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Testing basil as banker plant in IPM greenhouse
tomato crops
Pia Parolin
a
, Cécile Bresch
a
, Christine Poncet
a
, Ricardo Suay-Cortez
a
& Louise Van
Oudenhove
a
a
French National Institute for Agricultural Research (INRA), University of Nice Sophia
Antipolis, CNRS, UMR 1355-7254 Institute Sophia Agrobiotech, 06900 Sophia Antipolis,
France
Published online: 29 May 2015.
To cite this article: Pia Parolin, Cécile Bresch, Christine Poncet, Ricardo Suay-Cortez & Louise Van Oudenhove
(2015): Testing basil as banker plant in IPM greenhouse tomato crops, International Journal of Pest Management, DOI:
10.1080/09670874.2015.1042414
To link to this article: http://dx.doi.org/10.1080/09670874.2015.1042414
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Testing basil as banker plant in IPM greenhouse tomato crops
Pia Parolin*,C
ecile Bresch, Christine Poncet, Ricardo Suay-Cortez and Louise Van Oudenhove
French National Institute for Agricultural Research (INRA), University of Nice Sophia Antipolis, CNRS, UMR 1355-7254 Institute
Sophia Agrobiotech, 06900 Sophia Antipolis, France
(Received 22 May 2014; final version received 27 March 2015)
In order to reduce the use of pesticides, banker plants may be added to crop systems. They can improve the presence of
predatory arthropods by offering shelter or alternative food. In the present study, we analyzed basil as banker plant in a
greenhouse IPM tomato crop system testing different plant combinations in monocultures vs. dicultures. We measured the
influence of the presence of basil on the reduction of pest number, predator population stability, plant health and crop
yield. Significantly fewer pests were present in dicultures when basil was present, but the number of predators, crop yield
and leaf health was not statistically different between plant combinations. Under the given conditions, basil could not be
categorized as an efficient banker plant for Macrolophus pygmaeus.
Keywords: Ocimum basilicum; Macrolophus pygmaeus; Trialeurodes vaporariorum; Solanum lycopersicum ; whiteflies;
tomato; biocontrol plant; banker plant; biological pest control
1. Introduction
Basil (Ocimum basilicum L., Lamiaceae) is an aromatic
herb with repellent effects on different kinds of arthropods
(Bomford 2004; Del Fabbro and Nazzi 2008). Several
studies state its qualities as repellent or companion plant
to decrease aphid, thrips and hornworm attacks when
intercropped in fields, particularly of tomato crops
(Schader et al. 2005; Basedow et al. 2006; Sujatha et al.
2011; Parker et al. 2013). Jolliffe (1997) repor ts that inter-
crop yields averaged 13% higher with basil than in mono-
cultures. In okra fields in Ghana, intercropping with basil
caused 23% decrease in insect pests compared to pure
okra stands (Amoatey and Acquah 2010). Intercropping
basil with tomatoes in Brazil resulted in significantly
higher crop yields, higher than when intercropping with
other aromatic companio n plants (Carvalho et al. 2009).
Furthermore, basil does not compete with the crops in
terms of growth and nutrient use (Kuepper and Dodson
2001; Bomford 2004 , 2009). It is, therefore, an efficient
biocontrol plant, i.e. one “which is intentionally added to
a crop system with the intent to enhance crop productivity
by mutual benefit, pest attraction and/or pest regulation,
and thus contributes to an increase of the efficiency of bio-
logical control systems, which finally leads to increased
crop productivity” (Parolin et al. 2014). Biocontrol plants
reduce the need to employ potentially dangerous pesti-
cides whose effects are known to bear economic, social
and environmental risks, for example, for bees (Carvalho
et al. 2009; Johnson et al. 2013).
A particular type of biocontrol plants is banker plants
which improve the presence of predatory arthropods
by offering shelter and sometimes alternative food (Diaz
et al. 1999; Frank 2010; Huang et al. 2011; Cano et al.
2012). Ideally, the natural enemies of the pests which are
present in the system establish regular reproducing popu-
lations on the banker plants in a crop system. This way,
the banker plants enhance the efficiency of biological
control because the predators are ready to attack the pests
as soon as they appear on the crops in the IPM system
(Cano et al. 2012). The need for frequent inundative
releases and the expenses of biological control are then
reduced (Enkegaard 2005; Frank 2010; Huang et al.
2011). The potential of basil to act as banker plant is indi-
cated by its capacity to act as summer refuge for parasit-
oid populations (Hoelmer and Goolsby 2002; Cano et al.
2012). In preliminary experiments, we found that the
common predatory mirid bug Macrolophus pygmaeus
Rambur (Hemiptera: Miridae) replicated well on basil in a
greenhouse in the Mediterranean climate of Southern
France. Despite the fact that it is a zoophytop hagous spe-
cies that can feed not only on other arthropods but also on
plants, it is an important and well-established biocontrol
agent to control the widespread greenhouse whitefly pest
Trialeurodes vaporariorum Westwood (Hemiptera:
Aleyrodidae) which attacks tomato crops. Tomato, Sola-
num lycopersicum Mill. (Solanaceae), is traditionally an
important crop in greenhouses of the Mediterranean
region in Southern France. Therefore, we test the effi-
ciency of basil as banker plant for the predator M. pyg-
maeus to control the whitefly T. vaporariorum on tomato
crops. The main question is whether basil is a suitable bio-
control plant enhancing crop productivity and health, and
more specifically, if it acts as banker plant for predatory
insects in a tomato crop greenhouse system in Mediterra-
nean climate. Our hypothesis is that basil directly favors
the presence of a stable reproducing population of
*Corresponding author. Email: Pia.Parolin@sophia.inra.fr
Ó 2015 Taylor & Francis
International Journal of Pest Management, 2015
http://dx.doi.org/10.1080/09670874.2015.1042414
Downloaded by [Pia Parolin] at 23:52 29 May 2015
predators, indirectly contributes to a significant reduction
of whitefly pests in the tomato crop greenhouse system
and enhances crop fruit production and plant health. If
basil is an efficient banker plant in this species combina-
tion which is commonly used in IPM in the Mediterranean
region, we hypothesize that the presence of basil plants in
the greenhouse crop system has (Figure 1)
(1) direct positive effects on the population of preda-
tors which increases due to favorable conditions
provided by basil plants and cause a higher pres-
ence of predators when basil is present;
(2) indirect negative effects on the population of pests
which decreases, via the increase of the popula-
tion of predators on basil, causing (a) lower pest
numbers when basil is present, which is (b) due to
higher predator presence;
(3) indirect positive effects on crop leaf health: as less
pests are present, less leaves should be attacked;
and
(4) indirect positive effects on crop productivity as
measured by the number of tomatoes produc ed.
The intention was to include the whole complexity of
the entire plant and follow realistic growth conditions,
like those employed by the local producers. The final goal
is to enhance IPM methods in order to reduce the use of
pesticides by farmers in the region.
2. Materials and methods
We set up an experiment in a greenhouse using whole
plants in separate cages in order to test the effects of the
presence of basil. Pests were present in all cages. The
treatments differ ed in the presence or absence of basil
monoculture (two tomatoes or two basil plants) vs. dicul-
ture (one tomato C one basil plant) in each cage and in
the presence or absence of predators, so that the indirect
effects on the pests can be better understood. Basil was
employed as banker plant and tomato, S. lycopersicum
Mill. (Solanaceae), as crop plant. The plants were placed
in separate cages in three plant combinations: one dicul-
ture (basil C tomato, hereafter referred to as BT) and two
monocultures (basil C basil, tomato C tomato, hereafter
referred to as BB and TT, respectively). Half of the plants
were inoculated with pests (T. vaporariorum, 30 individu-
als) and predators (M. pygmaeus, 6 individuals), and half
with pests (30 individuals) alone. The treatments were
placed in separate cages (size: 2 £ 1 £ 1 m) made of mus-
lin (Figure 2) placed in a greenhouse covered with single
glass. A randomized block design was used for each treat-
ment. Adding up to 30 cages in total, we had five
Figure 1. Basil as banker plant and possible interactions with the pest (whitefly), beneficial (predatory mirid bug) and crop plant
(tomato) (adapted from Parolin et al. 2014). Simplified scheme of the interactions with a banker plant: one direct positive effect of preda-
tors, two indirect negative effects on the population of pests, three indirect positive effects on crop leaf health and four indirect positive
effects on crop productivity. Since the beneficial in this particular case is a zoophytophagous species which can also feed exclusively on
plants, the tritrophic interactions regulating the plantpreypredator relationships can be more complicated than those simplified in our
scheme.
2 P. Parolin et al.
Downloaded by [Pia Parolin] at 23:52 29 May 2015
replicates for each of the six combinations: BT with or
without predator (hereafter referred to as C/¡ predator),
BB C/¡ predator, and TT C/¡ predator.
After seven weeks, we counted the number of indi-
viduals o f predators and pests on all plants. We sepa-
rated between eggs, larvae and adults of the whitefly
pests. We counted the total number of adults and larvae
of the predatory insects as they were difficult to distin-
guish. The predator M. pygmaeus lays the eggs inside
the leaf parenchyma, thus they were not visible on the
leaf surfaces and were not counted. To count the flying
insects (adult pests, larvae C adults of predators), we
placed a yellow sticky trap in each cage. For detailed
counting of larvae and eggs (pests only), we randomly
harvested 15 leaves per basil and 15 leaflets per
tomato from each plant, transferred them cautiously into
zipplasticbagsandtookthemtothelabwherewe
counted them using a lens with a magnificat ion from
10xto30x.
2.1 Raising of plants and insects
Seeds of tomato and basil were sown in a mixture of 1:3
perlite and 2:3 loam in 20-cm pots, kept in a greenhouse
(25 § 2
C, RH 70% § 10%) and watered daily. The
plants were put in 1.3-l pots with 2/3 soil and 1/3 sub-
strate. When the experiment started, the plants were about
20 cm tall. Adult T. vaporariorum were collected from
greenhouse colonies on roses and bred on tomato plants.
Adult M. pygmaeus were ordered at Biotop, Valbonne,
France.
2.2 Inoculation of the plants
The treatments were isolated from one another using cages
made of fine mesh material (Figure 2) in order to prevent
insect transfer between treatments in the greenhouse.
Pesticide applications were strictly avoided. In every cage
of the modality (half of the plant combinations), the plants
were pre-inoculated with 6 M. pygmaeus predators (3
males and 3 females). As one week of inoculation is con-
sidered as a necessary introduction period (Ridray et al.
2001), after eight days we inoculated the plants with 30
individuals of whiteflies T. vaporariorum per cage, in all
cages. The whiteflies were placed in an open petri dish on
the floor of the cage. This was the beginning point of the
experiment which lasted seven weeks thereafter.
2.3 Environmental conditions
A fog and cooling system (Aria) was used to maintain cli-
matic conditions stable in the greenhouse around 25
Ctem-
perature and 70% relative air humidity. Additional shading
was rarely used as the cages already filtered much light. The
plants were watered and fertilized with a drip irrigation sys-
tem (set point values for EC D 1.5 ms, pH D 6).
2.4 Plant yield and leaf health
We counted the number of fruits on the tomato plants in
different plant combinations at the end of the experiment.
Plant growth data (height, leaf number) was not useful as
the tops of the plants had to be cut twice because the
tomatoes grew too high in the cages. We est imated the
Figure 2. Cages in the greenhouse at INRA Sophia Antipolis.
International Journal of Pest Management 3
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healthiness of plant leaves by choosing one of the two cat-
egories: healthy leaves where leaves hosted no whitefly
larvae and were green without signs of pest attack, or
damaged leaves where parts of the leaves were infested
by larvae and presented yellowish areas as a consequenc e
of pest attacks.
2.5 Statistical analysis
Different tests were used to analyze the four hypotheses.
When the aim was to analyze a single effect or two effects
without interaction, non-parametric KruskalWallis tes ts
were used. When several effects and their potential inter-
actions were studied, data were analyzed with generalized
linear models (GLMs):
(1) The effect of plant combination (BB, TB or TT)
on the number of predators (sum of larvae
and adults) in the cage was analyzed with a
KruskalWallis test.
(2) The effect of plant combination on the number of
pests was determined at the cage level for adults,
and at the plant level for both eggs and larvae. For
each level, data were analyzed by fitting a nega-
tive binomial GLM in order to model count data
taking over-dispersion into account. At the cage
level, the GLM considered the number of pest
adults per cage as the response variable, the pres-
ence of predator (binary variable) and the plant
combination as explanatory variables. At the plant
level, the GLM considered the number of pest
(larvae and eggs) as the response variable, and the
presence of predator, the plant (tomato or basil)
and the type of culture (monoculture or diculture)
as explanatory variables. For both GLM, a full
model was built including all explanatory varia-
bles and interactions. Then, non-significant effects
were progressively removed until obtaining the
most parsimonious model. Model selection was
based on the comparison of the resulting scaled
deviances with x
2
tests. Once the best supported
model was selected, the differences between the
different levels of each variable were identified
with Student’s t-tests.
(3) The effect of plant combination on crop leaf
health was analyzed with non-parametric tests. In
the cages containing two tomato plants, the num-
bers of leaves either healthy or damaged leaves
were summed. In each cage, the percentage of
damaged leaves was determined, and its variation
according to the type of culture and the presence
of predators was tested with KruskalWallis tests
with a D 0.025 for Bonferroni correction.
(4) The effect of plant combination on the number of
tomatoes assessed with non-parametric tests. The
mean number of tomatoes produced per cage was
tested according to both the presence of predator
and the type of culture with Kruskal–Wallis tests
with a D 0.025 for Bonferroni correction.
All analyses were carried out using R 3.1.1 (R Core
Team 2013).
3. Results
3.1 Abundance of predators
After seven weeks, the number of predators in the cages
was not significantly different between the three plant
combinations (Kruskal–Wallis x
2
(2) D 0.41, p D 0.81).
There were about seven M. pygmaeus predators per cage,
including larvae and adults (Figure 3).
3.2 Abundance of pest species
The presence of basil in the cage did not affect the number
of adult T. vaporariorum found on the yellow sticky trap.
Indeed, the number of adult whiteflies was not signifi-
cantly different between the three plant combinations
(x
2
(2) D 2.06, p D 0.36). However, the presence of M.
pygmaeus predators in the cage decreased the number of
adult T. vaporariorum pests (x
2
(1) D 13.12, p < 0.001),
(Figure 4).
At the plant level, whiteflies showed a clear preference
toward tomato plants (Figure 5). In the treatment without
M. pygmaeus predator, eggs and larvae of T. vaporario-
rum were more abundant on tomato plants than on basil,
regardless of the type of culture (monoculture: t D 2.86, p
D 0.004; diculture: t D 6.24, p < 0.001). Moreover, when
given a choice (in diculture), whiteflies prefer to lay eggs
on tomato plant rather than on basil (Figure 5). Indeed,
there were less eggs and larvae of T. vaporariorum on
basil in diculture than in monoculture (t D 2.47, p D
0.01), and vice versa on tomato plants (t D 2.09, p D
0.04). The basil seemed to have a repulsive effect on
whiteflies. In this experimental setup with two plants per
Figure 3. Box plot of the number of M. pygmaeus predators
(adults C larvae) in the different plant combinations after seven
weeks of experiment. BB: basil C basil, BT: basil C tomato, TT:
tomato Ct omato.
4 P. Parolin et al.
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cage, when predators were absents, diculture led to an
increase of whiteflies pests on tomato plant (Figure 5).
In diculture, the inoculation of plants with M. pyg-
maeus predators highly reduced the number of eggs and
larvae observed after seven weeks (Figure 5). This reduc-
tion was observed both for tomato plant (t D 3.00, p D
0.003) and basil (t D 5.00, p < 0.001). In basil monocul-
ture, the number of preimaginal stage of T. vaporariorum
was also reduced by the presence of M. pygmaeus preda-
tors (t D 3.39, p D 0.002). However, this reduction was
not significant in tomato plant monoculture (t D 0.48, p D
0.63). On tomato plants, the presence of M. pygmaeus
predators seemed to reduce significa ntly the population of
T. vaporariorum only in cages where basil was present.
However, since the number of preimaginal stage of T.
vaporariorum was much higher in the presence of basil
without M. pygm aeus, the abundance of T. vaporariorum
eggs and larvae in the presence of M. pygmaeus
was not different between monoculture and diculture
(t D 0.79, p D 0.43).
3.3 Damaged leaves
After seven weeks, the percentages of damaged leaves of
tomato plant was significantly lower in the presence of M.
pygmaeus predators (Figure 6; Kruskal–Wallis x
2
(1) D
5.33, p D 0.02). However, the plant health was not signifi-
cantly different between diculture and monoculture (Krus-
kalWallis x
2
(1) D 0.37, p D 0.54).
3.4 Crop yield: number of tomatoes produced
The number of tomato fruits did not differ significantly
between monoculture and diculture (KruskalWallis
x
2
(1) D 0.05, p D 0.82), nor between the treatments with
or without predators (Kruskal–Wallis x
2
(1) D 0.01, p D
0.94) (Figure 7).
Figure 4. Box plot of the number of adult whiteflies found in
the cage after seven weeks of experiment according to the pres-
ence or absence of predators. Colors indicate the different plant
combinations: white D BB (basil C basil), dark grey D BT (basil
C tomato), light grey D TT (tomato C tomato). Statistically sig-
nificant differences are marked with stars (
: p-value < 0.05;
:
p-value < 0.01;
: p-value < 0.001).
Figure 5. Box plot of the number of preimaginal stages of
whiteflies found on each plant according to the plant species,
and the cage treatment. BB: basil C basil, BT: basil C tomato,
TT: tomato C tomato. Colors indicate the absence (light grey) or
presence (dark grey) of M. pygmaeus predators in the cage. Sta-
tistically significant differences are marked with stars (n.s.: p-
value > 0.5;
: p-value < 0.05;
: p-value < 0.01;
: p-value
< 0.001).
Figure 6. Box plot of the percentage of damaged leaves (in %
of all observed leaves) on the tomato plants after seven weeks of
experiment according to the presence or absence of predators.
Colors indicate the different type of culture: dark grey D dicul-
ture (TB: basil C tomato), light grey D monoculture (TT: tomato
C tomato). Statistically significant differences are marked with
stars (
: p-value < 0.05;
: p-value < 0.01;
: p-value <
0.001).
International Journal of Pest Management 5
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4. Discussion
Our experiment showed that basil has an effect on the
control of whitefly pests in tomato crops employing the
predator M. pygmaeus. However, we could not show that
the predators produced more stable populations when
basil was present. Furthermore, the presence of basil did
not result in increased plant health and crop yield. There-
fore, despite the negative development of pest popula-
tions, with the present experiment we cannot define basil
as an efficient banker plant. Thus, our initial hypotheses
(Figure 1) are refuted. The results were as follows:
(1) No effects on the populations of predators: no
higher presence of predators when basil was
present.
(2) The negative effects of the predators on the popu-
lation of the pest species were higher in diculture,
probably due to the higher abundance of pests on
tomato plants in diculture.
(3) No indirect positive effects on crop leaf health.
(4) No effects on crop productivity as the number of
tomatoes produced was the same in the
treatments.
4.1 Stability of predator populations
The number of predators did not vary in a significant way
between different plant combinations, indicating that basil
did not provide the required increase of the predator popu-
lation. However, despite the same predator population
size, predator effect on tomato plant was higher in the
diculture (TB) than in the monoculture (TT). This result is
consistent with tests of different host plants for the preda-
tor M. pygmaeus. In Perdikis and Lykouressis (2000,
2004), eggplants appeared globally as a better host plan t
than tomato for M. pygmaeus. Perdikis and Lykouressis
(2000) underscored the successful development of the
predator on tomato, even in the absence of prey. The fact
that M. pygmaeus is zoophytophagous could have direct
impacts on the tomato crops, depending on relative preda-
tor-to-prey abundance (Casta
~
n
e et al. 2011). However,
other studies showed that the predatory bug could not
increase in number on tomato without prey, although it
could survive for a relatively long period (Perdikis and
Lykouressis 2004).
4.2 Reduction of the number of whiteflies
In the combinations where previously inoculated predators
were present, the number of whiteflies was reduced in the
combinations which included basil. This corresponds to the
requirements of a biocontrol plant. However, when preda-
tors were absent, more whiteflies were found on tomato in
the diculture (TB) than in tomato monoculture (TT), con-
tradicting its efficiency as biocontrol plant. Indeed, there
was a strong repellent effect of basil on whiteflies. There-
fore, when given a choice (in diculture TB), whiteflies pre-
fer tomato plant rather than basil. However, at a cage level,
the number of whiteflies was similar between different
plant combinations. The question, if the presence of basil
was responsible, indirectly, for the pest reduction when
predators were present, remains open here.
4.3 Crop health and yield
If basil was a good banker plant, significant differences
should have been detected in plant health in terms of per-
centage of attacked leaves of the tomato plants between
the monoculture and the diculture, which, however, was
not the case. If basil indirectly provided a favorable envi-
ronment for predators and protection from pests, then the
leaves of tomato should have been less attacked when
basil was present in the combination, which we cannot
state. The percentage of healthy tomato leaves was higher
when predators were present, in both monoculture and
diculture (Figure 6), indicating the indirect positive role
of preda tors for crop leaf health (Figure 1, interaction 3).
Basil leaves were less attacked in the dicultures with
tomato, probably because the whiteflies preferred to feed
on the tomato leaves when present (pers. obs.). The num-
ber of tomatoes produced did not differ significantly
between the treatments. The presence of basil did not
show significant effects on the yield produced. Neverthe-
less, tomato production might have been restricted by
space limitation. The tomato plants needed to be cut twice
as their growth exceeded the capacity of the muslin cages.
Tomato yield must be tested in larger compartments, so
that the plants do not need to be cut.
At this stage, basil cannot be suggested for application to
the local farmers in order to reduce the use of pesticides in
tomato crops as biocontrol plant. The use of other local bio-
control plants is perhaps more efficient and may contribute
to the required reduction of pesticide use and, on the long
run, to a reduction of costs for the local producers (van
Figure 7. Box plot of the number of fruits on tomato plant with
or without predators, after seven weeks of experiment. Colors
indicate the different type of culture: dark grey D diculture (TB:
basil C tomato), light grey D monoculture (TT: tomato C
tomato).
6 P. Parolin et al.
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Driesche et al. 2010). It is important to focus on local spe-
cies and to avoid the introduction of potentially invasive
alien species. In the case of basil, which originates from
tropical Asia and grows well in the Mediterranean climate
(Labra et al. 2004;DeMasietal.2006), this danger might
have been neglectable as it has been cultivated for
5000 years and was already mentioned by Theophrastus 300
BC to grow in the Mediterranean area (De Masi et al. 2006).
Conclusion
Basil is not a suitable banker plant in a tomato crop green-
house system in Mediterranean climate. Th e presence of
basil did not seem to increase M. pygmaeus population.
The differences in pest population densities seem rather
due to a direct repellent effect of basil rather than to an
indirect effect of basil through a hypothetical increased
predator population. Neither plan t health nor crop produc-
tion was affected by the basil presence. Under the given
conditions, basil could not be categorized as an efficient
banker plant for M. pygmaeus. Notwithstanding, this plant
species might play an active role as a different kind of bio-
control plant, such as companion plant, repellent plant, or
winter refuge for beneficial arthropods.
Acknowledgements
This research was part of the project PURE: Integrated pest man-
agement in farming systems of major importance for Europe
supported by EC-FP7-KBBE-2010-4.
Disclosure statement
No potential conflict of interest was reported by the authors.
Funding
PURE: Integrated pest management in farming systems of major
importance for Europe [Grant Number EC-FP7-KBBE-2010-4].
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