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Scientia Horticulturae 89 (2001) 103–113

Role of plant age in the resistance of Lycopersicon
hirsutum f. glabratum to the tomato leafminer
Tuta absoluta (Lepidoptera: Gelechiidae)
G.L.D. Leite1, M. Picanc¸o, R.N.C. Guedes*, J.C. Zanuncio
Departamento de Biologia Animal, Universidade Federal de Vic¸osa,
Vic¸osa, MG 36571-000, Brazil
Accepted 7 July 2000

Abstract
The objective of this work was to study the role of plant age in the resistance of tomato,
Lycopersicon hirsutum, to the tomato leafminer Tuta absoluta (Meyrick). Determination of the
levels of tridecan-2-one and undecan-2-one in L. hirsutum were made at three different ages (2–4
months after germination), as well as the leaf area, and the density and types of trichomes present
on L. hirsutum and L. esculentum. This information gathered was correlated with the following
biological characteristics of T. absoluta: rates of oviposition and egg eclosion; length of egg, larval
and pupal stages; mortality in the larval and pupal stages; pupal weight and proportion of females.
T. absoluta showed higher oviposition and egg eclosion, as well as lower mortalities and shorter
larval and pupal periods in L. esculentum than in L. hirsutum. The proportion of females obtained
was also higher on L. esculentum. The rate of egg eclosion and proportion of females as well as the
length of the larval stage and the mortality of larvae appeared to be affected by plant age. A higher
proportion of females, mortality of larvae and length of the larval period were obtained with older
plants of L. hirsutum, while the rate of egg eclosion was higher for L. hirsutum plants that were 3
months old. For L. esculentum, plant ageing increased larval mortality. It seems that the glandular
trichome density in L. hirsutum increased with plant age leading to an increase in the levels of
tridecan-2-one, which slows larval development. # 2001 Elsevier Science B.V. All rights reserved.
Keywords: Tomato plant; Tomato leaf miner; Tridecan-2-one; Undecan-2-one

*

Corresponding author. Tel./fax: þ55-31-899-2537.
E-mail address: guedes@mail.ufv.br (R.N.C. Guedes).
1
Present address: Departamento de Fitotecnia, Universidade Federal de Minas Gerais, Nu´cleo de
Cieˆncias Agra´rias, Av. Osmani Barbosa s/n, B. JK, Montes Claros, MG 39404-006, Brazil.
0304-4238/01/$ – see front matter # 2001 Elsevier Science B.V. All rights reserved.
PII: S 0 3 0 4 - 4 2 3 8 ( 0 0 ) 0 0 2 2 4 - 7

104

G.L.D. Leite et al. / Scientia Horticulturae 89 (2001) 103–113

1. Introduction
The tomato leafminer, Tuta (¼Scrobipalpuloides) absoluta (Meyrick) (Lepidoptera: Gelechiidae), is considered as one of the key pests of tomato
Lycopersicon esculentum in South America. Its control is based on intensive
insecticide applications (Guedes et al., 1994; Picanc¸o et al., 1996a,b). An
alternative strategy to reduce T. absoluta populations and their losses would be
the development of resistant commercial varieties. The wild tomato Lycopersicon
hirsutum f. glabratum (PI 134417) has been reported as resistant to this leafminer
(Giustolin and Vendramim, 1994; Leite et al., 1995). The resistance of L.
hirsutum to T. absoluta is usually attributed to the allomones tridecan-2-one and
undecan-2-one present in the leaf glandular trichomes of this tomato species,
which are absent in L. esculentum (Giustolin and Vendramim, 1994), but there is
very little work on the subject.
One of the limitations of using L. hirsutum f. glabratum (PI 134417) as a source
of resistance is that the resistance is affected by the plant growth and leaf
senescence. Schalk and Stoner (1976) and Kennedy and Sorenson (1985) found
that reduced resistance of L. hirsutum to Colorado potato beetle is due to the
reduction of the concentration of tridecan-2-one as leaves become older. The
objectives of this study were to identify the types and densities of trichomes,
quantify tridecan-2-one and undecan-2-one, and determine the effect of age (2–4
months after emergence) on the leaf area of L. hirsutum f. glabratum (PI 134417)
and how it affects the biology of T. absoluta.

2. Material and methods
Tomato plants were grown in 5 l polyethylene pots in greenhouse conditions.
The average minimum and maximum temperature were 11:7 0:8 and
35:0 0:4 C, respectively, and the photoperiod was 11:13 (L:D) during the
investigation. No supplemental light or heat was used. The 2 3 factorial
experiment was established in a randomized blocks design with 12 replicates for
the assessment of plant-related traits and 28 replicates for the assessment of insect
biological traits. Each replicate was constituted by one tomato plant in a 5 l
polyethylene pot. The influence of the following factors on the resistance of T.
absoluta were studied: tomato species (L. esculentum cv. Santa Clara and L.
hirsutum f. glabratum (PI 134417)) and plant age (2–4 months after emergence).
The daily recorded insect biology data (rates of oviposition and egg eclosion;
length of egg, larval and pupal stages; mortality in the larval and pupal stages;
pupal weight and proportion of females) as well as the levels of tridecan-2-one,
undecan-2-one, trichome density and leaf area were determined in leaves from
each third of the plant height and the average of these determinations was

G.L.D. Leite et al. / Scientia Horticulturae 89 (2001) 103–113

105

calculated for each plant and used for the statistical analyses. Two months old
plants presented flowers and small fruits sometimes, and 3 months old plants
presented flowers and non-mature fruits, while 4 months old plants presented
flowers and mature fruits.
Soil chemical analysis was made prior to the beginning of the experiment.
After this analysis, 200 mg of N and 100 mg of K/kg of soil were used for soil
fertilization. K was added as potassium chloride, which was mixed to the soil 1
month before starting the experiment. The N source was urea, added to the pots
as an aqueous solution 1 week before sowing. After the plants reached 2 months
of age, the fertilization with urea through aqueous solution was maintained, being
added to each pot a total of 0.30 g of urea. The N was applied in a single dose
for the 2 months old plants, but only 90 and 80% of the recommended N dose was
applied before sowing for the 2 and 4 months old plants, respectively. The
remaining N dose was applied as aqueous solutions in the 3 and 4 months
old plants after 30 and 60 days of sowing, respectively. The plants were daily
irrigated and enough water was provided to form a film at the soil surface of
each pot.
Groups of five 1 day old eggs of T. absoluta were placed on the third leaflet
fully expanded from each third of the plants with a total of 15 eggs/plant and 420
eggs/treatment. The plant branches containing the leaves with the insect eggs (in
their surface) were enclosed in cloth bags (0:2 m 0:28 m) and the length of the
egg incubation period, rate of eclosion, larval mortality, number of small and
large mines (mines smaller or larger than 0.5 cm in length were classified as small
or large mines, respectively) (Picanc¸o et al., 1995), length of the larval stage
(days), pupal weight and deformation, and sex ratio were daily evaluated. The
male and female pupae were identified, separated and placed in white 500 ml
plastic pots covered with cloth and incubated at 25 0:5 C and a 12 h
photoperiod until adult emergence (Giustolin and Vendramim, 1994). The pupae
were daily inspected until adult emergence for determination of the pupal
mortality and the length of the pupal period.
Another set of plants maintained in similar conditions to those used for the
insect studies were used for plant-related determinations. For extraction of
tridecan-2-one and undecan-2-one, 5 g of leaves were collected from each plant
in the morning and used for extraction with 50 ml of hexane (distilled) for 24 h.
The solution was decanted, dried over sodium sulfate and evaporated to dryness
at 308C in a rotatory evaporator. The oil was redissolved in 1 ml of hexane and
analyzed by gas chromatography using OV 17 (1%) packed in a 2 m glass
column. The chromatograph (Instrumentos GC, Sa˜ o Paulo, Brazil) was
programmed from 150 to 2208C at 68C/min. The injector and detector were
maintained at 260 and 2808C, respectively. The flow rates of H2/N2/air were 30,
30 and 300 ml/min, respectively. The tridecan-2-one and undecan-2-one
concentrations in the leaf hexane extracts were determined using standard

106

G.L.D. Leite et al. / Scientia Horticulturae 89 (2001) 103–113

calibration curves for both compounds (99% pure) obtained from Aldrich
(Milwaukee, WI). Three independent evaluations were made for each of the 12
replicates.
Trichome density was evaluated on three leaflets from each of the 12 replicates
for a total of 36 leaflets/treatment. Each leaflet was individually collected and
stained for 3 min in fast green dye (Johansen, 1940). These leaflets were collected
at the same period from the plants at the ages studied. The trichome density
(trichomes/mm2) was calculated using a light microscope and by counting the
number of trichomes present per leaflet area in both surfaces (Channarayappa
et al., 1992). Twenty-four fields of 0.6 mm2 were analyzed in the mid-part of each
leaflet area between the main vein and the leaflet edge. Leaflet area of three leaves
collected in the morning was determined using a leaf area meter LI-COR model
LI-3000 (Lincoln, NE). Three independent evaluations were made for each of the
12 replicates.
The trichomes were classified as glandular or non-glandular in both genotypes for the purpose of this work, despite the existence of seven distinct trichome
types within the genus Lycopersicon (Channarayappa et al., 1992). This was
done because the accessions of L. hirsutum show almost only glandular trichomes
(types I, IV, VIc, and VII), mainly of the type VI (producer of tridecan-2-one and
undecan-2-one), unlike L. esculentum which shows mainly non-glandular
trichomes (types III, Va, and Vb) (Channarayappa et al., 1992). L. hirsutum
does contain other allelochemicals important in insect plant interactions such as
a-tomatine, a-humulene, chlorogenic acid and rutin, but their concentrations in
the leaves are rather small (Elliger et al., 1981). The main leaf allelochemicals of
L. hirsutum are tridecan-2-one and undecan-2-one which are present mainly on
type VI glandular trichomes and have been correlated to insect pest resistance
(Dimock et al., 1982; Dimock and Kennedy, 1983; Kennedy and Sorenson, 1985;
Lin et al., 1987; Giustolin, 1991).
The results on T. absoluta biology and attack, and tomato leaf size, trichome
density, and concentrations of tridecan-2-one and undecan-2-one were subjected
to two-way analysis of variance and the Scott–Knott multiple range test
(P < 0:05) whenever appropriate (Scott and Knott, 1974). Regression analysis
were carried out separately for L. hirsutum and L. esculentum correlating
trichome density and concentrations of tridecan-2-one, undecan-2-one with plant
age and biological characteristics of T. absoluta.

3. Results
There was no significant difference (P < 0:05) of undecan-2-one concentration
(0.0015% based on fresh weight) in L. hirsutum as a function of plant age.
However, trichome density (mainly of type VI glandular trichomes) and,

G.L.D. Leite et al. / Scientia Horticulturae 89 (2001) 103–113

107

Fig. 1. Effect of plant age on trichome density for L. esculentum and L. hirsutum (A), and
relationship between level of tridecan-2-one (% weight fresh) and trichome density for L. hirsutum
(B). The symbols represent the average of 12 replicates and the vertical bars indicate standard errors
of the mean.

consequently, production of tridecan-2-one increased with plant age in L.
hirsutum (Fig. 1). Trichome density (mainly of non-glandular trichomes) also
increased with plant age in L. esculentum (Fig. 1), but there was no effect of plant
age on leaf area in any of the tomato species studied.
The highest rates of egg eclosion (56.5 and 61.3%), lowest larval mortality
(72.8 and 77.6%) and shortest larval (22.9 and 25.8 days) and pupal (7.3 and 8.0

108

Plant age

Egg incubation period

Egg eclosion

Proportion of females

L. esculentum
ðN ¼ 488; n ¼ 84Þ

L. hirsutum
ðN ¼ 548; n ¼ 84Þ

L. esculentum
ðN ¼ 1260; n ¼ 84Þ

L. hirsutum
ðN ¼ 1260; n ¼ 84Þ

L. esculentum
ðN ¼ 133; n ¼ 44Þ

L. hirsutum
ðN ¼ 123; n ¼ 40Þ

2 Months
3 Months
4 Months

6.4 aAa
6.5 aA
6.4 aA

6.5 aA
6.4 aA
6.5 aA

61.8 aA
63.0 aA
59.0 aA

52.3 bB
61.8 aA
55.5 aB

0.8 aA
0.8 aA
0.4 aA

0.4 aB
0.4 bB
0.8 aA

Mean

6.4 a

6.4 a

61.3 a

56.5 b

0.7 a

0.6 b

a

Means followed by the same low case letter in a row or the same capital letter in a column do not differ significantly by the Scott–Knott multiple
range test ðP < 0:05Þ.

G.L.D. Leite et al. / Scientia Horticulturae 89 (2001) 103–113

Table 1
Egg incubation period (days), egg eclosion (%) and proportion of females ðNo: females=ðNo: females þ malesÞÞ of T. absoluta in leaves of L.
esculentum and L. hirsutum of different ages (N indicates the number of insects and n indicates the number of experimental units used for each plant
species in each analysis)

G.L.D. Leite et al. / Scientia Horticulturae 89 (2001) 103–113

109

Table 2
Length of larval and pupal periods (days), mortality of larvae and pupae (%), and number of small
(<0.5 cm) and large (>0.5 cm) mines of T. absoluta reared on leaves of L. esculentum and L.
hirsutum from plants of different ages (N indicates the number of insects and n indicates the number
of experimental units used for each plant species in each analysis)
Plant age

2 Months
3 Months
4 Months
Mean

2 Months
3 Months
4 Months
Mean

2 Months
3 Months
4 Months
Mean

Length of larval period (days)

Length of pupal period (days)

L. esculentum
ðN ¼ 133; n ¼ 44Þ

L. hirsutum
ðN ¼ 123; n ¼ 40Þ

L. esculentum
ðN ¼ 103; n ¼ 33Þ

L. hirsutum
ðN ¼ 92; n ¼ 30Þ

22.5 bAa
23.3 bA
23.5 bA
22.9 b
Larval mortality (%)

23.1
26.3
32.9
25.8

7.1 bA
6.9 bA
7.8 bA
7.3 b
Pupal mortality (%)

8.3
8.5
8.5
8.0

L. esculentum
ðN ¼ 488; n ¼ 84Þ

L. hirsutum
ðN ¼ 548; n ¼ 84Þ

L. esculentum
ðN ¼ 133; n ¼ 44Þ

L. hirsutum
ðN ¼ 123; n ¼ 40Þ

63.5 bB
69.8
68.5 aB
66.5
86.3 bA
96.4
72.8 b
77.6
No. of small mines/leaf

aC
aB
aA
a

aB
aB
aA
a

aA
aA
aA
a

35.2 aA
18.6
21.3 aA
36.8
7.4 aA
11.5
22.8 a
25.3
No. of large mines/leaf

aA
aA
aA
a

L. esculentum
ðn ¼ 84Þ

L. hirsutum
ðn ¼ 84Þ

L. esculentum
ðn ¼ 84Þ

L. hirsutum
ðn ¼ 84Þ

3.4
2.1
3.0
2.8

5.1
8.7
8.1
7.3

11.9 bA
8.8 bB
4.2 aC
8.3 b

15.2 aA
16.1 aA
2.8 bB
11.3 a

aA
bA
bA
b

aB
aA
aA
a

a

Means followed by the same low case letter in a row or the same capital letter in a column do
not differ significantly by the Scott–Knott multiple range test ðP < 0:05Þ.

days) periods, number of small mines (2.8 and 25.8) and large mines (8.3 and
11.3) per leaf, and higher proportions of females were observed in L. esculentum
in comparison with L. hirsutum (Tables 1 and 2). However, no negative effects of
L. hirsutum were observed at different ages on the length of the egg incubation
period (6.4 days), pupal mortality (25.3%) and weight (3.1 mg), as well as pupal
(6.5%) and adult malformation (0.0%) of T. absoluta (Tables 1 and 2).
The ageing of L. hirsutum plants led to higher mortality of T. absoluta larvae,
longer larval periods, higher number of small mines and lower number of large
mines, besides higher proportion of females (Tables 1 and 2). In L. esculentum,
ageing apparently led also to a higher larval mortality and smaller number of
large mines (Table 2). The length of the pupal period was similar for females and
males of T. absoluta in both tomato species (7.2 and 7.3 days, respectively, when
on L. esculentum, and 7.4 and 8.4 days, respectively, when on L. hirsutum).

110

G.L.D. Leite et al. / Scientia Horticulturae 89 (2001) 103–113

4. Discussion
Dimock et al. (1982) reported a higher level of undecan-2-one (0.066 % on
fresh weight) in L. hirsutum f. glabratum (PI 134417) than that observed in this
study, probably because of the winter effect on our experiment (Nichoul, 1994).
The shorter photoperiod of winter days reduces the number of type VI trichomes
which are involved in undecan-2-one production (Nichoul, 1994). In addition,
type VI trichome density and exudate production are higher during the spring
than during the fall due to differences in photoperiod between these seasons
(Nichoul, 1994). On the other hand, the level of tridecan-2-one observed here
(0.30% on fresh weight) was similar to the level reported by Dimock et al. (1982)
(0.37 % on fresh weight).
A low number of small mines probably reflects the host suitability to the
caterpillars. Thus, when adequate food is available, the caterpillars lodge in the
leaf mesophyll and consume it increasing the size of the mines and transforming
the small into large galleries. L. hirsutum does not seem to be a suitable host to the
leafminer probably due to the antixenotic and antibiotic effect of tridecan-2-one,
as observed on the Colorado potato beetle by Kennedy and Sorenson (1985), in
addition to the possible existence of deterrent substances in the leaf mesophyll of
this tomato species (Lin and Trumble, 1986). The larvae of this leafminer insect
pest usually leave their mines to move over the leaf surface during sunny hours
(Moore, 1983). Such behavior favors the exposure of the insects to the noxious
substances of the leaf glandular trichomes of L. hirsutum. In a free choice test
performed by Picanc¸o et al. (1995), no significant difference in the number of
leaf galleries formed by T. absoluta was observed between L. esculentum and
L. hirsutum. However, Leite et al. (1995) showed that despite the lower
oviposition of T. absoluta in L. hirsutum, the larvae made higher number of
galleries in this genotype than in L. esculentum.
There was no effect of undecan-2-one on T. absoluta unlike what was observed
by Giustolin (1991). Giustolin (1991) observed that with the addition of 0.03% of
undecan-2-one in the insect diet, there was a decrease in larval mortality and
length of the larval period, as well as an increase in pupal weight and adult
longevity. When 0.06% of undecan-2-one was added to the diet, only 8.6% of
T. absoluta caterpillars reached the next developmental stage, but no other effect
on the insect biology was observed. In our study the amount of undecan-2-one
observed was lower (ca. 40 ) than the amount necessary to cause harmful effects
on the leafminer, therefore it is not surprising the absence of undecan-2-one
effects in our experiment. In addition, it seems that this compound is less toxic
than tridecan-2-one and active in fewer insect species (Farrar and Kennedy,
1991).
The increase in trichome density with plant age is related to an increase in the
leaf levels of tridecan-2-one in L. hirsutum which is expected since this

G.L.D. Leite et al. / Scientia Horticulturae 89 (2001) 103–113

111

Fig. 2. Length of the larval period of T. absoluta as a function of the level of tridecan-2-one (% of
weight fresh) (A), and trichome density in leaves of L. hirsutum (B).

compound is present in leaf glandular trichomes (Fig. 1). The higher levels of
tridecan-2-one associated with older plants of L. hirsutum, are related to a slower
larval development of T. absoluta (Fig. 2) when compared with insects reared on
L. esculentum. These results suggest that commercial varieties of tomato having
L. hirsutum as source of resistance may be more resistant to the leafminer
especially during the plant reproductive stage, the critical period of attack by this
pest in tomato plants (Ulle´ and Nakano, 1994).
The higher male (than female) mortality in the larval stage (evaluation
performed after pupation) in L. hirsutum than in L. esculentum suggests that male

112

G.L.D. Leite et al. / Scientia Horticulturae 89 (2001) 103–113

larvae are more susceptible to tridecan-2-one than female larvae. This may be due
to an increased concentration of tridecan-2-one/mg weight in male larvae as a
consequence of their smaller weight and the higher relative penetration rate of the
allelochemical through the male cuticle since the males have lower body volume
and consequently higher specific area than the females.
The shorter pupal development period of females of T. absoluta and the higher
proportion of females in the population suggests an adaptative advantage of the
insect species decreasing the period of the female exposure to adverse conditions
(Coelho and Franc¸a, 1987). Giustolin and Vendramim (1994) reported lower
pupal weights (of both males and females) for T. absoluta reared on L. hirsutum
than on L. esculentum. This could be due to antibiosis of tridecan-2-one on T.
absoluta caterpillars when feeding in L. hirsutum since the type VI glandular
trichomes which produce this compound are not present in L. esculentum.

5. Conclusion
T. absoluta showed better development in L. esculentum than in L. hirsutum.
The increase in trichome density with plant age led to an increase in the leaf
levels of tridecan-2-one in L. hirsutum and these higher levels of tridecan-2-one
may have led to a slower larval development of T. absoluta when compared with
insects reared on L. esculentum. These results suggest that commercial varieties
of tomato having L. hirsutum as source of resistance may be more resistant to the
leafminer especially in their reproductive stage, the critical period of attack by
this pest in tomato plants.

Acknowledgements
Appreciation is expressed to Dr. Patrick De Clerk, from the University of Gent,
Belgium, and two anonymous referees for reviewing this manuscript and
providing helpful comments and the Brazilian Government (CNPq) for financial
support.

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