Crop Protection 21 (2002) 419–425

Influence of plant volatiles on feeding damage caused

by the onion thrips Thrips tabaci
Elisabeth H. Koschiera,*, Katrin A. Sedya, Johannes Novakb
a
Institute for Plant Protection, University of Agricultural Sciences (BOKU) Vienna, Peter Jordanstrasse 82, 1190 Vienna, Austria
b
.
Institute for Applied Botany, University of Veterinary Medicine, Veterinarplatz 1, 1210 Vienna, Austria
Received 9 July 2001; received in revised form 28 September 2001; accepted 1 October 2001

Abstract

Leaf disc bioassays were conducted to determine the effects of essential oils and their volatile constituents from plant species
(Lamiaceae family) at three concentrations ranging from 0.01% to 1% on the feeding activity of adult female onion thrips (Thrips
tabaci Lindeman; Thysanoptera: Thripidae). The percentage of feeding damage area on leek (Allium porrum L.) leaf discs and the
adult survival was assessed after 24 h. Onion thrips were significantly deterred by the essential oils of marjoram (Origanum majorana
L.), lavender (Lavandula angustifolia L.) and mint (Mentha arvensis L.) at several concentrations, and by the oil of rosemary
(Rosmarinus officinalis L.) at 1% concentration. Furthermore, thrips feeding damage was reduced as a result of linalool and eugenol
application at three concentrations. Adult survival on the leaf disc surface was significantly decreased by application of terpinen-4-ol
at 1% concentration. Evaluation of the potential of biologically active plant volatiles against T. tabaci may provide a new approach
to the development of antifeedants and/or natural insecticides for use in both biological and integrated pest management strategies.
r 2002 Elsevier Science Ltd. All rights reserved.

Keywords: Thrips tabaci; Antifeedants; Essential oils; Volatile plant compounds

1. Introduction Lamiaceae family and their volatile constituents have

The onion thrips Thrips tabaci Lindeman (Thysanop-
tera: Thripidae) is the major foliage pest in field cultures
of onion (Allium cepa L.) and leek (Allium porrum L.)
in Austria (Kahrer, 1990, 1998). Economic damage by
onion thrips infestation is caused by both nymph and
adult feeding on green leaf parts. They feed by piercing
surface tissues and suck up the exuded cellular contents.
The empty cells show silvery-white spots on the leaf
surface, which are referred to as silver damage. As a
consequence, particularly leek plants become unmarke-
.
table (Cruger, 1991). Besides qualitative losses, quanti-
tative yield reductions in leek (Vierbergen and Ester,
2000) and onions (Kendall and Capinera, 1987) occur as
well.
Volatile chemicals of higher plants are part of the
chemical defense system against herbivores (Rice and
Coats, 1994). Essential oils from plant species within the
*Corresponding author. Tel.: +43-1-476-654-3353; fax: +43-1-476-
654-3359.
E-mail address: koschier@edv1.boku.ac.at (E.H. Koschier).
previously been found to have a broad spectrum of
biological activities such as antifeedant, repellent or
insecticidal properties against a number of agricultural
and stored products pests (e.g., Mansour et al., 1986;
Dube et al., 1989; Shaaya et al., 1991; Hori and
Komatsu, 1997; Lee et al., 1997; Tunc- and Sahinkaya,
1998; Hori, 1999a; Isman, 2000). Volatile monoterpenes
constitute the corpus of many of the essential oils of
Lamiaceae plants (Hallahan, 2000). Several monoter-
penes are known to act as feeding deterrents (Rice and
Coats, 1994), defined as chemicals which inhibit the
feeding activity of insects on a plant (Dethier et al.,
1960).
The effectiveness of plant volatiles as olfactory
attractants has been examined for various thrips species
(e.g., Morgan and Crumb, 1928; Penman et al., 1982;
Kirk, 1985; Pow et al., 1999). Surveys on repulsion
effects of volatile chemicals to thrips have been rarely
reported. Gaum et al. (1994) observed that floral
volatiles of several rose cultivars, even susceptible ones,
repelled western flower thrips (Frankliniella occidentalis
Pergande). Koschier et al. (2000) found salicylaldehyde

0261-2194/02/$ - see front matter r 2002 Elsevier Science Ltd. All rights reserved.
PII: S 0 2 6 1 - 2 1 9 4 ( 0 1 ) 0 0 1 2 4 - 7
420 E.H. Koschier et al. / Crop Protection 21 (2002) 419–425
to elicit negative responses of western flower thrips at
different concentrations. a-Tomatine has been demon-
strated to have a strong antifeedant activity against
Thrips palmi Karny on tomato leaves (Hirano et al.,
1994).
Key objective of the present study was to determine
the effects of essential oils from plant species within the
Lamiaceae family and some of their volatile main
constituents (i) on the feeding activity and (ii) on the
survival of T. tabaci. We selected essential oils and
volatile compounds that have previously been found to
have allomonic and/or toxic activity against various
mite and insect pest species.
2. Materials and methods
2.1. Rearing
A field collected population of T. tabaci was main-
tained in a laboratory rearing using the bean-pod
method, adopted and modified from De Kogel et al.
(1997) and Loomans and Murai (1997). Rearing units
consisted of a glass jar (0.5 l) that was covered by a fine
mesh cloth (105 mm) for ventilation. A piece of filter
paper was placed on the bottom of each glass jar as a
hiding place and pupation site for prepupae and pupae.
The insects were provided with fresh leek leaf pieces
(Allium porrum L.) for feeding and oviposition three
times a week. The jars were placed in a climate chamber
at 25711C and 7075% relative humidity with a
photoperiod of 16:8 (L:D)h.
Fresh leek cultivated in organic farming systems was
purchased weekly for thrips rearing and for bioassays.
2.2. Essential oils and their compounds
The essential oils of marjoram (Origanum majorana
L.), rosemary (Rosmarinus officinalis L.) and sage
(Salvia officinalis L.) were distilled by the Institute for
Applied Botany, University of Veterinary Medicine
Vienna using a 60 l distillation unit (Albrigi, Stallave-
na-Verona, Italy). Lavender (Lavandula angustifolia L.)
and mint oil (Mentha arvensis L.) are commercial
products of the Kurt Kitzing GmbH (Wallerstein,
Germany), and basil (Ocimum basilicum L.) is a product
of Roth (Karlsruhe, Germany). The pure compounds
linalool, 1,8-cineole, terpinen-4-ol, a-pinene and eugenol
used for the bioassays are products of Fluka Chemie
GmbH (Vienna, Austria).
The essential oils (5 ml) were diluted with CH2Cl2
(495 ml) prior to analyses. GC/MS-analyses were per-
formed on a HP 6890 coupled with a HP 5972 MSD and
fitted with a Rtx-5MS 30 m
0.25 mm capillary column
(0.25 mm film thickness). The analytical conditions were:
carrier gas Helium, injector temperature 2501C, split
ratio 50:1, temperature program 60–2401C with 31C/
min. Components were identified by comparing their
Kovats retention indices (Adams, 1995) and mass
spectra (McLafferty, 1989). The results are given in
Table 1.
2.3. Bioassay
Leek leaf discs (1.6 cm diameter) were taken from
inner layers of leek plants using a punch and were placed
in glass Petri dishes (9 cm diameter). All essential oils
and compounds were suspended in distilled water plus
Triton X-100 (0.5%) as wetting agent. Because of the
poor water solubility of 1,8-cineole an adequate amount
of ethanol was added to its suspension. The Petri dishes
with the leaf discs were sprayed by means of Potter
Spray Tower (Burkard Manufacturing Co Ltd., Rick-
mansworth, England) at constant air pressure with
dilution quantities at 1%, 0.1% or 0.01% concentration
to achieve an exact wet deposit of 0.1, 0.01 or 0.001 ml oil
or compound per cm2 leaf surface. Control leafs discs
were treated with Triton X-100 (0.5%) in distilled water
or in the case of 1,8-cineole with Triton X-100 (0.5%)
plus ethanol in distilled water only. After drying, treated
leafs were placed singly in bioassay units which
consisted of glass Petri dish bottoms (6 cm diameter).
The underside of the leaf discs were embedded into an
agar film (Agar, Fluka Chemie GmbH, Vienna, Austria)
on the base of each bioassay unit to keep the leaf discs
moist and to prevent thrips crawling underneath them.
Ten adult female thrips of unknown ages were collected
randomly from the mass rearing. They were shortly
anaesthetized with CO2 and introduced to each bioassay
unit using a fine brush. Each unit was covered with a
thin plastic film (Toppitss, Melitta Bentz KG, Minden,
Germany) which was perforated by means of insect pins
(0.4 mm diameter) for ventilation (three holes per cm2
on average). The bioassay units were stored in a climatic
chamber at 25711C and 7075% r.h. with a photo-
period of L16:D8. Data on temperature and relative
humidity were recorded continuously (Testostor 171
Logger, Testo GmbH, Lenzkirch, Germany). All units
were randomized to exclude any position effects in the
climatic chamber.
Each experiment was replicated six times for each
volatile at each concentration. Differences in color and
age of the plant material caused differences in the level
of damaged area between the experimental series. To
equalize these differences we included an untreated
control plot in each experimental series using material
from a single plant. After 24 h thrips were removed from
the leaf discs. The number of dead, i.e. not moving when
picked up with a fine brush, and live individuals were
determined indicating adult survival. The percentage of
silver damage area on the leaf discs was measured with
an image analysis system that consisted of a binocular
 E.H. Koschier et al. / Crop Protection 21 (2002) 419–425 421

Table 1
Relative percentage (>1%) of volatile compounds in the respective essential oil detected on GC-MS

Compounds Essential oil

Marjoram Rosemary Basil Sage Mint Lavender

a-Thujene 1.3
a-Pinene 10.0 3.4
Camphene 2.6 4.9
Sabinene 5.2
b-Pinene 1.8
Myrcene 1.6
Para-cymene 11.3 6.3
Limonene 1.6 6.5 2.1 3.3
1,-8 Cineole 50.8 3.9 15.5
g-Terpinen 1.6
Trans-sabinene hydrate 5.8
Cis-sabinene hydrate 19.1
Linalool 3.1 9.6 42.3
a-Thujone 12.0
b-Thujone 4.1
Camphor 6.3 26.3
Isopulegol 1.1
Menthone 24.0
Isomenthone 10.5
Neo-menthol 6.9
Borneol 5.1 5.6 1.5
Menthol 33.5
Terpinen-4-ol 22.5 1.2 1.7
Isomenthol 1.5
a-Terpineol 4.8 5.0 1.2
Methyl chavicol 78.5
Pulegone 1.4
Cis-sabinene hydrate acetate 10.2
Linalyl acetate 7.4 40.2
Bornyl acetate 1.2 2.7
Lavandulyl acetate 3.4
Menthyl acetate 5.0
a-Terpinylacetate 1.3
b-Caryophyllene 2.0 4.1 1.5 2.4
a-Humulene 7.8
Viridiflorol 4.9
Caryophyllene oxide 1.7

microscope, a digital camera (Nikon D1 Professional 3. Results

Digital SLR Camera, Nikon Cooperation, Tokyo,
Japan) and an image analyzing software (Lucia Gs,
LIM Laboratory Imaging s.r.o., Prague, Czech
Republic).
2.4. Statistical analysis
Data were analyzed using STATGRAPHICS Plus for
Windows 4.0. Homogeneity of variances was tested
using Bartlett’s test. In case of inhomogeneity
pffiffiffiffiffiffiffiffiffiffiffi
variances data were transformed to x0 ¼ arcsin ðx=nÞ:
A one-way-ANOVA was performed to determine
differences between percentage of damaged area on the
leaf discs or adult survival of each treatment, respec-
tively. Means were compared using Tukey’s HSD test
(a ¼ 0:05).
Application of the essential oils of marjoram,
rosemary, mint and lavender showed significant dose-
dependent effects on the feeding activity of adult T.
tabaci females (Table 2). A clear antifeedant activity
against the onion thrips was found by application of the
essential oil of marjoram. Depending on the oil
concentration from 0.01% to 1% the silver damage on
the treated leaf discs was reduced between 91.8% and
of
93.9% compared to the untreated control leaf discs.
Lavender oil did not deter onion thrips females at
the lowest 0.01% concentration, but significantly
reduced feeding damage at 0.1% and 1% solution. No
significant reduction of the silver damage caused by
T. tabaci feeding occurred after application of rosemary
oil at 0.1% and 0.01% concentration. The highest
422 E.H. Koschier et al. / Crop Protection 21 (2002) 419–425

Table 2
Effects of essential oils on feeding activity (percentage of leaf area damaged) of T. tabaci adult females after 24 h

Concentration Essential oil

Marjoram Rosemary Basil Sage Mint Lavender

% damaged area (7s.e.)

Control 2.79 (70.6) aa 3.80 (70.7) aa 2.32 (70.7) aa 6.05 (71.6) 11.27 (71.5) aa 10.86 (70.8) aa
0.01% 0.23 (70.1) b 4.70 (70.4) a 2.89 (70.7) a 6.69 (71.3) 9.51 (71.3) ab 11.93 (71.3) a
0.1% 0.17 (70.1) b 3.71 (70.4) a 0.82 (70.3) a 5.68 (71.4) 7.27 (70.7) ab 6.83 (70.8) b
1.0% 0.21 (70.1) b 1.31 (70.2) b 0.88 (70.4) a 5.54 (70.9) 6.44 (70.7) b 6.63 (70.7) b
F-ratio 20.2 *** b 9.2 *** 3.3 * 0.2 n.s. 4.1 * 8.8***
Df 3,20 3,20 3,20 3,20 3,20 3,20
a
Mean percents in each column followed by the same letter are not significantly different (Tukey’s HSD test, pp0:05).
pffiffiffiffiffiffiffiffiffiffiffi
b
F probability was based on the ANOVA of x0 ¼ arcsin ðx=nÞ transformed data.

Table 3
Effects of essential oil components on feeding activity (percentage of leaf area damaged) of T. tabaci adult females after 24 h

Concentration Component

a-Pinene 1,8-Cineole Terpinen-4-ol Linalool Eugenol

% damaged area (7s.e.)

Control 0.37 (70.2) 8.30 (71.2) 0.08 (70.02) 0.31 (70.05) aa 1.36 (70.5) aa
0.01% 0.26 (70.1) 8.82 (71.5) 0.02 (70.01) 0.05 (70.01) b 0.35 (70.1) b
0.1% 0.30 (70.1) 7.69 (70.5) 0.04 (70.01) 0.01 (70.00) b 0.11 (70.1) b
1.0% 0.23 (70.1) 7.34 (71.0) 0.03 (70.02) 0.03 (70.01) b 0.17 (70.1) b
F-ratio 0.2 n.s. 0.3 n.s. 2.0 n.s. 48.4 *** b 7.4 ** b
Df 3,20 3,20 3,20 3,20 3,20
a
Mean percents in each column followed by the same letter are not significantly different (Tukey’s HSD test, pp0:05).
pffiffiffiffiffiffiffiffiffiffiffi
b
F probability was based on the ANOVA of x0 ¼ arcsin ðx=nÞ transformed data.

Table 4
Effect of essential oils on survival of T. tabaci adult females after 24 h

Concentration Essential oil

Marjoram Rosemary Basil Sage Mint Lavender

a
Number of alive adult females (7s.e.)
Control 9.83 (70.2) ab 9.83 (70.2) 9.33 (70.3) 10.00 (70.0) 9.83 (70.2) 9.67 (70.3)
0.01% 9.83 (70.2) a 9.83 (70.2) 9.67 (70.2) 10.00 (70.0) 9.83 (70.2) 9.50 (70.2)
0.1% 9.83 (70.2) a 9.67 (70.2) 8.83 (70.5) 10.00 (70.0) 9.83 (70.2) 9.00 (70.4)
1.0% 9.00 (70.4) a 9.67 (70.2) 9.33 (70.3) 9.67 (70.2) 9.50 (70.3) 9.17 (70.4)
F-ratio 3.2* 0.3 n.s. 1.0 n.s. 2.5 n.s. 0.6 n.s. 0.8 n.s.
Df 3,20 3,20 3,20 3,20 3,20 3,20
a
Results indicate number alive adult females out of 10.
b
Mean numbers of females followed by the same letter are not significantly different (Tukey’s HSD test, pp0:05).

concentration tested (1%) of this essential oil signifi-
cantly reduced the percentage of damaged leaf area by
feeding of thrips females. A similar result was found
with mint oil, even though inhibition of thrips feeding
activity was not as clearly pronounced as using
rosemary. A leaf surface treatment with basil oil or
sage oil did not show clear deterrent properties.
Two of five tested volatile compounds significantly
deterred feeding by T. tabaci (Table 3). Thrips feeding
damage was reduced as a result of linalool and eugenol
application at the three tested concentrations ranging
from 0.01% to 1%. a-Pinene, 1,8-cineole and terpinen-
4-ol did not interfere with the feeding activity of the
onion thrips.
Adult survival of T. tabaci females on the leaf surface
was not significantly affected by essential oil application
after 24 h (Table 4). A similar result was found with four
of the five tested plant volatiles (Table 5). Application
of a-pinene, 1,8-cineole, linalool and eugenol had no
lethal effect on thrips females. In contrast, terpinen-4-ol
 E.H. Koschier et al. / Crop Protection 21 (2002) 419–425
Table 5
Effect of essential oil components on survival of T. tabaci adult females after 24 h
Concentration Component
a-Pinene 1,8-Cineole
a
Number of alive adult females (7s.e.)
Control 9.83 (70.2) 10.00 (70.0)
0.01% 9.83 (70.2) 10.00 (70.0)
0.1% 9.67 (70.2) 9.67 (70.2)
1.0% 9.67 (70.2) 9.83 (70.2)
F-ratio 0.3 n.s. 1.4 n.s.
Df 3,20 3,20
a
Results indicate number alive adult females out of 10.
b
Mean numbers of females followed by the same letter are not significantly different (Tukey’s HSD test, pp0:05).
caused 14.3%, 8.9% and 16.1% mortality at 0.01%,
0.1%, and 1% concentration, respectively.
4. Discussion
Bioassays used in antifeedant research should match
field conditions as far as possible (Koul, 1993). There-
fore, for the present study we chose a leaf-disc bioassay
that permits volatilization of the essential oils and the
compounds from the plant surface where T. tabaci
individuals were feeding. For crop pests such as T.
tabaci a no-choice test is appropriate, because this
situation is representative of monocultures in agricultur-
al systems (Lewis and Van Emden, 1986; Koul, 1993).
In general, our results provide evidence for a dose-
dependent antifeedant activity of the tested essential oils
and compounds against T. tabaci. Previous studies on
western flower thrips (F. occidentalis) indicated that the
plant odor concentration is crucial for its attractiveness
or repellency (Pow et al., 1999; Koschier et al., 2000).
Our results demonstrate that the essential oil from
Origanum majorana (L.) significantly reduced the area of
feeding damage caused by T. tabaci females on treated
leek leaf discs. Additionally, we observed a somewhat
lower adult survival at 1% oil concentration. The
marjoram oil used in this study contained 22.5% of
the monoterpene terpinen-4-ol. Bioassays with terpinen-
4-ol showed a slight feeding inhibition and a decreased
survival at the three tested concentrations, even though
the responses were very low. Lee et al. (1997) and Isman
(2000) found repellence and toxicity of terpinen-4-ol to
the two-spotted spider mite (Tetranychus urticae Koch)
and several insect species.
No other plant chemical tested in the present study
caused reduced survival of onion thrips. Allelochemicals
such as monoterpenes are part of the defense system of
plants against herbivores. Their antifeedant properties,
which inhibit the ingestion of an insect, appear more
important than any acute toxic effect (Rice and Coats,
1994).
423
Terpinen-4-ol Linalool Eugenol
9.33 (70.2) ab 9.67 (70.2) 9.67 (70.2)
8.00 (70.4) ab 9.67 (70.2) 9.83 (70.2)
8.50 (70.3) ab 9.50 (70.3) 9.17 (70.2)
7.83 (70.4) b 9.67 (70.2) 9.33 (70.3)
3.5 * 0.1 n.s. 1.8 n.s.
3,20 3,20 3,20
The essential oil of Lavandula angustifolia (L.)
deterred thrips females from feeding at 0.1% and 1%
concentration. Hori (1998, 1999a) found lavender oil to
act as repellent and antifeedant against the green peach
aphid, Myzus persicae (Sulzer). The analysis of the
lavender oil used in the present study revealed linalool
(42.3%) as major component. Our results with linalool
further indicate that the feeding deterrence of lavender
oil to T. tabaci is probably due to the monoterpene
linalool, which is a common compound found in floral
fragrances (Knudsen et al., 1993). The feeding damage
was clearly reduced with linalool even at a low (0.01%)
concentration. In contrast, linalool has been demon-
strated to elicit positive responses of F. occidentalis
(Koschier et al., 2000). Linalool (9.6%) was also
identified as major component in the essential oil of
Ocimum basilicum L. and is probably responsible for the
slightly decreased onion thrips feeding activity on the
0.1% and 1% basil oil treatment, though the response
was very low. In contrast to our basil oil chemotype,
basil often contains the phenylpropanoid eugenol as
main compound (Fugmann and Adam, 1997). Eugenol
clearly inhibited the feeding activity of T. tabaci in our
leaf disc assays. In conclusion, we expect a eugenol-
containing basil chemotype to feature a more distinct
antifeedant activity against the onion thrips than the
chemotype used in our experiments. Isman (2000)
observed an antifeedant effect of eugenol to the green
peach aphid and other insect species. Contrarily,
eugenol was previously found to attract cereal thrips
species (Holtmann, 1963) and western flower thrips
(Koschier et al., 2000) at 1% concentration.
The essential oil of Rosmarinus officinalis L. has been
demonstrated to provoke negative responses of two
aphid pest species (Hori and Komatsu, 1997; Hori,
1998, 1999a, b). We found rosemary oil to deter thrips
females at the 1% concentration. Since 1,8-cineole is the
main component in rosemary oil, we expected it to be
the main factor of deterrence in the oil, but neither 1,8-
cineole nor a-pinene, another main constituent inhibited
feeding activity of onion thrips. Minor compounds or
424 E.H. Koschier et al. / Crop Protection 21 (2002) 419–425
non-volatile components are probably involved in the
antifeedant properties of rosemary oil. Mentha arvensis
(L.) essential oil does not contain any of the tested
volatile compounds, though application of mint oil
resulted in reduced feeding damage. Testing of the main
constituents such as the p-menthanones or menthol,
might reveal the component responsible for its anti-
feedant effect.
In conclusion, the results of the present study provide
evidence that essential oils from plants within the
Lamiaceae family and their volatile constituents inter-
fere with the feeding activity of T. tabaci females. If
effective in field cultures of leek, application of plant
volatiles might prevent qualitative crop losses due to
severe feeding damage on green leaf parts. The prospect
of crop protection against thrips pests by means of
behavior-controlling natural products such as plant
volatiles is promising since they are generally regarded
as not toxic to non-target organisms and do not persist
in the environment (Isman, 1999).
Acknowledgements
The authors thank the colleagues at ZID BOKU for
the technical support and Dr. A. Ploner and Assistant
Prof. Dr. K. Moder for their valuable comments on the
statistical analysis. This project is supported by the
Austrian Science Fund (FWF, project P 14172-BIO).
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