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Environmental Science and Pollution
Research
ISSN 0944-1344
Environ Sci Pollut Res
DOI 10.1007/s11356-017-9795-6
Oviposition deterrent activity of basil
plants and their essentials oils against Tuta
absoluta (Lepidoptera: Gelechiidae)
Boni Barthélémy Yarou, Thomas
Bawin, Antoine Boullis, Stéphanie
Heukin, Georges Lognay, François Jean
Verheggen & Frédéric Francis
1 23
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CHEMISTRY, ACTIVITY AND IMPACT OF PLANT BIOCONTROL PRODUCTS
Oviposition deterrent activity of basil plants and their essentials
oils against Tuta absoluta (Lepidoptera: Gelechiidae)
Boni Barthélémy Yarou
1
&Thomas Bawin
1
&Antoine Boullis
1
&Stéphanie Heukin
2
&
Georges Lognay
2
&François Jean Verheggen
1
&Frédéric Francis
1
Received: 23 January 2017 /Accepted: 19 July 2017
#Springer-Verlag GmbH Germany 2017
Abstract The leafminer Tuta absoluta Meyrick
(Lepidoptera: Gelechiidae) is one of the most important pests
of tomato, reducing crop yields by up to 100% in greenhouses
and fields, in several countries globally. Because synthetic
insecticides lead to resistance and have adverse effects on
natural enemies and the health of producers, alternative con-
trol methods are needed. In this study, we assessed the
oviposition-deterring effect of basil plants, Ocimum
gratissimum L. and O. basilicum L. (Lamiaceae), using
dual-choice behavioural assays performed in flight tunnels.
We found that both plants significantly reduced T. absoluta
oviposition behaviour on a tomato plant located nearby. To
evaluate the potential effect of basil volatile organic com-
pounds, we formulated essential oils of both plant species in
paraffin oil, and observed a similar oviposition-deterring ef-
fect. Gas chromatography analyses detected 18 constituents in
these essential oils which the major constituents included thy-
mol (33.3%), p-cymene (20.4%), γ-terpinene (16.9%),
myrcene (3.9%) in O. gratissimum and estragol (73.8%), lin-
alool (8.6%), β-elemene (2.9%) and E-β-ocimene (2.6%) in
O. basilicum. Twenty and 33 compounds were identified of
the volatiles collected on O. gratissimum and O. basilicum
plants, respectively. The main components include the
following: p-cymene (33.5%), γ-terpinene (23.6%), α-
terpinene (7.2%), α-thujene (6.7%) and E-α-bergamotene
(38.9%) in O. gratissimum, and methyl eugenol (26.1%),
E-β-ocimene (17.7%), and linalool (9.4%) in O. basilicum.
Four compounds (α-pinene, β-pinene, Myrcene, Limonene)
were common in essential oils and plants. Our results suggest
the valuablepotential of basil and associated essential oils as a
component of integrated management strategies against the
tomato leafminer.
Keywords O. basilicum .O. gratissimum .Essential oil .
Behaviour .Oviposition .Tuta absoluta .Integrated
management
Introduction
Tomato , Solanum lycopersicum L. (Solanaceae), production is
damaged by the leafminer, Tuta absoluta Meyrick
(Lepidoptera: Gelechiidae), which is considered as one of
the most important pests in both greenhouse and outdoor pro-
duction (Desneux et al. 2010; Desneux et al. 2011; Campos
et al. 2017). Native to South America, this Microlepidoptera
has recently become an invasive pest on tomato crops in
Europe (Desneux et al. 2011) and Africa (Brévault et al.
2014; Tonnang et al. 2015). Tuta absoluta larvae destroy all
aerial parts of plants (leaves, stems, fruits, buds and flowers),
resulting in severe yield losses (i.e. up to 100%) (Desneux
et al. 2010; Urbaneja et al. 2013). Although tomato is the
preferred host plant of T. absoluta, it has also been recorded,
and/or develops, on other cultivated and non-cultivated plants.
Its host-range includes other Solanaceae species, such as po-
tato (Solanum tuberosum L.), aubergine (Solanum melongena
L.), black nightshade (Solanum nigrum L.) and bittersweet
nightshade (Solanum dulcamara L.). It also targets species
Responsible editor: Philippe Garrigues
*Boni Barthélémy Yarou
entomologie.gembloux@ulg.ac.be
1
Functional and Evolutionary Entomology, Agro Biochem
Department, Gembloux Agro-bio Tech, University of Liege (ULg),
Passage des Déportés, 2, 5030 Gembloux, Belgium
2
Analytical Chemistry, Agro Biochem Department, Gembloux
Agro-bio Tech, University of Liege (ULg), Passage des Déportés, 2,
5030 Gembloux, Belgium
Environ Sci Pollut Res
DOI 10.1007/s11356-017-9795-6
Author's personal copy
from other plant families, such the Fabaceae, including the
groundnut (Arachis hypogaea L.), cowpea (Vigna unguiculata
L.) and bean (Phaseoulus vulgaris L.) (Desneux et al. 2010;
Bawin et al. 2015a,b).
The control of T. absoluta is mainly based on synthetic
insecticides (da Silva Galdino et al. 2011; Valchev et al.
2013); however, these methods have limits. For instance,
many active compounds, including abamectin, spinosad,
indoxacarb (Siqueira et al. 2001;Camposetal.2015); delta-
methrin, methamidophos (Lietti et al. 2005); and
cypermethrin, chlorpyriphos (Roditakis et al. 2013), induce
the emergence of resistant pest populations. Moreover, these
insecticides also negatively impact the natural enemy popula-
tions of T. absoluta (Arnó and Gabarra 2011; Biondi et al.
2013; Abbes et al. 2015) owing to multiple potential side
effects as described by Desneux et al. (2007) in a review.
The effectiveness of biological control using entomopatho-
genic organisms (González-Cabrera et al. 2011;Ben
Khedher et al. 2015) beneficial insects (Ferracini et al. 2012;
Chailleux et al. 2012; Chailleux et al. 2013;Öztemiz2013;
Salehi et al. 2016) present valuable alternative methods to
synthetic pesticides. For example, the use of certain Miridae
as Macrolophus pygmaeus Rambur or Dicyphus maroccanus
Wagner appears to be a reliable biological alternative for the
control of T. absoluta (Urbaneja et al. 2013; Abbas et al. 2014;
Jaworski et al. 2015). Until now, prophylactic methods
(Urbaneja et al. 2013) supplemented with pheromone traps
(Filho et al. 2000; Vacas et al. 2011; Cocco et al. 2013)have
facilitated the development of a reliable and sustainable form
of management of T. absoluta. In addition to the methods
mentioned above, plant breeding could be a promising ap-
proach for T. absoluta management also. Indeed, some tomato
cultivars appear to be less susceptible to T. absoluta damage
according to Sohrabi et al. (2016).
The essential oils of many plants species have been dem-
onstrated to have beneficial effects that manipulate insect
pests and reduce damage to crops (Fayalo et al. 2014; Liu
et al. 2014). For instance, the biocidal effects (insecticidal,
repellent) of Ocimum spp. (Lamiaceae) have been well stud-
ied, mainly on malaria insect vectors (Kazembe and Chauruka
2012; Belong et al. 2013; Akono Ntonga et al. 2014)andon
pests of stored products (Ilboudo et al. 2010; Adeniyi et al.
2010; Koubala et al. 2013). In intercropping systems, Ocimum
spp. are also effective in controlling additional crop pests,
including moths, leaf beetles, aphids and whiteflies (Beizhou
et al. 2011;Songetal.2013). However, knowledge remains
limited on how aromatic plants (especially Ocimum species)
affect the tomato leafminer T. absoluta.
In this study, we evaluated the repellent and oviposition-
deterring effect of two Ocimum species, Ocimum gratissimum
L. and Ocimum basilicum L.,on the tomato leafminer
T. absoluta. Both aromatic species are used as food ingredients
and medecinal plants (Selvakkumar et al. 2007;Prabhuetal.
2009;Bilaletal.2012), so they could be considered harmless to
humans. Our results are expected to indicate the potential of
Ocimum species as alternative pest biocontrol methods.
Materials and methods
Plant and insect rearing
Ocimum gratissimum (African basil) and O. basilicum
(European basil) seeds were provided by the Vegetable
Crops Program of the National Institute of Agricultural
Research of Benin (INRAB), West Africa. Tomato
S. lycopersicum cv. Moneymaker and basil plants were indi-
vidually grown under greenhouse (Gembloux, Belgium,
25 ± 5 °C, 50–70% relative humidity [RH], 16:8-h light: dark
photoperiod) in plastic pots (8 × 8 × 9 cm) filled with potting
soil (VP113BIO, Peltracom, Belgium) and were watered ev-
ery 2 days. Plants were used in experiments when they
reached 4 weeks (S. lycopersicum and O. basilicum)and
6weeks(O. gratissimum) after seeding (i.e. at about 20–
25 cm in height).
The larvae of T. absoluta (third instar) were first collected
in July 2011 from a commercial tomato plantation (SAS
Rougeline, Saint-Andiol, France, 43° 49′53.1″N4°58′
20.1″E). The T. absoluta colony was subsequently main-
tained on tomato plants in 45 × 45 × 45 cm net cages
(BugDorm, MegaView Science, Taichung, Taiwan) in a
L2Q laboratory (24 ± 1 °C, 60–70% RH, and 16:8-h light:
dark photoperiod).
Oviposition assays
Effect of Ocimum plants on Tuta absoluta oviposition
response
Flight tunnel experiments were used to evaluate how basil
plants in the vicinity of a tomato plant impacted the oviposi-
tion of T. absoluta adults. The flight tunnels
(230 × 45 × 45 cm) were divided into three areas (Caparros
Megido et al. 2014; De Backer et al. 2015,2016): a central
area for insect release and two areas at opposite sides contain-
ing the plants. Two modalities were tested: (1) a tomato plant
associated with either an O. gratissimum or O. basilicum plant
versus (2) a tomato plant without Ocimum plant (control). For
each replicate, 15 unsexed T. absoluta adult individuals were
randomly sampled from the rearing population and were re-
leased in the central area of the tunnel. After 48 h, the number
of males and females in each area of the tunnel, as well as the
number of eggs laid on each plant, were recorded. The exper-
iments were conducted under the following conditions:
20 ± 1 °C, 65 ± 5% RH, and a 16:8-h light: dark photoperiod
Environ Sci Pollut Res
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under cool white LED lights (77 μmoL/sqm/s). These condi-
tions were monitored using an automatic data logger (HOBO
RH/TEMP 8 K; Onset Computer Corporation, Bourne, MA,
USA). Six replicates were assessed for each modality (i.e.
basil species association).
Effect of Ocimum essential oils on Tuta absoluta
oviposition response
Essential oils (EO) from O. gratissimum and O. basilicum
were purchased from local manufacturers in Benin. Two con-
centrations of each EO (i.e. 5 and 10 mg/ml) were formulated
in paraffin oil (PO), and were tested for their oviposition de-
terring following the same experimental design as described in
the previous section. A 1 ml cylindrical rubber septum
(17 mm high × 10 mm diameter) (VWR International,
Radner, PA, USA) loaded with 100 μl solution (formulated
EO or PO alone) was placed on each plant as a semiochemical
dispenser. The tested modalities were: (1) a tomato plant with
100 μl of a 5 mg/ml corresponding to 0.5 mg of EO
(O. gratissimum or O. basilicum) versus a tomato plant with
100 μl of PO and (2) a tomato plant with 100 μl of a 10 mg/ml
corresponding to 1.0 mg of EO (O. gratissimum or
O. basilicum) versus a tomato plant with 100 μl of PO. Six
replicates were assessed for each treatment.
Analysis of essential oils and characterisation
of plant volatiles components
Essential oil analysis of basil
Essential oil components were analysed by both gas
chromatography-flame ionisation detection (GC-FID) and
gas chromatography-mass spectrometry (GC-MS). GC-FID
analyses were performed using an Agilent Technologies
(Santa Clara, CA, USA) 6890 gas chromatograph fitted with
a flame ionisation detector and capillary column HP-5 (5%
Phenyl Methyl, 30 m, 0.25 mm i.d., 0.25 μmfilmthickness).
Helium was used as carrier gas at a constant flow rate of
1.5 ml/min. The temperature program was started at 40 °C
for 2 min, and then increased by 8 °C/min to 280 °C, with
final hold at this temperature for 5 min. Injector (splitless
mode) and detector (H
2
, 35 ml/min; air, 350 ml/min) temper-
atures were 280 and 290 °C, respectively.
GC-MS was carried out on an Agilent Technologies (Santa
Clara, CA, USA) 6890 gas chromatograph coupled to an
Agilent 5973 mass spectrometer. The analysis was performed
under similar conditions (previous section) using a capillary
column (HP-5 MS: 30 m, 0.25 mm i.d., 0.25 μmfilmthick-
ness), with helium as the carrier gas. The mass spectra scan-
ning range was set from 35 to 350 amu (EI mode at 70 eV).
The components were identified by comparing the recorded
mass spectra with the Wiley 275.L spectral database using
Chemstation software (Agilent Technologies, Palo Alto, CA,
USA). Further identification was carried out by injecting a
homologue series of n-alkanes (C
7–
C
30
) under identical chro-
matographic conditions, and calculating non-isothermal reten-
tion indices (RI). These RI were compared to those reported in
the published literature (Adams 2007; Babushok et al. 2011;
Pherobase 2016). Relative proportions of the components
were calculated based on GC-FID peak areas using response
factors of 1 for each component.
Plant volatiles analysis
To identify the volatile organic compounds (VOCs) emitted
by each basil species, plants at 4 to 6 weeks after seed emer-
gence were placed in separate 4 L glass jars to collect VOCs
with solid-phase micro-extraction (SPME, 10-mm fibre with a
50/30 μm carboxen-ivinylbenzene-polydimethylsiloxane
coating-Supelco). Before sampling, the fibres were condi-
tioned at 250 °C for 1 h in a GC injector. Volatiles were
collected at 25 ± 1 °C for 1 h. GC-MS analyses were per-
formed in triplicate (three SPME sampling on three different
plants of each species) under the aforementioned conditions
(EO analysis section). The components were identified by
comparing the recorded mass spectra with the Wiley 275.L
spectral database using Chemstation software (Agilent
Technologies, Palo Alto, CA, USA). The relative proportion
of each component was calculated by dividing the peak area of
each compound by the total peak area.
Statistical analyses
Binomial proportion tests (equal distribution hypothesised)
were used to compare the number of (1) females and males
recorded in each area of the tunnel and (2) the number of eggs
laid on basil-associated and non-associated tomato plants dur-
ing the dual-choice behavioural assays. These statistical tests
were performed using Minitab® v.17 software (Minitab,
Coventry, UK). The significant threshold was P<0.05.
Results
Effect of Ocimum plants on the distribution
and oviposition response of Tuta absoluta
Both males and females were equally distributed (P>0.05)on
both sides of the flight tunnels in all of the tested modalities
(Table 1). However, females laid significantly more eggs on
non-associated tomato plants compared to tomato plants asso-
ciated with O. gratissimum (P<0.001)andO. basilicum
(P<0.001)(Fig.1).
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Effect of Ocimum essential oils on the distribution
and oviposition response of Tuta absoluta
The number of insects that responded to the EOs and the
distribution of individuals are presented in Table 1. Males
were equally distributed on both sides of the flight tunnels in
almost all tested modalities. In contrast, T. absoluta female
attraction differed between tomato plants associated with PO
and those associated with the EOs of either O. gratissimum
(P= 0.04 with 1.0 mg) or O. basilicum (P= 0.03 with 0.5 mg).
Females also laid significantly more eggs on tomato plants
associated with PO compared to tomato plantsassociated with
an O. gratissimum or O. basilicum EO dispenser, except when
the dose of O. gratissimum was 0.5 mg (Fig. 2).
Chemical components of essential oils and plant volatiles
For each essential oil, 18 compounds were identified
(Table 2). Mass spectra and RI reflected the data from the
published literature (Runyoro et al. 2010; Babushok et al.
2011; Kpadonou Kpoviessi et al. 2012). Six of these com-
pounds were common to both Ocimum species (α-pinene,
Camphene β-pinene, Myrcene, Limonene and Borneol).
Monoterpenoids were the main components, with thymol
(33.3%), p-cymene (20.4%), γ-terpinene (16.9%) and
myrcene (3.9%) in the EO of O. gratissimum EO and estragol
(73.8%), linalool (8.7%), β-elemene (2.9%) and E-β-ocimene
(2.6%) in the EO of O. basilicum. Twenty and 33 compounds
were identified from plant volatile analyses for
O. gratissimum and O. basilicum, respectively (Table 3).
Thirteen compounds are common between the volatile blends
of the two Ocimum species (α-thujene, α-pinene, β-pinene,
Sabinene, Myrcene, α-phellandrene, δ-3-carene, α-terpinene,
p-cymene, Limonene, Cis-ocimene γ-terpinene and Allo-
ocimene). Monoterpenoids were, again, the main components
Tabl e 1 Results of dual choice evaluating the distribution of T. absoluta associated with Ocimum plants (Og: O. gratissimum;Ob:O. basilicum)and
essential oil (EO) and non-associated tomato plants (T). Values are the total numbers of individuals for each combination (six replicates)
Insects tested number Responding insects (%)
a
Choice behaviour with plants pvalue
96 83 (86.5) T T + Og
Males 18 14 0.59
Females 28 23 0.58
95 73 (76.8) T T + Ob
Males 10 15 0.42
Females 24 24 1
Choice behaviour with essentials oils
93 64 (68.8) T + PO T + Og (5 mg/ml)
Males 8 14 0.29
Females 22 20 0.88
88 64 (72.7) T + PO T + Og (10 mg/ml)
Males 10 9 0.80
Females 34 18 0.03
93 73 (78.5) T + PO T + Ob (5 mg/ml)
Males 10 9 0.80
Females 34 18 0.03
86 64 (74.4) T + PO T + Ob (10 mg/ml)
Males 10 14 0.54
Females 17 23 0.43
a
Responding insects include living individuals present in one of the two side areas of the tunnel
Fig. 1 Average number of eggs laid by T. absoluta females on a tomato
plant (control) versus a tomato plant associated either with
O. gratissimum or O. basilicum 2 days after releasing the insects in a
dual-choice flight tunnel. Values are number (Mean ± SE) of the total
eggs laid per combination (six replicates). ***P<0001
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with p-cymene (33.5 ± 2.3%), γ-terpinene (23.6 ± 1.6%), α-
terpinene (7.2 ± 0.5) and α-thujene (6.7 ± 0.8%) in
O. gratissimum and E-α-bergamotene (38.9 ± 10.3%), methyl
eugenol (26.1 ± 10.9%), E-β-ocimene (17.7 ± 4.0%) and lin-
alool (9.4 ± 3.2) in O. basilicum.Fourcompounds(α-pinene,
β-pinene, Myrcene, Limonene) were common in essential oils
and plants VOCs.
Discussion
Our study demonstrated the oviposition-deterring effect of
O. gratissimum and O. basilicum plants and their respective
essential oils against T. absoluta. However, no differences
were found in terms of the spatial distribution of these insects
in the flight tunnels. Therefore, we could not confirm that basil
plants and their essential oils repel adult leafminers.
Our results about Ocimum oviposition-deterring effect con-
firm those of previous studies. For instance, O. basilicum
ethanolic extracts negatively influenced the oviposition be-
haviour of another leafminer Phthorimaea operculella Zell.
(Lepidoptera: Gelechiidae) (Sharaby et al. 2009). Ocimum
spp. essential oils were previously shown to reduce the ovipo-
sition behaviour of Agrotis ipsilon H.(Lepidoptera:
Noctuidae) on cotton plants (Shadia et al. 2007). Moreover,
O. gratissimum,O. basilicum and O. sanctum L. extracts or
EOs were found to have a repellent and oviposition deterrent
effect on the insect pests (Dryophthoridae, Curcuilionidae,
Bostrichidae, Tenebrionidae, Bruchidae) of various stored
products (Asawalam et al. 2008; Ogendo et al. 2008;
Kiradoo and Srivastava 2010). The observed reduction in eggs
laid on tomato plants when associated with basil plants or
essential oils demonstrates that Ocimum spp. had a deterred
effect on the oviposition behaviour of T. absoluta females.
The ability of insects to locate host plants for feeding or
reproduction (eggs laying) is affected by the VOCs that it
perceives (Bruce et al. 2005; Webster et al. 2010;Soléetal.
2010; Bruce and Pickett 2011). In addition to directly repel-
ling insects, non-host odours also mask host plant volatiles
(Zhang et al. 2013). Thus, in both approaches, the compound
identified by our study in basil EO and VOCs might have
changed the chemical environment by masking tomato
VOCs and preventing T. absoluta females from recognising
these plants. Indeed, an oviposition deterrent effect of certain
compounds (such as β-caryophyllene, α-pinene, β-pinene,
limonene, terpineol-4, thymol and eugenol) has been
highlighted for several pests of stored food products
(Regnault-Roger and Hamraoui 1995; Ferrarini et al. 2008;
Tab l e 2 Chemical components of the essential oils found in
O. gratissimum (Og) and O. basilicum (Ob)
Compound Og (%)
a
Ob (%)
a
RI
b
α-thujene 3.6 –926
α-pinene 1.5 0.2 934
Camphene 0.2 0.1 951
Sabinene 0.2 –974
β-pinene 0.5 0.3 979
Myrcene 3.9 0.9 989
α-phellandrene 0.3 –1007
δ-3-carene 0.3 –1010
α-terpinene 1.9 –1018
p-cymene 20.4 –1026
Limonene 0.6 0.3 1031
E-β-ocimene –2.6 1047
γ-terpinene 16.9 –1060
p-cymenene 1.5 –1091
Estragol –73.9 1195
1,8-cineol –1.5 1034
Linalool –8.7 1101
Camphor –0.8 1150
Borneol 0.6 0.2 1169
Terpineol 4 1.9 –1183
Methyl thymol ether 0.5 –1231
Thymol 33.3 –1291
β-elemene –2.9 1393
E-caryophyllene –0.1 1426
α-bergamotene 0.2 –1435
γ-elemene –0.7 1437
α-guaiene –0.4 1440
α-humulene –0.2 1462
β-selinene –0.6 1496
γ-cadinene –0.9 1519
‘–’not-detected
a
Relative percentage
b
Retention index
Fig. 2 Average number of eggs laid by T. absoluta females on a tomato
plant associated with a rubber septum filled with paraffin oil versus a
tomato plant with a rubber septum filled with O. gratissimum or
O. basilicum essential oil 2 days after releasing the insects in a dual-
choice flight tunnel. Values are number (Mean ± SE) of the total eggs
laid per combination (six replicates), ***P< 0.001, ns (not significant)
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Chaubey 2012) and also on the potato tuber moth,
P. o p r c u l e l l a (Sharaby et al. 2009). This finding might explain
the difference in oviposition rates by T. absoluta females be-
tween associated and non-associated tomato plants.
Various explanations exist for the observed random distri-
bution of adult individuals in our study. For instance, during
the first moments of the trials (≤24 h), females made a choice
to orient towards one of the two tomato plants. This choice
was probably related to the perception of VOCs released by
the plants, with tomato plant VOCs having well-defined pro-
portions that characterise it and stimulate oviposition.
Previous studies have confirmed that T. absoluta females rely
on olfactory cues during host-searching and the assessment of
hosts as suitable larval substrate for oviposition (Proffit et al.
2011). A shorter observation period might have allowed us to
determine the most attractive area for ovipositing females.
Males are generally attracted by sex pheromones emitted by
females (Ramaswamy 1988); thus, their distribution was
probably influenced by the behaviour of females.
The two basil species had a similar effect on T. absoluta
behaviour. Both species could be used to manage this pest.
The fact that they had quite different chemical profiles (ac-
cording to the relative proportions) could be advantageous in
management schemes, as their use could be alternated to avoid
insect resistance or habituation. We suggest two approaches
for their use. First, EOs could be used as semiochemical dif-
fusers in greenhouse production. However, to optimise the
effectiveness of these EOs, it is important to assess the release
dynamics of EOs. Second, intercropping might be a more
suitable approach in developing countries, where tomato pro-
duction is primarily conducted under field conditions. These
two basil species are extensively used in the diet of many
families, particularly in West African countries (such as
Benin and Togo). In these countries, these basil species occu-
py an important place among vegetable crops, and are sold at
local markets throughout the year. Thus, farmers might be
more inclined to adopt using basil plants in association sys-
tems, providing benefits both for pest management strategies
and the commercialization and consumption of basil.
To date, few studies have focused on associating toma-
toes with the culturing of other plant species to manage
T. absoluta infestations. To our knowledge, the only exam-
ple in the published literature is the association of coriander
herbs (Coriadrum sativum L.) (Apiaceae) and gallant sol-
dier (Galinsoga parviflora Cav.) (Asteraceae) to reduce
T. absoluta abundance and increase auxiliary predators, like
ladybugs and spiders, when intercropping with tomato
plants (Medeiros et al. 2009). Our study was the first to
highlight the repellent effect of Ocimum species on
T. absoluta females. The use of these species as companion
plants might reduce damage to tomato plants by lowering
the number of eggs laid. In conclusion, our suggested ap-
proach is relatively easy to implement, and could be com-
bined with other methods in an integrated management
strategy against T. absoluta. This approach could also re-
duce the use of synthetic insecticides, especially in the rural
communities of West Africa.
Tab l e 3 Chemical volatile organic compounds emitted by
O. gratissimum (Og) and O. basilicum (Ob) plants,
Components Og (%)
a
Ob (%)
a
RT
b
α-thujene 6.7 ± 0.8 0.9 ± 0.6 10.8
α-pinene 2.5 ± 0.4 2.9 ± 1.9 11.0
Camphene 0.5 ± 0.1 –11.5
Sabinene 0.7 ± 00 0.9 ± 0.3 12.2
β-pinene 1.9 ± 0.6 2.5 ± 0.6 12.3
Myrcene 3.8 ± 0.2 5.1 ± 0.9 12.7
α-phellandrene 1.0 ± 0.1 1.2 ± 0.6 13.1
δ-3-carene 0.5 ± 00 1.2 ± 0.4 13.3
α-terpinene 7.2 ± 0.5 1.0 ± 0.5 13.5
p-cymene 33.5 ± 2.3 4.5 ± 3.1 13.8
Limonene 3.8 ± 0.3 6.0 ± 4.4 13.9
1,8-cineole –2.9 ± 1.5 14.0
Cis-ocimene 0.6 ± 0.1 1.1 ± 0.3 14.1
E-β-ocimene –17.7 ± 4.0 14.4
γ-terpinene 23.6 ± 1.6 1.5 ± 1.3 14.8
α-terpinolene –3.6 ± 2.2 15.7
p-cymenene 2.6 ± 0.2 –15.7
Linalool –9.4 ± 3.2 16.0
(E)-4,8-dimethyl-1,3,7-nonatriene –0.8 ± 0.3 16.5
Allo-ocimene 0.2 ± 0.1 2.0 ± 0.4 16.8
Neo-allo-ocimene –2.8 ± 0.4 17.2
Thymol methyl Ether 3.0 ± 1.3 –19.9
Thymol 4.4 ± 3.1 –21.4
α-cubebene –0.7 ± 0.3 23.1
Eugenol –0.5 ± 0.3 23.3
α-copaene –0.9 ± 0.1 23.8
β-elemene –1.0 ± 0.2 24.2
Methyl eugenol –26.1 ± 10.9 24.4
E-caryophyllene 0.8 ± 0.4 –25.0
E-α-bergamotene –38.9 ± 10.3 25.3
Aromadendrene –0.5 ± 0.0 25.5
E-β-farnesene –5.0 ± 1.3 25.7
α-humulene –1.0 ± 0.1 25.9
Epi-bicyclosesquiphellandrene –0.6 ± 0.2 26.1
Ar-curcumene –0.6 ± 0.0 26.4
β-Selinene 1.9 ± 0.8 –26.7
E-Methyl isoeugenol –2.3 ± 0.9 26.7
α-selinene 0.7 ± 0.3 –26.9
γ-cadinene –1.2 ± 0.2 27.3
1S,cis-calamenene –2.9 ± 0.7 27.5
‘–’not-detected
a
Relative percentage (Mean ± SD of triplicates)
b
Retention time
Environ Sci Pollut Res
Author's personal copy
Acknowledgements This research was funded by Erasmus Mundus
program. Tuta absoluta facilities were funded by the Service Public de
Wallonie (SPW–DGO3. D31-1344). The authors thank Frédéric Dresen
for technical support in tomato plant cultures and insect rearing; Danny
Trisman and Franck Michels for technical support in GC-MS analyses.
Compliance with ethical standards
Conflict of interest The authors declare that they have no conflict of
interest.
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