cresol 1.4
cis-3-hexenyl benzoate 1.1
eugenol 3.0
methyl palmitate 1.2
isophytol 7.5
cis-phytol 15.0
Grape- Citrus a-pinene 0.4 3053 Uysal et al.,
fruit oil paradisi sabinene 0.3 2011
Mac- p-pinene 0.8
fadyen, p-myrcene 0.7
1830 a-terpinene 0.7
limonene 91.5
linalool 1.1
trans-limonene oxide 0.9
citronellal 0.4
a-terpineol 0.3
nerol 0.3
neral 0.4
geraniol 0.3
geranial 0.4
Euca- Eucalyp- a-pinene 5.65 770 Abdossi
lyptus tus globu- p-pinene 0.31 et al., 2015
oil lus Labil- sabinene 0.65
lardiere, limonene 0.84
1861 1,8-cineole 76.65
y-terpinene 0.63
terpinen-4-ol 0.37
a-terpineol 1.96
a-terpineol acetate 4.85
isoledene 0.54
a-gurjunene 0.85
p-gurjunene 0.36
alloaromadendrene 3.98
aromadendrene 0.51
Rose- Rosma- a-pinene 14.90 1342 Gachkar
mary oil rinus camphene 3.33 et al., 2007
officinalis p-octanone 1.61
Linnaeus, myrcene 2.07
1753 1,8-cineole 7.43
linalool 14.90
camphor 4.97
borneol 3.68
terpinen-4-ol 1.70
verbenone 1.94
piperitone 23.70
bornyl acetate 3.08
p-caryophyllene 2.68
Plant compounds concentration, % ISO References
stance
cis-p-farnesene 1.26
a-bisabolol 1.01
Cin- Cinna- heptanal 1.09 - Jayapra-
namon momum nonanal 1.09 kasha et al.,
oil verum a-copaene 23.05 2002
J. Presl, a-bergamotene 27.38
1825 rans-cinnamyl acetate 2.41
aromadendrene 1.79
a-humulene 6.19
germacrene D 2.10
viridiflorene 3.29
a-muurolene 2.70
y-cadinene 1.57
S-cadinene 5.97
ledol 1.29
spathulenol 2.02
globulol 1.67
p-bisabolol 1.26
tetradecanol 4.27
epi-a-bisabolol 2.08
Spruce Picea santene 2.27 - Radulescu
oil abies a-pinene 5.40 et al., 2011
(Lin- camphene 7.55
naeus) H. limonene 9.29
Karsten., borneol 1.11
1881 bornyl acetate 11.78
a-muurolene 1.61
y-cadinene 1.54
S-cadinene 9.49
nerolidol 1.01
a-muurolol 11.01
S-cadinol 1.48
a-cadinol 21.39
manool 3.58
Thuja Thuja a-thujene 1.46 - Jirovetz
oil occi- a-pinene 3.33 et al., 2006
dentalis camphene 2.55
Linnaeus, a-fenchene 2.04
1753 sabinene 12.14
p-pinene 1.14
myrcene 4.05
p-cymene 2.37
a-terpinene 1.83
limonene 2.36
p-phellandrene 1.65
y-terpinene 2.29
rans-sabinene hydrate 1.09
terpinolene 2.32
fenchone 12.87
linalool 1.89
a-thujone 2.76
p-thujone 9.48
camphor 1.24
terpinen-4-ol 3.32
linalyl acetate 1.24
sabinyl acetate 16.55
terpinyl acetate 1.17
p-caryophyllene 1.23
S-cadinene 1.29
Sandal- Ptero- cis-a-santalol 31.67 3518 Subasinghe
wood carpus epi-a-bisabalol 1.44 et al., 2013
oil santalinus epi-ß-santalol 2.36
Linnaeus cis-ß-santalol 14.50
filius, cis-nuciferol 1.02
1782 y-curcumen-12-ol 1.68
ß-curcumen-12-ol 2.35
Ginger Zingiber camphene 3.0 16928 Singh et al.,
oil officinale ß-phellandrene 1.4 2008
Roscoe, 1,8-cineole 1.9
1807 borneol 2.1
neral 7.4
geraniol 3.4
geranial 25.9
ar-curcumene 6.6
a-zingiberene 9.5
Plant compounds concentration, % ISO References
stance
(E,E)-a-farnesene 7.6
p-sesquiphellandrene 5.1
trans-nerolidol 1.5
zingiberenol 1.7
p-eudesmol 1.0
Cedar Cedrus a-terpinene 1.02 9843 Derwich
oil atlantica cis-ocimene 1.62 et al., 2010
(Endli- humulene 1.30
cher) G. p-caryophyllene 3.14
Manetti o-himachalene 7.62
ex Car- cis-a-atlantone 6.78
rière, 1855 himachalol 5.26
a-himachalene 4.15
a-pinene 14.85
p-pinene 1.35
himachalene 10.14
cadinene 3.02
isocaryophillene 1.10
p-himachalene 9.89
germacrene-D 3.52
p-copaene 2.26
cymene 1.05
3-carene 1.10
verbenol 2.24
limonene 2.01
ylangene 2.20
p-phellandrene 2.19
y-amorphane 2.22
Juniper Juniperus a-thujene 1.68 8897 Chatzo-
oil communis a-pinene 41.25 poulou &
Linnaeus, sabinene 17.38 Katsiotis,
1753 p-pinene 2.05 1993
myrcene 2.66
a-terpinene 1.22
limonene 4.23
1,8-cineole 1.21
y-terpinene 2.09
terpinolene 1.16
terpinen-4-ol 2.78
p-caryophyllene 1.69
a-humulene 1.56
germacrene D 1.83
Gera- Pelargo- linalool 5.60 4731 Boukhris
nium oil nium rose oxide-trans 2.01 et al., 2012
graveo- iso-menthone 4.42
lens p-citronellol 21.93
L'Héritier, geraniol 11.07
1789 citronellyl formate 13.24
geranyl formate 6.22
p-bourbonene 3.14
trans-caryophyllene 1.02
germacrene D 4.33
viridiflore 2.35
S-cadinene 2.38
5-cadinene 1.33
a-agarofuran 1.28
10-epi-y-eudesmol 7.92
geranyl tiglate 2.39
Orange Citrus a-pinene 0.36 3140 Singh et al.,
oil sinensis sabinine 0.37 2010
(Lin- p-myrcene 1.71
naeus) octanal 0.43
Osbeck limonene 90.66
(pro. sp.) linalylacetate 2.80
t-sabinine hydrate 0.42
laevo-p-pinene 0.46
geranyl formate 0.65
Tea tree Melaleuca a-pinene 2.1 4730 Cox et al.,
oil alternifo- a-terpinene 8.3 2001
lia (Mai- p-cymene 2.3
den & limonene 1.1
Betche) 1,8-cineole 4.5
Cheel, y-terpinene 17.8
1925 a-terpinolene 3.3
terpinen-4-ol 39.8
a-terpineol 3.4
Sub- Chemical composition
Plant ISO References
stance compounds concentration, %
aromadendrene 1.2
viridiforene 1.2
5-cadinene 1.5
Lime oil Citrus 2,3-dimethyl-2,3-butanediol 1.67 - Sandoval-
aurantii- resorcinol 3.65 Monte-
folia 1-methoxycyclohexene 8.00 mayor
(Christ- linalool oxide 1.18 et al., 2012
mann) corylone 6.93
Swingle, terpinen-4-ol 1.66
1913 a-terpineol 5.97
3-methyl- 1,2-cyclopentanedione 8.27
3,7-dimethyl-(z)-2,6-octadienal 1.09
geraniol 1.15
citral 2.21
7-methyl-(Z)-8-tetadecen-1-ol acetate 2.83
geranyl acetone 1.84
bergamotene 1.00
(z)-8-methyl-9-tetradecenoic acid 1.24
trans-a-bisabolene 1.02
caryophyllene oxide 3.02
spathulenol 1.95
umbelliferone 4.36
palmitic acid 6.89
5,7-dimethoxycoumarin 15.80
5-methoxypsoralen 1.14
5,8-dimethoxypsoralen 6.08
Pepper Mentha 1,8 cineole 6.69 856 Saharkhiz
mint oil piperita menthone 2.45 et al., 2012
Linnaeus, menthofuran 11.18
1753 neomenthol 2.79
menthol 53.28
menthyl acetate 15.10
(z)-caryophyllene 2.06
germacrene D 2.01
Jojoba Sim- saturated acids (C20-C26) 1.64 - Knoepfler
oil mondsia palmitoleic acid 0.24 et al., 1958
chinensis oleic acid 0.66
(Link) eicosenoic acid 30.30
C. K. docosenoic acid 14.20
Schneider, eicosenol 14.60
1912 docosenol 33.70
hexacosenol 2.00
Lemon Citrus p-pinene 5.20 855 Espina
oil limon p-cymene 3.29 et al., 2011
(Lin- limonene 59.10
naeus) y-terpinene 9.66
Osbeck, cis-p-mentha-1 (7), 8-dien-2-ol 1.33
1765 geranial 2.11
cis-thujopsene 2.38
p-bisabolene 3.61
Lavende Lavan- camphene 1.37 3515 Jianu et al.,
r oil dula p-myrcene 2.03 2013
angusti- D-limonene 2.10
folia p-phellandrene 16.00
Miller, 1,8-cineole 15.69
1768 terpinen-4-ol 9.57
borneol 5.07
a-terpineol 6.00
santalene 4.50
caryophyllene 24.12
Almond Prunus myristic acid 0.0-0.07 - Fernandes
oil dulcis palmitic acid 4.7-15.8 et al., 2017
(Miller) palmitoleic acid 0.1-2.5
D. A. stearic acid 0.3-2.5
Webb, oleic acid 50.4-81.2
1967 linoleic acid 6.2-37.1
linolenic acid 0.0-11.1
arachidic acid 0.04-0.20
campesterol 2.5
stigmasterol 2.5
p-sitosterol 55.9-95.1
A5-avenasterol 8.5-28.2
a-tocopherol 97.3
y-tocopherol 2.8
Note. * - number of ISO standard.
Each cup was covered with a separate plastic cover to prevent mixing of odours of essential oils. The experiment was made in five replications (n = 10, i.e. five tests with two replications for each essential oil). The duration of each experiment was 48 hours. At the end of this period, the flour from the cups was sieved for counting live and dead insects.
At the second stage of our research, we evaluated the influence on movement activity of T. confusum by the most efficient essential oils found at the first stage. In the container, we put wheat flour in 1 cm layer and 45 cups with 25 imagoes of T. confusum, of which 40 contained cotton pads processed with essential oil (one variant in four cups), and five cups - control. We used the earlier found repellent (J. officinale, R officinalis, T. occidentalis, M. altemifolia, C. verum, L. angustifolia) and attractant (Z. officinale, C. atlantica, J. communis, C. sinensis) essential oils. The experiment which lasted 24 hours was made in three replications (n = 12, i.e. four cups in three experiments for each ofthe 10 essential oils).
At the third stage, we checked the patterns found earlier in the effect of the essential oils on the migration activity of T. confusum imagoes. In the experiment, we used polyethylene tubes of 4 cm in diameter and 150 cm in length with measurement marks put at each 10 cm of length. On either ends of the tubes, a cotton disk with essential oil was put: with a repellent on one, and with an attractant on the other. Evaporations of the following essential oils were used, selected during the two previous stages: thuja - ginger and jasmine - cedar. Each variant of the experi-
70 65 60
ment lasted 24 h and was performed in eight replications (n = 8). By the end of the experiment, for each 10 cm of tube, the flour was collected and sieved through a laboratory sieve for measuring its presence in each section. All stages of the experiment were conducted in the laboratory with constant illumination and temperature, out of reach of direct sun light. Fluctuations in temperature over 24 hours did not exceed 2 °С (+21.. .+23 °С), duration of daylight in October-November 2018 was 8.30-11.00 hours and was prolonged to 14 h a day by artificial illumination, and air humidity equaled 60-70%.
The results were statistically analyzed in Statistica 8.0 software (Statsoft Inc., USA). Differences between the selections were considered reliable at P < 0.05 (one-way ANOVA).
Results
The property of the studied essential oils to attract or repel T. confusum (Fig. 1) at the first stage of the experiment allowed us to determine that repellent activity against T. confusum imago was exhibited by essential oils of J. officinale, R officinalis, T. occidentalis, M. alternifolia, L. angustifolia and C. verum. Attractant property was also demonstrated by essential oils of J. communis, Z. officinale, C. sinensis and C. atlantica. The rest of the examined essential oils had no significant effect on movement activity of T. confusum.
Fig. 1. Effect of essential oils on migratory ability of T. confusum in 48 h laboratory experiment
At the second stage of the research, we determined that the most active repellents against T. confusum were essential oils of T. occidentalis and J. officinale, and the most active attractants - essential oils of Z. officinale and C. atlantica (Fig. 2, 3, Table 2, 3).
At third stage, we studied distribution of T. confusum under simultaneous effect of repellent and attractant essential oils in evaporations of thuja-ginger and jasmine-cedar. Essential oil of T. occidentalis repelled imagoes at a distance of up to 30 cm. Essential oil from Z. officinale lured insects, but with low efficiency: attractant properties were exhibited at a distance of up to 20 cm (Fig. 4). Essential oil of J. officinale exhibited repellent properties at a distance of up to 20 cm, and C. atlantica lured insects with the same efficiency as oil of Z. officinale (Fig. 5).
If 50
Discussion
The data we obtained indicate that essential oils of J. officinale and T. occidentalis exert notable repellent activity on T. confusum at low concentrations, whereas essential oils of Z. officinale and C. atlantica attract insects.
Fig. 2. Attractant effect of essential oils on T. confusum in conditions of laboratory experiment
60
40
ч 30
20
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0
Absence of notable effects in the rest of the examined samples can be related to insufficient concentration of essential oils or resistance of this pest species. Resistance of the insects to evaporations of essential oils can be related to activity of cytochrome P450-dependent monoox-ygenase, carboxyl esterase, superoxide dismutase and catalase (Ryan & Byrne, 1988; Boyer et al., 2011).
Table 2
Effect of essential oils on distribution of T. confusum in the food substrate
Essential oil
Number of attracted individuals (x ± SD), spec.
P
F
Fo.(
4f 8
—1 r— I—
— 1—'
Citrus sinensis 8.92 ± 8.88
Jasminum officinale 8.67 ± 8.72
Zingiber officinale 13.08 ± 7.83
Cedrus atlantica 12.42 ± 6.39
Cinnamomum verum 6.33 ± 5.40 0.161 1.489 1.966
Lavandula angustifolia 6.83 ± 4.17
Rosmarinus officinalis 7.58 ± 6.46
Thuja occidentalis 4.50 ± 4.03
Melaleuca alternifolia 9.50 ± 12.49
Juniperus communis 9.92 ± 6.91
Table 3
Effect ofessential oils on distribution of dead individuals of T. confusum in food substrate
Essential oil
Number of attracted individuals (x ± SD), spec.
P
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to >S >s c
ji <3> .C
■a o a "co co d
o o c CO o "co o
-J §
Fig. 3. Attractant effect of essential oils on T. confusum in conditions of laboratory experiment
Citrus sinensis Jasminum officinale Zingiber officinale Cedrus atlantica Cinnamomum verum Lavandula angustifolia Rosmarinus officinalis Thuja occidentalis Melaleuca alternifolia Juniperus communis
1.92 ± 1.62 1.33 ± 1.70 1.83 ± 2.00 2.33 ± 1.56 1.92 ± 1.88 3.67 ± 2.81 1.25 ± 1.14 1.58 ± 1.38 1.58 ± 1.31 1.58 ± 1.38
0.056 1.922 1.966
Fig. 4. Effect of essential oils of T. occidentalis and Z. officinale on distribution of T. confusum in fodder substrate
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£ 10
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2 0
Fig. 5. Effect of essential oils from J. officinale and C. atlantica on distribution of T. confusum in fodder substrate
6
4
2
F
F
0
Effect of essential oils and their components on T. confusum and other economically harmful species is described in a number of publications. Haouas et al. (2007) studied biological activity of essential oils from Chrysanthemum spp. against T. confusum. Most efficient was the essential oil obtained from leaves of Ch grandiflorum, which caused decrease in tempi of growth (by 0.03 mg/mg/24 hours), efficiency of consumption of food (by 50.7%), amount of ration consumed (by 66.4%) and increase in mortality of T. confusum larvae up to 80% after 7 days of the experiment. Under exposure to essential oil of leaves of Ch coronarium, mortality reached 18%. Contact toxicity of essential oil of Ch. grandiflorum in 1% concentration equaled 27% after 7 days.
In a study of contact toxicity of essential oil of Eucalyptus saligna for T. confusum, Tapondjou et al. (2001) determined that complete mortality of insects was achieved at a dose of 0.78 and 1.56 mL/cm2 over 4 days. Furthermore, cymol, one of the main components of the studied oil, in 1.30 mL/cm2 concentration caused complete death of T. confusum after 24 hours.
Khani et al. (2017) studied toxicity and repellency of essential oils of Juniperus polycarpus and J. sabina against T. confusum. Mortality significantly increased with increase in concentration and duration of influence. Highest mortality (90%) was achieved at impact of essential oil of J. poly carpus in 611 mcL/L of air and 82% at impact of oil from J. sabina in concentration 666 ^iL/L of air after 24 hours. Values of LC50 for essential oils from J. polycarpus and J. sabina equaled 368.4 and 301.9 ^L/L of air, respectively. Repellency of the tested substances depended on the concentration and equaled 96.0% at concentration of 15 ^L/L of acetone and 6.7% and 10.0% at 1 ^L/L concentration of essential oils from J. polycarpus and J. sabina respectively.
Isikber et al. (2006) investigated toxicity of essential oils of bay laurel (L. nobilis) and rosemary (R. officinalis) for different stages of T. confusum. Highest mortality (95.0%) after 24 hours of exposure was achieved at maximum tested concentration of essential oil from R officinalis (431.5 mg/L of air), whereas essential oil of L. nobilis caused only 15.5% mortality in the same conditions. These parameters are related to insufficient time of exposure. The highest resistance to essential oil of rosemary, was presented by pupae, with LT90 equaling 89.5 hours at 172.6 mg/L of air, whereas imagoes were more sensitive (LT90 = 37.5 hours). Against essential oil of bay laurel, the most resistant were larvae (LT90 = 77.2 hours), whereas pupae were more sensitive (LT90 = 39.3 hours). Though essential oils from L. nobilis and R of ficinalis have fumigant toxicity for all life stages of T. confusum, with different effectiveness, causing complete death of the insects requires using high doses.
Insecticidal action of essential oil from C. sinensis on T. confusum was researched by Oboh et al. (2017). At concentrations of 50 and 75 |iL/L of air, essential oil exhibited moderate insecticidal activity: 50% and 60% mortality respectively after 24 h of impact. At maximum concentration of 150 ^L/L of air, insecticidal activity was high with 100% mortality and LC50 38.9, 26.9 and 14.5 ^iL/L after 24, 48 and 72 hours of impact respectively. Also there was studied inhibiting activity of acetylcholinesterase and Na+/K+-ATPase of T. confusum exposed to essential oil of C. sinensis: IC50 for these enzymes equaled 7.94 and 60.25 ^L/L of air, respectively. Yunis (2014) studied impact of essential oil of C. sinensis on T. confusum: essential oil in 10% concentration caused 70.0%, 86.6% and 100.0% mortality after 1, 2 and 3 days, respectively. In a study of toxicity of essential oil of C. sinensis for T. confusum, Callosobruchus maculatus (Fabricius, 1775) and Rhy-zopertha dominica, Tandorost & Karimpour (2012) determined that LC50 equaled 259, 158 and 124 ^L/L of air after 24 hours of impact and 134, 106 and 93 ^L/L of air after 48 hours for each insect respectively. Highest mortality was achieved using concentrations 53, 41 and 31 ^L-1 against T. confusum, C. maculatus and R. dominica, respectively after 48 hours of exposure.
Campolo et al. (2013) studied biological activity of essential oils from five species of Citrus spp. against T. confusum. Maximum mortality was reached after 24 hours at absence of flour and concentration of 17.2 • 10-3% of essential oils from C. sinensis, C. aurantium and C. limon. Addition of 10 mm of flour reduced efficiency, and 100% mortality was caused by maximum concentration of 69.0 • 10-3% for these
samples. LD50 equaled 4.03, 4.08 and 5.09, and LD90 for C. sinensis, C. aurantium and C. limon equaled 11.14, 11.90 and 15.46 respectively.
Russo et al. (2015) tested insecticidal action of essential oil from Eucalyptus globulus on T. confusum. Insecticidal effect increased depending on time and concentration of the studied substance. High concentration of 1.25 ^L/cm2 eliminated 90% of the pests after 30 minutes of exposure. After two hours of exposure, mortality close to maximum (98.3%) was reached by a lower concentration of 1.00 ^L/cm2, whereas at four-hour exposure, 100% mortality required 0.75 ^L/cm2.
Fathi & Shakarami (2014) undertook a research on larvicidal effect of essential oils of Eucalyptus spp. for T. confusum and T. castaneum. Larvae of T. confusum were more sensitive than T. castaneum. After 24 hours of exposure, LC50 for essential oils of E. camaldulensis, E. vi-minalis, E. microtheca, E. grandis and E. sargentii against larvae of T. confusum were 41.5, 20.7, 53.4, 26.4 and 110.5 ^L/L of air respectively, whereas values of LC50 of these essential oils for larvae of T. castaneum equaled 110.3, 48.1, 117.0, 71.9 and 155.8 ^L/L of air, respectively.
Khalis Ali (2013) explored toxicity of different plant extracts for T. confusum. Indicators of maximum mortality ranged depending on the plant species, therefore for Anethum graveolens, it equaled 56.7% at 4.5 hours exposure, for Apium graveolens - 93.3% after 5 h, Eucalyptus glauca - 90.0%i after 2 h, Malvaparviflora - 96.7% after 3 h, M. longi-folia - 93.3% after 4 h and for Z. officinalis - 100% after 2 h.
Khani et al. (2012) studied toxicity of essential oil from Aloysia ci-trodora Palau, 1784 for T. confusum and Callosobruchus maculatus. It was determined that C. maculates was more sensitive for this fumigant (LC50 = 10.2 ^L/L of air) than T. confusum (LC50 = 497.8 ^L/L of air) at 24 h impact.
Karci & Isikber (2007) researched ovicidal activity of different essential oils in concentration of 100 ^L/L of air against T. confusum over 24, 48 and 72 h. Strong ovicidal effect was exhibited by essential oils from Allium sativum, A. cepa, Pimpinella anisum, Origanum dubium and Foeniculum vulgare, with mortality parameters equaling 99.3%, 100.0%, 95.6%, 100.0% and 96.9% after 72 h of exposure and LT90 equaling 1.1, 22.1, 22.4, 13.8 and 51.1 h, respectively. I^ikber et al. (2009) studied ovicidal action of different essential oils in concentration of 20 ^L/L of air on T. confusum over 24 h. Use of essential oils of
A. sativum, Betula lenta and Cinnamonum zeylanicum caused 100% mortality of the insects, whereas for Pimpinella anisum, this parameter equaled 50.7 ± 1.8%. Values of LC90 for essential oils of A. sativum,
B. lenta, C. zeylanicum and P. anisum were 6.9, 4.5, 3.1 and 33.5 ^iL/L of air respectively.
Stamopoulos et al. (2007) studied biological effect of five monoter-penoids, components of essential oils, against different stages of T. con-fusum. The most toxic for all studied stages were terpinen-4-ol with LC50 in range of 1.1-109.4 ^L/L of air, (R)-(+)-limonene (LC50 - 4278 ^L/L of air) and 1.8-cineole (LC50 - 3.5-466 ^L/L of air). Toxicity of linalool was lower with LC50 in range of 8.6-183.5 ^L/L, and gera-niol was the least toxic: LC50 - 607-1627 ^L/L of air. In all cases, except geraniol, most sensitive stage was third age larvae, and most tolerant - three-days old eggs. Also, a reduction of fertility and production of eggs was observed under the effect of evaporations of tested substances. Tripathi et al. (2001) investigated contact toxicity, fumigant and an-tifeedant activity of 1.8-cineole extracted from Artemisia annua against T. castaneum. The study revealed that adult individuals were more sensitive to 1.8-cineole than larvae. At 121.9 mg/g concentration, antifee-dant effect for imago equaled 81.9%. 1.8-Cineole applied to the filter paper in 3.2-16.1 mg/cm2 concentration significantly reduced number of matured eggs.
Malacrino et al. (2016) studied toxicity of enantiomers of limonene, a component of different essential oils, against T. confusum. Mortality of insects depended on concentration of substance and temperature. In tests without flour, R-(+)-limonene caused 100% mortality at 20 °C and concentration equaling 85 mg/L of air, whereas mortality at impact of S-(-)-limonene was 80.0 ± 2.5% in the same conditions and heightened with increase in temperature up to 88.0 ± 4.9% at 30 °C and to 100% at 40 °C. Mortality caused by R-(+)-limonene at 40 °C reached maximum already at concentration of 42.5 mg/L of air. Addition of
flour significantly reduced the parameters. At presence of 10 mm layer of flour and temperatures of 20 and 30 °C, no mortality of T. confusum was observed, whereas at 40 °C, effectiveness of R-(+)-limonene and S-(-)-limonene equaled 86.0 ± 2.5% and 66.0 ± 6.8%, respectively at maximum concentration (85 mg/L of air).
Kalita & Bhola (2012) investigated toxicity and repellence of different plant extracts for T. castaneum. Highest parameters of mortality of insects were obtained using extracts from Viola arvensis (68%), M. chamomilla (57%), Brassica campestris (56%) and Jacaranda mimosifolia (49%) after 7 days of impact. Furthermore, J. mimosifolia, M. chamomilla and Tagetes minuta exhibited high repellence (IR = 0.04) against T. castaneum. In the research by Al-Jabr (2006) on toxicity and repellence of different essential oils, M. chamomila had high repellence, equaling 81.9 and 84.7 at 1.0% concentration, against O. surinamensis and T. castaneum respectively.
Bhaskar Mi & Tripathi (2011) studied repellence of different essential oils for Sitophilus oryzae (Linnaeus, 1763) and T. castaneum. At highest tested concentration (0.2%) of essential oils from S. aromati-cum, A. marmelos, C. sativum and C. reticulate, their repellence equaled 90.0%, 85.0%, 83.3% and 78.3% for S. oryzae, and 90.0%, 86.6%, 83.3% and 80.0% for T. castaneum respectively.
Maedeh et al. (2012) evaluated toxicity and repellence of essential oil from Z. officinale against T. castaneum. Values of LC50 and LC90 after 48 hours were assessed as 374.9 and 1124.2 ^L/L of air, respectively. Repellence of essential oil was high even at low concentrations and reached 85% at 1.6 ^L/L of air.
Wang et al. (2006) investigated fumigant and repellent action of essential oil from A. vulgaris on T. castaneum. Essential oil in concentration of 0.6 ^g/mL and higher efficiently repelled the insects, and 100% mortality of imagoes was caused at 8.0 ^g/mL. Mortality of larvae of different ages equaled 49-52%. The oil also had high fumigant activity against eggs of T. castaneum and caused 100% mortality at concentrations of10, 15 and 20 g/L of air over 96 h.
Huang & Ho (1998) evaluated toxicity and antifeedant activity of cinnamaldehyde (one of the components of essential oil from C. verum) against T. castaneum. Cinnamaldehyde had no antifeedant effect on imagoes of T. castaneum in concentration up to 13.6 mg/g of fodder, though it significantly reduced consumption of food and growth of larvae in concentrations of 27.2 and 54.4 mg/g of fodder. With increase in concentration of cinnamaldehyde, antifeedant effect increased.
Arabi et al. (2008) studied insecticidal activity of essential oil from Perovskia abrotanoides Karelin (1841) against S. oryzae and T. castaneum. Maximum concentration of 645 mcL/L of air eliminated 80% of T. castaneum after five hours of exposure, whereas for S. oryzae, this parameter equaled around 10% in the same conditions. Maximum mortality was obtained using 322 ^L/L concentration over 7 and 13 h, respectively. Values ofLT50 for S. oryzae ranged from 8 hours for highest concentration (645 ^iL/L of air) to 11.5 hours at lowest concentration (32 ^L/L of air), whereas for T. castaneum, LT50 was achieved in 4.5 hours (32 ^L/L of air) and 2.8 hours (645 ^L/L of air), which confirms the high sensitivity of T. castaneum to the fumigant compared to S. oryzae.
Keita et al. (2001) investigated insecticidal effect of essential oil from T. occidentalis against C. maculatus. Using 100 g of kaolin powder aromatized with 3 ^L of essential oil caused 95% mortality among females and 100%o among males after 6 hours of exposure. LD50 for essential oil from T. occidentalis was 323, 162 and 52 ^L/g at 12, 24 and 48 hours of exposure.
Abou-Taleb et al. (2015) studied toxicity of various essential oils for T. castaneum and determined that the most efficient were essential oils from O. vulgare (LC50 = 9.9 mg/L of air), C. sinensis (LC50 = 24.6 mg/L), C. lemon (LC50 = 25.5 mg/L) and M. communis (LC50 = 26.5 mg/L of air). Parameters of inhibition of acetylcholinesterase (IC50) for C. aurantifolia, C. lemon, O. vulgare and R. officinalis equaled 105.8, 35.3, 24.4 and 20.8 mg/L, and for inhibition of ATPases - 44.4, 10.2, 10.2 and 11.4 mg/L respectively.
Despite the relevance of analysis of methods of using essential oils or their components as insecticidal preparations, their introduction into technology of integrated pest control remains impossible due to the
insufficient amount of information on this problem, and difficulty of standardizing and ensuring quality control of the plant products.
Conclusions
Notable repellent activity against T. confusum has been demonstrated by essential oils from J. officinale and T. occidentalis at concentration of 0.48 mL/cm2. Essential oils from Z. officinale and C. atlantica attracted the insect. We observed manifestations of repellent properties of essential oils from R. officinalis, M. alternifolia, L. angustifolia and C. verum, and that essential oils of J. communis and C. sinensis attracted the pest. Thus, out of 20 studied essential oils, only four samples had notable biological effect on migratory ability of T. confusum. These data support a lot of other research on insecticidal activity of essential oils and possibility of using them as natural pesticides. Study of biological activity of essential oils against economically harmful species of insects is a relevant task necessary for development of ecologically-based pest control.
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