Phytopathol. Mediterr.

(2006) 45, 238–246

Effect of essential oils and plant extracts on hatching, migration

and mortality of Meloidogyne incognita

SAID K. IBRAHIM, ABDALLAH F. TRABOULSI and SAMIH EL-HAJ

Faculty of Agricultural Sciences, Lebanese University, Beirut, Lebanon

Summary. The nematicidal activity of the essential oil/pure components and plant extracts of naturally grown
aromatic plant species against hatching, migration and mortality of the root knot nematode Meloidogyne incognita
was investigated. The pure components carvacrol, thymol, and linalool at 1, 2 and 4 mg liter-1 concentrations were the
most toxic against M. incognita second-stage juveniles (J2s) followed by terpineol and menthone. Hatching was
completely inhibited at low concentrations (2, 4 mg liter-1) of carvacrol, thymol, and linalool. Clove extracts (1 mg
liter-1) of Allium sativum significantly reduced hatching activity to below 8%, followed by flower extracts of Foeniculum
vulgare which reduced hatching to below 25%. These extracts were also toxic against J2s of M. incognita (LC50 43)
followed by leaf extracts of Pinus pinea, Origanum syriacum, Mentha microcorphylla, Eucalyptus spp. and Citrus
sinensis with an estimated LC50 of 44, 50, 65, 66 and 121 ppm respectively. Flower extracts of F. vulgare had the
highest effect on J2 mortality in sand (86%). The highest concentration of essential oils (6%) was detected in leaf
extracts of Origanium syriacum. Over 30 major components were identified in all the plant extracts tested.

Key words: aromatic plants, root-knot nematodes, carvacrol, thymol, linalool.

Introduction health (Dinham, 1993). There is therefore an ur-

gent need to replace pesticides with alternative
Agrochemicals have had a major role in improv-
means of control that are less toxic and more envi-
ing yields in food production. However, concern has
ronmentally friendly. Plant extracts containing
arisen about the negative impact that such chem-
volatile compounds, especially essential oils, have
icals have on human health and the environment.
been found to possess antimicrobial, insecticidal
Some pesticides have active ingredients that act
and nematicidal activity (Marban-Mendoza et al.,
as hormone disruptors, and may cause loss of fer-
1987; Thackray et al., 1990; Digrak, 1999; Okoko,
tility, carcinogenesis and mutagenesis. The wide-
et al., 1999). Certain plants kill or repel pests, dis-
spread application of agrochemicals to most cash
rupt their life cycle, or discourage them from feed-
crops has meant that pesticides are present in the
ing. Plant extracts may contain volatile and non-
ecosystems, aquifers and water systems of most
volatile components (Brown and Morra, 1997).
agricultural areas. In the long term, this could have
Some of these components may be detected at a
repercussions on both the environment and human
distance by olfaction and act as attractants or re-
pellents (Pickett and Stephenson, 1980).
Nematodes occur in virtually all soil types.
Corresponding author: S.K. Ibrahim Worldwide annual crop losses caused by nema-
E-mail: sibarhim58@[Link] todes, mainly the root-knot nematodes (RKN) Me-

238 Phytopathologia Mediterranea

 Effect of essential oils and plant extracts on Meloidogyne incognita

loidogyne spp. and the cyst nematodes Globodera was collected using a technique similar to that of
spp. and Heterodera spp., are estimated at approx- Fenwick (1949). Soil leachate was collected from
imately US$ 100 billion (Sasser and Freckman, pots containing the same soil but without plants.
1987). They attack a wide range of economically
important crops in horticultural, agricultural and Isolation of essential oils
forest systems. Nematode control is essentially Essential oils were extracted from plant mate-
preventive; because once a plant is parasitized it rial collected from different regions of Lebanon
is impossible to kill the nematode without also de- (Table 1) as described by Traboulsi et al., 2002.
stroying the plant. Controlling nematodes is very Between one hundred and 200 g of fresh plant
difficult and relies heavily on the use of soil fumi- material per species was used. Ethyl alcohol (99%)
gants with toxic and expensive nematicides. Me- was used as a solvent in the extraction process and
thyl bromide is effective against nematodes, but it 100 ml was used for each extraction run. Every
is harmful to human beings and to the environ- run consisted of four extraction cycles, 20 min each,
ment including beneficial organisms. There is an at a heating temperature of 100ºC and a cooling
urgent need to look for natural compounds with temperature of 50ºC. The plant extracts were re-
less toxicity and a low environmental impact. Leb- duced to a volume of 50 ml using a rotary evapora-
anon is committed to phasing out methyl bromide tor (90ºC) and stored at 4ºC until required. To de-
and to find alternatives to this biocide. Little in- termine the concentration of an essential oil in each
formation has been published on the status of nem- plant extract, 10 g of each plant part was weighed
atodes and their control in Lebanon. One way to and evaporated in an oven at 100ºC. The remain-
search for environmentally benign nematicidal con- ing material was then weighed and the percent-
trol agents is to screen naturally occurring com- age of essential oil determined.
pounds in plants. Lebanon is a country rich in
medicinal and aromatic plants (Nehmeh, 1978). Gas chromatography/mass spectrometry analysis
Volatile compounds from such plants, especially The plant material, air-dried at room temper-
their essential oils, have been found to possess ature for about one week, was subjected to hydro-
antimicrobial and insecticidal activity (Oka et al., distillation for 4 h according to the standard meth-
2001), but so far only a few essential oils and their od using a Clevenger-type distillation apparatus
components have been evaluated for their nemati- (Traboulsi et al., 2002). Plant components were
cidal properties. determined by gas chromatography (GC)
This paper reports the identification of the (Hewlett-Packard) coupled to an HP 5871A mass
chemical components of essential oils isolated from spectrometer detector and equipped with an on-
eighteen plants, and the effect that plant extracts column DBI (30 m⫻0.20⫻0.05 µm). The temper-
and pure essential oil components, from various ature programme consisted of an initial tempera-
medicinal and aromatic plants have on the hatch- ture of 50ºC; hold 3 min-1; ramp rate 3ºC min-1,
ing, migration and mortality of M. incognita sec- final temperature 220ºC; hold 65 min-1; column
ond-stage juveniles (J2s). flow rate 0.6 ml d’He/mi constant. The injection
temperature was 200ºC with an injection volume
of 2 µl/min. The mass spectrometer settings were:
Materials and methods
electron impact ionization mode with 70 eV elec-
Nematode inocula tron energy, scan mass range m/z 50–400. Detec-
Meloidogyne incognita isolates were main- tion temperature was 270ºC using the retention
tained on tomato (Lycopersicon esculentum L.) in time and peak area as a mean of measure (Table
pot cultures. Inocula of freshly hatched J2s were 2). Components were identified by comparing the
obtained from egg masses in distilled water. Only GC retention and mass spectra with those report-
J2s that hatched within a 24-h period were used ed in the literature. Pure essential oils of com-
for experiments. mercial origin were kindly supplied by Jean-Marie
Bessiere (Ecole Nationale Supérieure de Chimie
Tomato-root diffusate and soil leachate de Montpellier, France). Each oil was separated
Tomato-root diffusate from 4-week-old plants from water with a Pasteur pipette, dried by fil-

Vol. 45, No. 3 December, 2006 239

S.K. Ibrahim et al.

Table 1. Location, plant parts, harvest dates, and percentage of essential oils of the plants.

Essen-
Plant species Common name Location Plant part Harvest date tial oil
(%)

Andropogon nardus Lemon grass Mount Lebanon Leaves April 2005 2.1
Allium sativum Garlic Market Cloves April 2005 0.3
Chrysanthemum coronarium Chrysanthemum Mount Lebanon Whole plant April 2005 n.a.
Cinnamomum zeylanicum Cinnamom Mount Lebanon Bark April 2005 0.5
Citrus sinensis Sweet orange Bourghoulieh Leaves June 2000 0.25
Eucalyptus spp. Eucalyptus Hazmieh (Mount) Leaves April 2000 0.4
Foeniculum vulgare Fennel Tabaria (South) Flowers May 2000 1.0
Lavandula stoechas Lavander Pchemon (Mount) Flowers June 1998 0.8
Laurus nobilis Laurel Nabay Leaves April 2000 0.5
Matricara discoidea Chamomile Mount Lebanon Whole plant April 2005 0.9
Mentha microphylla Mint Mashgara (Bekaa) Leaves, flowers April 1998 1.1
Myrtus communis Myrtle Akkar Leaves September 1998 1.2
Origanum syriacum Oregano Mashgara (Bekaa) Leaves September 1998 6.0
Pelargonium graveolens Pelargonium Mount Lebanon Leaves April 2005 0.8
Pimpinella anisum Anise Mount Lebanon Seeds April 2005 2.6
Pinus pinea Pine Mashgara (Bekaa) Leaves April 2000 0.5
Pistacia lentiscus Mastic tree Mashgara (Bekaa) Leaves September 1998 0.2
Salvia officinalis Sage Mount Lebanon leaves, stem, flowers April 2005 1.3
Tagetes patula Marigold Mount Lebanon Leaves, stem, flowers April 2005 1.3

n.a., Data not available.

tration over anhydrous sodium sulphate and Effect of plant extracts on J2 migration in sand
stored at -20°C in a sealed dark bottle until anal- One hundred freshly hatched J2s were placed
ysis. The oil yields (Table 1) were calculated rela- in cavity watch glasses containing 5, 10, 25, 50,
tive to the mass of dry plant material. 100 mg l-1 of each plant extract in 2 ml DW. After
2, 4, 8 and 24 hours the J2s were removed and
Effect of plant extracts with tomato-root diffusate on placed on a sand column (height 1.5 cm, sand par-
J2 hatching ticle size 250–500 µm) in plastic tubes (3.0⫻4.0 cm)
Bioassays were carried out as described by Ib- sealed at the bottom with a nylon mesh (53-µm
rahim et al. (1993), using four batches of 25 egg aperture). The column was placed upright in a cav-
masses in 2 ml of each test solution in cavity watch ity watch glass containing DW and kept at room
glasses at 25–27ºC. Distilled water (DW), soil lea- temperature (25–27°C). After 24 hours the number
chate (SL) and tomato-root diffusate (TRD), dilut- of J2s that had migrated through the mesh was
ed 1:3, and were used as control. TRD were dilut- recorded. The controls for all experiments without
ed with various concentrations (0.125, 0.250, 0.5, plant extracts were treated identically. This exper-
1, 2, 4 mg l-1) of plant extracts or pure components iment was conducted twice.
as treatments.
Hatched J2s were counted weekly for 6 weeks. Effect of plant extracts on mortality of J2s
Every week, a fresh solution with or without TRD Fifty newly hatched J2s were placed in cavity
plus plant extracts or pure components was added. watch glasses containing 5, 10, 25, 50 and 100 mg
At the end of each experiment, the egg masses were l-1 of each plant extract in 2 ml of DW. After 24
disrupted and the number of eggs containing un- hours, plant extracts were removed and the J2s
hatched J2s was counted to determine the percent- placed in DW and left to recover. The J2s were ex-
age of hatching. All treatments were replicated four amined after 24 hours and only J2s that were un-
times and each experiment was repeated twice. able to move were considered dead. This experi-

240 Phytopathologia Mediterranea

 Effect of essential oils and plant extracts on Meloidogyne incognita

Table 2. Percentage of major components of the essential oil in each plant species tested.

Plant species (plant part) Major components of essential oil (%)

Allium sativum (cloves) Allicina, allyl-methyltrisulphidea, diallyldisulphidea, diallytrisulphidea,

diallytetrasulphidea, allypropyldisulphidea
Cinnamomum zeylanicum (bark) Methyl chavicol (74%), 1,8 cineole (2.1%), linalool (2.6%), caryophyllene
(3.1%), eugenol (2.5%)
Andropogon nardus (leaves) Citral (75.2%), myrcene (18.5%), geraniol (16.9%), limonene (7.7%)
Citrus sinensis (leaves) Limonene (40%), linalyl acetate (37%), minolool (16%), bergaptene (4%)
Eucalyptus spp. (leaves) 1,8 cineol (30%), camphor (18%), α-pinene (19%), borneol (17%), borneol
acetate (5%)
Foeniculum vulgare (flowers) Limonene (63.6%), anethole (25.5%), fenchyl acetate (2.6%), α-pinene
(0.97%), myrcene (0.98%), estragole (1.1%)
Laurus nobilis (leaves) 1,8 cineole (44%), α-pinene (4.5%), β-pinene (2.2%), sabinene (8.9%), methyl
eugenol (1.9%)
Lavandula stoechas (flowers) Fenchone (40.1%), 1,8 cineole (11.7%), bornyl acetate (5.8%), myrtenyl
acetate (2.8%), myrtenol (2.1%), α-pinene (2%), viridiflorol (1.9%)
Matricaria discoidea (whole plant) Chamazulene (14%), farnesenea, farnesola
Mentha microphylla (leaves, flowers) Piperitenone (54.2%), pulegone (10.7%), piperitenone oxide (11.3%),
menthone (3.3%), 1,8 cineole (2.8%)
Myrtus communis (leaves) 1,8 cineole (40%), α-pinene (17%), linalool (9.9%), α-terpineol (7.9%), geranyl
acetate (4.5%), myrtenyl acetate (7%), α-terpinyl acetate (2%)
Origanum syriacum (leaves) Carvacrol (61%), thymol (21.8%), δ-terpinene (4%), p-cymene (5.5%),
myrcene (1.2%), α-terpinene (1.3%)
Pelargonium graveolens (leaves) Citronellol (21.7%), linalool (17.3%), geraniol (14%), menthone (1.1%)
Pimpinella anisum (seeds) Transanethole (85%), estragole (0.57%), linalool (1.5%), α-terpineol (1.5%)
Pinus pinea (leaves) Limonene (74.6%) 1,8 cineole (4.3%), α-pinene (3.8%), myrcene (2.7%), β-
caryophyllene (3.4%)
Salvia officinalis (leaves, stem, flowers) α-thujene (26.8%), limonene (22.9%), linalyl acetate (17.4), ocimene (10.7%),
linalool (5.7%), myrcene (3.6%) α-pinene (2.5%)
Tagetes patula (leaves, stem, flowers) Linalool (26.8), limonene (22, 9%), linalyl acetate (17.4%), ocimene (10.7%)

a
, Data not available.

ment was repeated twice. Data were subjected to common in a number of plants: Cinnamomum zey-
Sigma Stat 2.0 analysis of variance (ANOVA), and lanicum, Laurus nobilis, Lavandula stoechas, Men-
the LC was calculated using probit analysis. tha microphylla, Myrtus communis, Pelargonium
graveolens, Pimpinella anisum, Pinus pinea, Salvia
officinalis and Tagetes patula. Myrtenyl acetate was
Results
detected in both L. stoechas and M. communis. The
Plant extracts highest percentage of essential oils occurred in P.
The highest concentration (6%) of essential oil anisum (transanethole, 85%), followed by Cymbopo-
was detected in leaf extracts of Origanum syria- gon citrates L. (citral, 75.2%), Cinnamomum zeyla-
cum L. (Table 1). nicum (eugenol, 71.3%), and Origanum syriacum (car-
In the plant extracts more than 30 major compo- vacrol, 61%) (Table 1). The lowest percentage was
nents were detected. 1,8- cineole and linalool were detected in P. anisum (estragole, 0.57 %) (Table 1).

Vol. 45, No. 3 December, 2006 241

S.K. Ibrahim et al.

Effect of pure components with or without tomato- no significant differences between P. pinea, O.
root diffusate on J2 hatching syriacum, M. microcorphylla and Eucalyptus spp.
The effect of pure components on hatching of However, C. sinensis was significantly different
J2s was related to component concentration (Ta- from all the other essential oils.
bles 3 and 4). More than 95% of the J2 hatched
when the egg masses were incubated with TRD Effect of plant extracts on mortality of J2s
and DW alone, but when the egg masses were Even at the lower concentrations all the six
exposed to the pure components with TRD at 1 plant extracts began to affect J2s after minutes
mg l-1 for 6 weeks, the response varied according of exposure, with slower body movements becom-
to the component. Thymol, carvacrol, and linalool ing apparent. As concentrations increased from 5
almost completely inhibited hatching, while men- to 100 mg l-1 there was a corresponding increase
thone and (1S)-(-)-α-pinen reduced hatching to 27 in J2 mortality (Table 7). At a concentration of
and 39% respectively. 1,8 cineol or cineole and 100 mg l-1, F. vulgare had the greatest effect on J2
(1R)-(+)-α-pinen were less effective against hatch- mortality, followed by P. pinea, O. syriacum and
ing (Table 3). Hatching did not exceed 6% with Eucalyptus. C. sinensis had the lowest effect on
carvacrol, thymol and linalool at 2 mg l-1, but at 4 mortality.
mg l-1 hatching was completely inhibited from the
first week with carvacrol, thymol and linalool and
was reduced significantly to 15% with menthone Discussion
(Table 4). At concentrations of 0.125, 0.250, 0.5
Allelochemicals are plant-produced compounds
mg l-1, however, carvacrol, linalool, thymol and
that affect the activity of other organisms and are
menthone did not significantly affect hatching
thought to be toxins and secondary metabolites that
(data not shown). Hatching in SL did not exceed
act as attractants or deterrents (Dodds, 1996;
46% (Tables 3 and 4).
Brown and Morra, 1997). Sudan grass, for exam-
Effect of plant extracts with or without tomato-root ple, contains a chemical called d’hurrin that de-
diffusate on hatching of J2s grades into hydrogen cyanide, which is a powerful
The effect of 11 of the essential oils on the hatch- nematicide (Wider and Abawi, 2000). In our study
ing of J2s is shown in Table 5. Essential oil of A. all the essential oil/plant extracts tested had a
sativum had the greatest inhibitory effect, reduc- nematicidal effect and affected the hatching of M.
ing hatching to less than 8% followed by F. vul- incognita J2s even at the lowest concentration (1
gare, which reduced hatching to 25.3%. The effect mg l-1). Inhibition of hatching persisted as long as
of A. sativum was detected from the first week, the eggs were exposed to the essential oil/plant
when hatching was only 3.3% and increased very extracts. When the eggs were transferred to TRD
slowly during the remaining 5 weeks. F. vulgare alone, hatching resumed, but the degree of recov-
also had an inhibitory effect, but it was not fully ery was affected by the essential oil/ plant extracts
expressed until the second week. C. coronarium, concentrations to which the eggs had previously
P. palestina, M. discoidea and T. patula did not sig- been exposed.
nificantly inhibit hatching as compared with the Essential oils from various plants have shown
control. In the majority of the treatments, the promise as sources for nematicides. Most of these
greatest reduction in hatching occurred in the first plants are aromatic and culinary herbs that con-
two weeks. tain nematicidal compounds such as carvacrol and
thymol (Oka et al., 2000). Over 20 major com-
Effect of essential oils on the migration of J2s pounds of the essential oils were identified, but
Toxicity lines were established for each of the the most toxic against M. incognita J2s were car-
six plant extracts and the LC50 and LC90, slope val- vacrol, linalool, thymol and menthone. At very low
ues were calculated (Table 6). Extracts from F. vul- concentrations (1 mg l-1) several oils immobilized
gare were the most toxic (LC50 43 ppm) followed by the juveniles and some also reduced hatching. Es-
extracts of P. pinea, O. syriacum, M. microcorphyl- sential oils from: caraway, fennel, applemint,
la, Eucalyptus spp. and C. sinensis with an LC50 of spearmint, Syrian oregano, and oregano showed
44, 50, 65, 66 and 121 ppm respectively. There were a very high nematicidal activity (Oka et al., 2000).

242 Phytopathologia Mediterranea

 Table 3. Effect of pure components concentrations on hatching of second stage juveniles of Meloidogyne incognita.

Pure component concentration (1.0 mg l-1 a.i.) b + TRDa

Exposure DWa SLa TRDa
time (week) Carvacrol Thymol Linalool Terpineol Menthone 1,8 Pinen (1S) Pinen (1 R)
Cineole -(-)-α -(+)-α

1 244 a 739 b 504 a 40 c 7c 22 c 117 d 427 d 835 ade 232 de 424 de

2 856 811 1287 224 8 234 582 765 1210 681 860
3 1029 1394 2991 54 4 2 1684 170 405 494 903
4 4459 728 1762 7 5 17 364 7 125 569 1009
5 2043 190 1325 3 4 4 256 5 250 753 1609
6 114 45 712 0 0 0 62 1 125 675 325
Total 8745 3907 7869 328 28 301 3065 1374 2950 3404 5130
% hatch 95.4 46.0 98.1 3.8 0.5 5 35.0 27.0 56.0 39.0 59.0

a
DW, distilled water; SL, soil leachate, TRD, tomato root diffusate.
b
Data are the means of four replications. Data followed by the same letter are not significantly different at P<0.05.

Table 4. Effect of pure component concentrations on hatching of second stage juveniles of Meloidogyne incognita.

Pure component concentration +TRD b

Exposure DWa SLa TRDa 2 mg l-1 4 mg l-1
time (week)
Carvacrol Thymol Linalool Menthone Carvacrol Thymol Linalool Menthone

1 244 a 739 b 504 a 404 129 71 342 17 4 4 124

2 856 811 1287 82 1 52 176 3 0 0 274
3 1029 1394 2991 13 1 22 508 1 0 0 327
4 4459 728 1762 4 0 109 529 0 0 0 996
5 2043 190 1325 1 0 20 333 0 0 0.7 89
6 114 45 712 0 11 8 246 0 0 0 383
Total 8745 3907 7869 504 140 282 2134 21 4 4.7 2193
% hatch 95.4 46 98.1 5.3 1.5 6 20 0.2 0.04 0.04 15

Vol. 45, No. 3 December, 2006

a
DW, distilled water; SL, soil leachate, TRD, tomato root diffusate.
b
Data are the means of two replications. Data followed by the same letter are not significantly different at P<0.05.

243
Effect of essential oils and plant extracts on Meloidogyne incognita
S.K. Ibrahim et al.

Table 5. The effect of plant extracts at 1.0 mg l-1 on the hatching of second stage juveniles of Meloidogyne incognita
(accumulative percentage).

Hatching time a
Plant species (plant part)
Week 1 Week 2 Week 3 Week 4 Week 5 Week 6

Allium sativum (cloves) 3.32 4.1 6.2 6.8 7.3 7.6 a

Andropogon nardus (leaves) 17.35 33.3 37.2 38.3 38.5 38.6 b
Chrysanthemum coronarium (whole plant) 28.4 82.2 95.5 96.8 97.3 97.4 d
Cinnamomum zeylanicum (bark) 43.4 67.3 70.5 71.4 72.1 72.6 c
Foeniculum vulgare (flowers) 20.61 22.5 24.4 25.1 25.2 25.3 b
Matricaria discoidea (whole plant) 38.5 82.3 92.7 93.6 93.9 94.1 d
Pelargonium graveolens (leaves) 15.65 42.1 48.5 49.9 50.2 50.5 bc
Pimpinella anisum (seeds) 33.94 43.3 44.7 45.7 46.1 46.3 bc
Pistacia palestina (leaves) 46.7 82.1 94.4 96.5 96.8 96.9 d
Salvia officinalis (leaves, stem, flowers) 22.54 53.5 64.2 65.4 65.6 65.8 c
Tagetes patula (leaves, stem, flowers) 36.1 70.6 86.4 89.9 91.4 92.0 d
Control (TRD) 42.1 82.8 96.3 97.0 97.3 97.31 d

a
Data are the means of two replications. Data followed by a common letter are not significantly different at P<0.05.

Table 6. The effect of plant extracts on the migration of second stage juveniles of Meloidogyne incognita.

Plant species LC50 (ppm) LC90 (ppm) LC50 ppm (95% confidence limits) Slope Slope function

Citrus sinensis 121 260 146.168 - 100.165 1.208 3.324

Eucalyptus sp. 66 132 80.058 - 54.410 1.213 3.015
Foeniculum vulgare 43 105.5 50.697 - 36.471 1.179 2.523
Mentha microphylla 65 160 82.615 - 51.140 1.271 3.888
Origanum syriacum 50 124 61.900 - 40.387 1.238 3.342
Pinus pinea 44 129 54.824 - 35.313 1.246 4.178

Data are the means of two replications.

Table 7. Mortality of second stage juveniles (J2s) of Meloidogyne incognita after 24 h exposure to plant extracts.

Mortality (%) at different concentrations (mg l-1)a

Plant species J2s tested (No.)
5 10 25 50 100

Foeniculum vulgare 79 1 8 24 63 86
Pinus pinea 81 10 19 37 63 69
Origanum syriacum 82 5 16 29 52 65
Mentha microphylla 85 6 21 31 45 56
Eucalyptus sp. 79 3 10 28 46 63
Citrus sinensis 84 4 12 14 25 29

a
Data are means of two replications.

244 Phytopathologia Mediterranea

 Effect of essential oils and plant extracts on Meloidogyne incognita

Concentration of 1,8-cineole (40%) is high in 1997). Moreover, T. patula was the most effective
many plants (Kojima et al., 1998; Obeng-Ofori and plant in lowering RKN populations (Ogden, 1997).
Reichmuth, 1999). In our study 1,8 cineole, car- In our study marigold crude extracts stimulated
vacrol, linalool, and thymol were detected in sev- the hatching of M. incognita J2s, however, it may
eral plants but at different concentrations. The have affected the life cycle after invasion. Belcher
highest concentration of carvacrol (61%) was de- and Hussey (1977) reported that T. patula acted as
tected in O. syriacum plant extracts. Some pure a trap crop to M. incognita, preventing giant cell
essential oils (carvacrol, linalool, thymol and men- initiation. On the other hand, a minimum concen-
thone) exhibited nematode-suppressive character- tration of 1 mg l-1 of A. sativum and F. vulgare sig-
istics equivalent to that of cadusafos, a synthetic nificantly (P<0.05) decreased M. incognita J2s to
pesticide (Ibrahim and Haydock, 1999). The EC 7.6 and 25% respectively. Leaf extracts of Crota-
value for essential oils against hatching M. incog- laria virgulata subsp. grantina had a nemostatic
nita J2s was very low (0.125-4.0 mg l-1) compared effect on the J2s of M. incognita at the same low
with 0.49 mg l-1 for oxamyl and 4.63 mg l-1 for aldi- concentration (Jourand et al., 2003). Pérez et al.
carb against Globodera rostochiensis on agar (2003) reported that C. coronarium extracts also
plates. A concentration of 0.1 and 0.5 mg l-1 of ox- reduced hatching, J2 survival and the reproduc-
amyl inhibited the orientation of M. incognita to- tion rate of M. artiellia in vitro, but in our research
ward the host roots (Wright et al., 1980) and af- C. coronarium stimulated the hatching of M. in-
fected the orientation of G. rostochiensis J2s to- cognita J2s. Leaf powder of rock fleabane (Inula
ward potato roots (Evans and Wright, 1982). The viscose) at a concentration of 0.1% in sand reduced
mode of action of essential oils against nematodes hatching of J2s of M. javanica and the citrus nem-
is still not fully understood. In insects, several es- atode (Tylenchulus semipenetrans), though the
sential oils inhibit acetylcholinesterase activity stem-bulb nematode Ditylenchus dipsaci was un-
(Ryan and Byrne, 1988). Some common components affected (Oka et al., 2001). Clove extract and Nim-
of essential oils such as carvacrol, t-anethole, and becidine did not show any potential to control Aphe-
thymol have been found to be insecticidal (Isman, lenchoides fragariae (Jagdale and Grewal, 2002).
1999, Traboulsi et al., 2002) and some, to be nema- Further research is required to explore the natu-
ticidal (Oka et al., 2000). Zuckerman and Esnard ral biocidal activity of plant extracts against nem-
(1994) reported that some plant extracts with atodes.
nematicidal activity may also affect behaviour, such
as the nematode’s ability to recognise the host.
Acknowledgements
As a result of enzyme degradation, Brassica spp.
like rapeseed and mustard release bio-products The authors thank the Lebanese National Coun-
that have a nematode-suppressive effect by inter- cil for Scientific Research for financial support, and
fering with the nematode reproductive cycle Elias Chnais, Charbel Hage and Hassana Ghanem
(Brown and Morra, 1997). Rapeseed and Sudan for technical assistance.
grass green manures grown prior to potatoes in a
field reduced RKN in potatoes by 86% (Stark, 1995).
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Accepted for publication: November 13, 2006

246 Phytopathologia Mediterranea