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3 FiguresEffect of Rootstock on Mango Fruit Susceptibility to Infestation by Anastrepha obliqua
Abstract
The effect of the use of rootstock Criollo on the susceptibility of Manila mango fruit to infestation by Anastrepha obliqua (Macquart) was determined in the present investigation. Growth, quality parameters (Soluble solids content (%), citric acid content (%), firmness, monoterpene volatiles (3-carene, limonene, alpha-pinene, and beta-myrcene), main flavonoids content (milligrams/100 g fresh pulp), and larvicidal activity of methanolic extracts, as well as the degree of infestation during preharvest development of the fruits were analyzed. The results indicated that the rootstock did not have any significant effect on growth, soluble solids content, or citric acid content; although it increased firmness as well as 3-carene levels and main flavonoids content, resulting in a greater resistance to infestation by A. obliqua. The obtained results support the use of this rootstock because of the beneficial effects observed on the resistance of mango fruit to infestation by this fruit fly.
Figures
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Effect of Rootstock on Mango Fruit Susceptibility to Infestation by
Anastrepha obliqua
Author(s) :A. Vazquez-Luna, F. Rivera-Cabrera, L. J. Perez-Flores, and R. Diaz-
Sobac
Source: Journal of Economic Entomology, 104(6):1991-1998. 2011.
Published By: Entomological Society of America
DOI:
URL: http://www.bioone.org/doi/full/10.1603/EC11223
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HORTICULTURAL ENTOMOLOGY
Effect of Rootstock on Mango Fruit Susceptibility to Infestation by
Anastrepha obliqua
A. VAZQUEZ-LUNA,
1
F. RIVERA-CABRERA,
2
L. J. PEREZ-FLORES,
2,3
AND R. DIAZ-SOBAC
4
J. Econ. Entomol. 104(6): 1991Ð1998 (2011); DOI: http://dx.doi.org/10.1603/EC11223
ABSTRACT The effect of the use of rootstock Criollo on the susceptibility of Manila mango fruit
to infestation by Anastrepha obliqua (Macquart) was determined in the present investigation.
Growth, quality parameters (Soluble solids content (%), citric acid content (%), Þrmness,
monoterpene volatiles (3-carene, limonene,
␣
-pinene, and

-myrcene), main ßavonoids content
(milligrams/100 g fresh pulp), and larvicidal activity of methanolic extracts, as well as the degree
of infestation during preharvest development of the fruits were analyzed. The results indicated
that the rootstock did not have any signiÞcant effect on growth, soluble solids content, or citric
acid content; although it increased Þrmness as well as 3-carene levels and main ßavonoids content,
resulting in a greater resistance to infestation by A. obliqua. The obtained results support the use
of this rootstock because of the beneÞcial effects observed on the resistance of mango fruit to
infestation by this fruit ßy.
KEY WORDS biomolecules, fruit ßy, graft, mango, preharvest
Mango (Mangifera indica L.) is one of the fruits with
greater commercial importance worldwide. In Vera-
cruz, Manila, and Criollo mango fruits are commer-
cialized and some producers use the Criollo mango as
rootstock for the Manila variety (SAGARPA 2010).
Even though Manila mango fruit is appreciated for its
organoleptic characteristics, it is aimed principally for
the national market and it is not considered for ex-
portation. This variety is very susceptible to infesta-
tion by fruit ßies of the genus Anastrepha.
Quarantine treatments are used to control the
fruit ßy on the varieties for exportation, whereas on
Manila mangoes, the control is based principally on
the setting of McPhail type traps. This method of
biological control is based upon knowledge of the
reproduction and life cycles of the insect, as well as
on its capability to perceive volatiles that have an
attracting effect causing its physical immobilization
(Aluja and Pin˜ero 2004, Pelz-Stelinski et al. 2006).
It has been reported that several fruits and plants
synthesize monoterpenes, aldehydes, and ßa-
vonoids, among other biomolecules, during certain
stages of their development. Some of these com-
pounds may be attractive or repellent to the ßies,
whereas others work as antinutrients or larvicides
regulating the development of the different types of
ßy larvae (Morimoto et al. 2003, Malo et al. 2005,
Va´zquez-Luna et al. 2007, Massa et al. 2008).
The attracting effect of some fruit volatiles such
as ethyl butyrate, 3-hexen-1-ol, 1-hexanol, ethyl
hexanoate, hexyl acetate, 3-hexenyl butyrate ethyl
octanoate, ethyl propionate, ethyl decanoate, ethyl dode-
canoate, D-limonene, benzoic acid, and dimethyl anthra-
nilate upon the ßies of the Anastrepha genus has been
studied by different authors (Malo et al. 2005, Massa et al.
2008). Herna´ndez-Sa´nchez et al. (2001) analyzed the
volatiles present in Tommy Atkins mango fruits, Þnding
that the terpenes with the greatest attracting effect on
Ceratitis capitata (Wiedemann) were p-cimene and li-
monene.
Araujo et al. (2003) reported that the monoterpenes
␦
-3-carene and 1,8-cineole from the essential oil of
Hyptis martiusii Benth presented insecticidal activity
against the eggs and insects of the white ßy (Bemisia
argentifolii [Bellows and Perring]), a common pest
affecting edible fruits like cantaloupe and water-
melon. Ibrahim et al. (2001) reported that the mono-
terpenes limonene,
␣
-pinene, and

-myrcene, of citric
fruits, showed an insecticidal activity against domestic
ßies.
However, it has been reported that ßavonoids
5-hydroxy-3,6,7,8,4⬘- pentamethoxyßavone, 5-hy-
droxy-3,6,7,8-tetramethoxyßavone, 5,6-dihydroxy-
3,7-dimethoxyßavone and 4,4⬘,6⬘-trihydroxy-2⬘-me-
thoxy-chalcone, present in extracts of Gnaphalium
affine D. Don., showed antialimentary action against
the larvae of Spodoptera litura F. in their Þrst stage
of development, which attacks fruits such as canta-
loupe (Morimoto et al. 2000). Likewise, Morimoto
1
Posgrado en Biologõ´a Experimental, Universidad Auto´noma Met-
ropolitana-Iztapalapa. Av. San Rafael Atlixco No. 186, Col. Vicentina,
Delegacio´n Iztapalapa, C.P. 09340 Me´xico, D.F.
2
Departamento de Ciencias de la Salud, Universidad Auto´noma
Metropolitana-Iztapalapa. Av. San Rafael Atlixco No. 186, Col. Vi-
centina, Delegacio´n Iztapalapa, C.P. 09340 Me´xico, D.F.
3
Corresponding author, e-mail: ljpf@xanum.uam.mx.
4
Instituto de Ciencias Ba´sicas, Laboratory de Quõ´mica y Biologõ´a
Molecular de Frutas. Universidad Veracruzana. Av. Luis Castelazo
s/n, Col. Industrial Animas, C.P. 91190.
0022-0493/11/1991Ð1998$04.00/0 䉷2011 Entomological Society of America
et al. (2003) isolated ßavones from Sculletaria ba-
icalensis Georgi root, observing that nobiletin, chry-
sin, and wogonin had the highest antialimentary
activity against larvae of S. litura.
In addition to the biomolecules that have different
effects on adult ßies and the larvae development,
changes in chemical composition (reduction in acid-
ity) as well as changes in physical structure (decrease
in Þrmness) that occur during fruit development also
play an important role on the success of the infesta-
tion. On this regard, Pen˜a and Mohyuddin (1997)
revised the resistance to Anastrepha obliqua (Mac-
quart) and suggested that the difference in the sus-
ceptibility to the infestation may be because of the
presence of toxic compounds, the absence of essential
nutrients or resin ducts which give resistance to the
fruit against both, the ovipositor and the movement of
the larvae. Hennessey and Schnell (2001) analyzed 18
mango cultivars to determine their resistance to the
Caribbean ßies larvae (Anastrepha suspensa (Loew))
development, according to the developmental stage of
the fruit. In this research, slices of ripe and unripe fruit
were obtained and inoculated with ßy eggs. Despite a
higher resistance to infestation displayed by some
cultivars, in all cases, ripe mangoes were better sub-
strates for the larvae development than unripe fruit,
suggesting that there is a critical step of the oviposited
egg and its transition to the larva stage, which depends,
among other factors, on the composition of the fruit
pulp and nutrients availability. In addition, it has been
observed that in the earlier stages of development, C.
capitata larvae grows better in diets with a high con-
tent of glucose and sucrose compared with those that
contain starch and maltose (Zucoloto 1993). Addi-
tionally, it was found that when C. capitata larvae are
put in a piece of fruit with a lower nutritive content,
they prefer to move to another with a higher nutri-
tional quality which presents higher quantities of sol-
utes and sugars (Fernandes-da-Silva and Zucoloto
1993). The objective of the current study was to an-
alyze the effect of the use of Criollo rootstock in the
susceptibility of Manila mango fruit to infestation by
A. obliqua and to correlate it with the presence of
biomolecules in the fruit and some characteristics of
the pulp composition during the preharvest develop-
ment.
Materials and Methods
Biological Material. The study was carried out in
mango orchards in Jalcomulco, Veracruz (Mexico),
located at 96⬚55⬘43.4⬙west longitude, with an annual
average temperature of 24⬚C and an annual average
rainfall of 1,100 mm. The mean maximum and mini-
mum temperature was 31.1 and 17.7⬚C, respectively
(INIFAP 2010). Eight years old mango trees, Manila,
Criollo, and Manila grafted in Criollo were selected by
using an aleatory-number criterion. The study was
begun in March, during the Þrst seasonal mango ßow-
ering. Fruit samples were obtained every 7 d, from 37
to 100 d after ßowering. The experimental unit con-
sisted of three replicates of three fruits for each day of
sampling and for every type of mango studied. In the
case of fruit ßy infestation the experimental unit was
of nine independent fruits with three replicates for
each day of sampling and for every type of mango
studied. Growth rate was determined through the
estimation of the fruit volume from the measuring of
the diameters with a Vernier according to the equation
established by Castro-Neto and Reinhardt (2003):
Volume ⫽longitudinal diameter ⫻ventral diameter ⫻
dorsal diameter.
Firmness. It was determined through a universal
texturometer (model 18-2321 Sommer & Runge)
measuring the force to penetrate the fruit. The re-
sults obtained were expressed in Newtons using the
following equation: Firmness (N) ⫽Kg-force ⫻
9.807. Firmness on mango fruit was measured on
both sides of the equatorial diameter because that is
the part with the most quantity of pulp, which re-
ßects adequately the Þrmness of the fruit (Sugiyama
et al. 2005).
Soluble Solids Content (SSC). The juice from 5 g of
pulp was extracted by hand maceration and SSC was
determined with a digital manual refractometer
(ATAGO, Mod. Pal-1). Results were expressed in per-
centage.
Citric Acid Content (%). Percentage of acidity was
reported as the percentage of citric acid, the predom-
inant acid in mango, using the following equation:
citric acid content (%) ⫽[0.064 ⫻vol of NaOH used ⫻
concentration of NaOH solution (0.1 mol/L)]/wt of
sample (g).
Volatiles. 3-carene, limonene,
␣
-pinene, and

-myrcene.
To obtain the volatiles, an entire fruit of known weight
from each type of mango at the different preharvest
development stages was placed in a 900 ml glass con-
tainer with an airtight cap, to which a rubber septum
was attached for gas sampling, after 2 h, 5 ml of the
mixture of air-volatiles were extracted and injected
into a HewlettÐPackard HP6890 Gas Chromatogra-
pher coupled to a quadrupole mass detector Agilent
Technologies 5973. A packed column HP5MS of 5%-
phenyl-95% methylpolysiloxane, 30 m long ⫻0.25 mm
in diameter with a 25
m thickness of stationary phase
was used to separate and detect the volatiles. Helium
was used as the carrier gas with a ßow rate of 1 ml/min.
The initial oven temperature was 50⬚C that was kept
constant for 4 min, and a ramp of 25⬚C/min was used
until reaching 250⬚C; the Þnal temperature was kept
constant during 4 min, the total time of the operation
was 16 min. The temperature of the injector and of the
electronic ionization detector (EID) was 250⬚C (An-
drade et al. 2000, Pino et al. 2005). The structural
information generated from the mass spectrum was
identiÞed using the library of compounds of the
ChemStation program with a 90% of reliability. Pure
standards were used to quantify
␣
-pinene,

-myrcene,
3-carene, and limonene (Sigma-Aldrich, St. Louis,
MO).
Methanolic Extracts. Ten grams of pulp from the
mangoes studied were used. To remove chlorophyll
which interferes with ßavonoid detection, the pulp
was extracted in 100 ml of hexane at room temperature
1992 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 104, no. 6
for 48 h. A second extraction of the pulp was carried
out with 100 ml of methanol for 96 h at room tem-
perature. After this time, the methanolic extract was
separated by centrifugation at 4500 g, during 5 min at
30⬚C and afterwards it was Þltered and kept in dark-
ness at 10⬚C.
Main Flavonoids Content in Methanolic Extracts.
The quantiÞcation of mangiferin, quercetin, and cate-
quin in the methanolic extracts was performed using
the method described by Merken and Beecher (2000)
with a high performance liquid chromatography (Var-
ian model ProStar 210) equipped with a 4.6 ⫻250 mm
5
m C18 reversed phase column (Agilent, Wilming-
ton, DE) and an ultra violet detector. The mobile
phase consisted of 2% (vol:vol) acetic acid in water
(eluent A) and 2% (vol:vol) of acetic acid in MeOH
(eluent B). The gradient program was as follows: 40Ð
60% B (5 min), 30Ð70% B (20 min), 15Ð85% B (35
min). The injection volume for all samples was 20
l.
Authentic chemical standards (Sigma-Aldrich) were
used to conÞrm and quantify these ßavonoids. The
sum of mangiferin, quercetin, and catequin was ⬎60%
of the ßavonoids extracted from the mangoes studied.
Results were expressed as milligrams of main ßa-
vonoids (sum of mangiferin, quercetin, and cate-
quin)/100 g fresh pulp.
Fruit Fly Infestation (A. obliqua). Fruit ßy infes-
tation during the preharvest development of the man-
goes studied was determined by evaluating the pres-
ence of larvae in the fruit through the visual inspection
method described by Pelz-Stelinski et al. (2006). In-
festation was reported as the number of larvae/kilo-
gram of fruit (Manrhakan and Price 1999, Pelz-Stel-
inski et al. 2006).
Larvicidal Activity of Methanolic Extracts. Metha-
nolic extracts of pulp from the three types of mangoes,
obtained during the preharvest development, were
used to determine larvicidal activity according to the
method described by Albuquerque et al. (2004). Dif-
ferent volumes of methanolic extracts (50 and 100
l)
were taken and adjusted to a Þnal volume of 20 ml with
a mixture of water (98.5 ml): dimethylsulfoxide (1.5
ml). Twenty Þve larvae of A. obliqua from the Þrst
stage were placed in petri dishes containing the 20 ml
of the methanolic extracts solutions at each of the
concentrations mentioned above. After 24 h at room
temperature, the number of dead larvae was counted
and the percentage of mortality for each of the con-
centrations was calculated. Methanol-water dimethyl
sulfoxide solutions in the same proportion as the sam-
ples were analyzed at the same time and used as
controls.
Statistical Analysis. The results from the physical
and chemical characteristics as well as the fruit ßy
infestations during the preharvest development and
the larvicidal activity of the methanolic extracts were
analyzed using analysis of variance (ANOVA) (
␣
⫽
0.05; Minitab release 12.2). When the main effects
were signiÞcant (P⬍0.05) differences between
means were separated using Tukey multiple compar-
ison test.
Results
The average growth rate between the 72nd and the
100th days for Manila mango was 4.44 ⫾0.28 cm
3
/d
and 4.64 ⫾0.32 cm
3
/d for the Manila grafted in Criollo
rootstock, with no signiÞcant differences observed
between them; whereas mango Criollo presented an
average growth rate signiÞcantly lower (2.64 ⫾0.19
cm
3
/d) (F⫽6.7; df ⫽2; P⫽0.016) (data not shown).
Firmness was determined as the force required to
penetrate the fruit expressed in Newtons (Fig. 1a). A
gradual decrease was observed in the Þrmness of the
fruits during the preharvest development, although
the softening was greater between the 72nd and 100th
days. In general, Manila mango presented a signiÞ-
cantly greater softening (F⫽3.98; df ⫽2; P⫽0.023)
than Criollo mango, which showed a greater Þrmness;
whereas Manila mango grafted in Criollo showed in-
termediate values. The use of the rootstock affected
Fig. 1. (a) Firmness, (b) SSC (%), and (c) citric acid
content (%) during the preharvest development of Manila,
Criollo, and Manila grafted in Criollo (graft) mango fruits.
Each value was obtained from the mean of three independent
samples ⫾SE.
December 2011 VA´ZQUEZ-LUNA ET AL.: MANGO FRUIT SUSCEPTIBILITY TO A. obliqua 1993
the Þrmness of the fruit because values higher than
those of the Manila mango were found.
The percentage of SSC increases considerably dur-
ing the development of mango fruit (Fig. 1b). In gen-
eral, the SSC observed in Criollo mango were signif-
icantly lower (F⫽4.13; df ⫽2; P⫽0.016) than those
from the other types of mango during the Þrst 79 d of
the preharvest development. No signiÞcant differ-
ences were found in the levels of SSC between Manila
mango and Manila grafted in Criollo in this period. In
following periods, the levels of SSC were similar
among the different types of mango studied. At 100 d
of preharvest development (considered as harvestable
maturity for Manila), the fruits reached a maximum
value of 9.57 ⫾0.11 for Manila, 9.05 ⫾0.15 for Criollo,
and 10.03 ⫾0.07% for Manila grafted in Criollo.
The citric acid content (%) showed a gradual de-
crease during the preharvest development in the three
types of mango (Fig. 1c). The highest values of citric
acid (F⫽5.87; df ⫽2; P⫽0.009) were found in Criollo
and the lowest were observed in Manila and Manila
grafted in Criollo, with no signiÞcant differences be-
tween these types of mango. Citric acid content in the
physiological or harvestable maturity (100 d) for Ma-
nila, Criollo, and Manila in Criollo rootstock was
0.99 ⫾0.04, 1.52 ⫾0.13, and 1.18 ⫾0.07%, respectively.
The most abundant monoterpene found in the stud-
ied mangoes was 3-carene, followed by limonene,
␣
-pinene, and

-myrcene (Fig. 2). In general, a grad-
ual decrease in the four volatiles was observed during
the Þrst 72 d of development in the analyzed mangoes.
Later, a more drastic decrease in the levels of these
volatiles was observed until undetectable values were
reached at 100 d of development. In the case of
3-carene, the highest levels were detected in Criollo
and the lowest in Manila; whereas Manila in Criollo
rootstock showed intermediate values signiÞcantly su-
perior to those in Manila (F⫽8.95; df ⫽2; P⫽0.001).
Regarding the levels of limonene and
␣
-pinene, the
highest levels were found in Criollo, with no signiÞ-
cant differences observed between Manila and Manila
grafted in Criollo. No signiÞcant differences were ob-
served in the levels of

-myrcene among the mangoes
analyzed during the whole period of preharvest de-
velopment.
A behavior similar to that of the volatiles studied
was observed for the levels of the main ßavonoids (Fig.
3), this is, the main ßavonoids content decreased dur-
ing the preharvest development. Between 37 and 72 d
of development, there was a gradual decrease in the
main ßavonoids content for the three types of mango;
from this stage on, the decrease was more pronounced
reaching values close to 5.3 mg/100 g of fresh pulp
weight at 100 d of development. The main ßavonoids
content was signiÞcantly lower in Manila during the
entire development, whereas Criollo presented the
highest values and Manila in Criollo rootstock pre-
sented intermediate values (F⫽6.52; df ⫽2; P⫽
0.005).
The evaluation of the degree of infestation was
measured through the quantiÞcation of the number of
larvae present in the pulp of the sampled mangoes.
The results are shown in Table 1. No larvae were found
in any of the mangoes analyzed between 37 and 65 d
of development. The number of larvae present was
signiÞcantly different (F⫽63.23, df ⫽2, P⫽0.001) for
Fig. 2. Levels of 3-carene, limonene,
␣
-pinene, and

-myrcene obtained during the preharvest development of Manila,
Criollo, and Manila grafted in Criollo (graft) mango fruits. Each value was obtained from the mean of three independent
samples ⫾SE.
1994 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 104, no. 6
the three types of mango studied, after 72 d of devel-
opment. The larvae were detected in earlier stages of
development (72 d) and in higher number for Manila
mango (F⫽15.91, df ⫽2, P⫽0.001. In Manila grafted
in Criollo, larvae were observed starting from 79 d of
development, while Criollo mango showed the great-
est resistance to infestation because the larvae were
detected at 86 d of development and its number was
signiÞcantly lower from 86 to 100 d of preharvest
development (F⫽50.21, df ⫽2, P⫽0.001).
The results of the percentage of larvicidal activity in
the methanolic extracts obtained from the pulp of
Manila, Criollo, and Manila grafted in Criollo mango
fruit at different days of the preharvest development
are shown in Fig. 4. In general, it was observed that the
larvicidal activity was greater during the Þrst days of
development even in the mixture with lower metha-
nolic extract concentration and that the extracts with
the greatest activity were those from Criollo mango.
Furthermore, it was observed that the greater the
content of methanolic extracts, the greater the larvi-
cidal activity (Fig. 4b). It is interesting to emphasize
that the extracts obtained from fruits from the grafted
mango showed a signiÞcantly higher activity that
those observed in Manila mango (F⫽68.94, df ⫽2, P⫽
0.001) and (F⫽95.68, df ⫽2, P⫽0.001) for 50 and 100
l of methanolic extracts, respectively.
Discussion
Important changes in chemical composition and
physical structure occur during the preharvest devel-
opment of mango fruit. For its economic relevance the
increase in volume as an indicator of fruit develop-
ment is an important parameter (Guzma´n-Estrada et
al. 1996, Pereira de Lucena et al. 2007). Our results
indicate that the use of Criollo as rootstock for Manila
did not signiÞcantly affect the growth of the fruits
because these reached volumes similar to those ob-
tained by Manila (data not shown).
Kader (2008) reported that the Þrmness in Haden
and Kent mangoes decreases during their physiolog-
ical development on the tree and the biggest fruits
have less Þrmness and ripen faster than the smaller
Fig. 3. Main ßavonoids content in methanolic extracts of
fresh pulp of Manila, Criollo, and Manila grafted in Criollo
(graft) mango fruits during the preharvest development
were expressed as the sum of mangiferin, quercetin, and
catequin. Each value was obtained from the mean of three
independent samples ⫾SE.
Table 1. Analysis of fruit fly infestation (A. obliqua) during preharvest development of Manila, Criollo, and Manila grafted in Criollo
(graft) mango fruits
Fruit Day of preharvest development
37 44 51 58 65 72 79 86 93 100
Manila 000002⫾0.08 6 ⫾0.79 21 ⫾1.59 40 ⫾2.26 26 ⫾1.94
Criollo 00000 0 0 3⫾0.09 8 ⫾0.87 5 ⫾0.28
Graft 00000 0 1⫾0.04 7 ⫾0.75 13 ⫾1.18 9 ⫾0.92
Data are the mean of three samples ⫾SE.
Fig. 4. Larvicidal activity (% of mortality) of the metha-
nolic extracts from the pulp of Manila, Criollo, and Manila
grafted in Criollo (graft) mango fruits during the preharvest
development; (a) 50
l of methanolic extracts and (b) 100
l
of methanolic extracts. Each value was obtained from the
mean of three independent samples ⫾SE.
December 2011 VA´ZQUEZ-LUNA ET AL.: MANGO FRUIT SUSCEPTIBILITY TO A. obliqua 1995
ones harvested from the same tree and at the same
developmental stage. In the present work, Criollo
mangoes were signiÞcantly smaller and have less Þrm-
ness than the other mangoes studied, corroborating
the proposal by Kader. However, the fruits of mango
Manila grafted in Criollo rootstock, despite having
volumes similar to those of Manila, presented higher
values of Þrmness. Petit-Jime´nez et al. (2007) propose
that the reduction in Þrmness is because of changes in
the microstructure, chemical composition, and thick-
ness of the skin during the preharvest development.
Several enzymes related to the degradation of pectins
and hemicelluloses participate in these changes, such
as pectin methylesterases, polygalacturonases,

-ga-
lactosidases, xyloglucanases, and xylanases (San˜udo et
al. 2008). To understand the causes of the greater
Þrmness in the fruits from Manila grafted in Criollo
rootstock, it is necessary to study the structure, chem-
ical composition, and thickness of the mango skin and
the activity of the enzymes participating in the fruit
softening.
The SSC values obtained in this work were similar
to those reported for Tommy Atkins, Haden, Keitt, and
Kent mangoes (Baez-San˜udo et al. 1999, Kader 2008,
Dick et al. 2009). The results obtained in this study
indicate that the use of the rootstock did not signiÞ-
cantly affect the SSC in the fruit.
The decrease in acidity has been attributed to a
reduction in the concentration of the acids which
participate in metabolic processes associated with the
preharvest maturation (Wills et al. 1989). The results
obtained from the mangoes studied in this work were
similar to those reported by Baez-San˜udo et al. (1999)
for Tommy Atkins mangoes. Using Tommy Atkins
mangoes, Dantas de Morais et al. (2002) determined
that the higher the acidity values, the greater the fruit
Þrmness. A positive correlation between citric acid
content and Þrmness in all studied mangoes (r⬎0.85)
was also found in this work (data not shown). Our
results indicate that the use of the rootstock did not
modify the citric acid content in Manila. As was pre-
viously mentioned, the use of the rootstock did not
affect the levels of SSC (%) either; therefore the
maintenance of the sensorial quality could be ex-
pected.
Among the most abundant volatiles during the pre-
harvest development of the three types of mangoes,
the monoterpenes 3-carene, limonene,
␣
-pinene, and

-myrcene were found. These monoterpenes have
also been identiÞed as being among the most abundant
in other mango cultivars such as: Tommy Atkins,
Ataulfo, Haden, Manga amarilla, Macho, Manga
blanca, San Diego, Manzano, Smith, Florida, Keitt,
Kent, Delicioso, Super Haden, Ordon˜ez, Filipino, La
Paz, and Minin (Bender et al. 2000, Franco et al. 2004,
Pino et al. 2005, Salazar-Sandoval et al. 2007). The
levels of these volatiles were analyzed in this study
because it has been reported that they have an at-
tracting or insecticidal activity against different spe-
cies of ßies such as: C. capitata, Musca domestica L., B.
argentifolii, and the larvae of Culex quinquefasciatus
Say mosquito and Aedes aegypti L. ßy (Herna´ndez-
Sa´nchez et al. 2001, Ibrahim et al. 2001, Araujo et al.
2003). Salazar-Sandoval et al. (2007) also observed in
Ataulfo mango that the major component of the vola-
tiles was the 3-carene, its concentration being greater
in unripe than in ripe fruits. The results obtained in
this study indicate that the use of the Criollo rootstock
signiÞcantly increased the levels of 3-carene in the
fruits, with no signiÞcant differences observed in the
levels of limonene,
␣
-pinene, and

-myrcene.
The results of this work indicate that the Criollo
rootstock increased the main ßavonoids content in the
fruit. Berardini et al. (2005) analyzed the content of
ßavonoids and xanthones in 14 varieties of ripe mango,
among them: Manila, Tommy Atkins, Haden, and
Kent, the highest concentrations being observed in
the skin of the fruit and not in the pulp. It was also
found in the current study that the levels of total
ßavonoids were higher in the skin than in the pulp
(data not shown).
The absence of larvae in the Þrst stages of devel-
opment (37Ð65 d) in the mangoes studied, coincides
with the report of Hennesey and Schnell (2001), who
observed that unripe fruits from 18 mango cultivars
were more resistant to the development of the Carib-
bean ßy (A. suspensa) (Loew) than ripe fruits. These
authors attribute these differences to the fact that ripe
mangoes contain a greater availability of nutrients for
the development of the larvae. Moreover, after 72 d of
development, a drastic decrease was observed in the
levels of volatiles and the main ßavonoids, which have
been reported to possess insecticidal, larvicidal, and
antialimentary properties, and this would explain the
presence of larvae in the three types of mango from
this stage of development onward.
The greater resistance to infestation observed in the
Criollo mango in this study (Table 1) can be explained
by several factors: these fruits showed the highest
values of Þrmness (Fig. 1a), the lowest levels of SSC
(Fig. 1b) and the highest acidity (Fig. 1c), which
would generate unfavorable conditions for oviposition
and egg development. Furthermore, they also had the
highest levels of the monoterpene volatiles 3-carene,
limonene, and
␣
-pinene (Fig. 2), which have been
reported to have insecticidal activity. Besides, its
methanolic extract showed the highest larvicidal ac-
tivity (Fig. 4). The differences observed in the infes-
tation between Manila grafted in Criollo and Manila
mangoes (Table 1) could be explained through the
differences observed in the levels of 3-carene that
were signiÞcantly higher in the grafted mango, be-
cause no signiÞcant differences were observed in the
quality parameters analyzed (Fig. 1). Another expla-
nation that requires further study is the presence of
other phytochemicals in the methanolic extracts that
could be responsible for the larvicidal activity ob-
served. In this sense it would be interesting to analyze
the composition of the methanolic extracts to identify
the compound(s) that possess larvicidal activity.
The results obtained in this study indicate that the
use of Criollo mango as rootstock for Manila did not
signiÞcantly affect the growth of the fruits because
these had volumes similar to Manila mango. It did not
1996 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 104, no. 6
affect the SSC levels either, or the content of citric
acid, because no signiÞcant differences were found in
these parameters. In general, the use of the rootstock
increased the Þrmness of the fruit. Additionally, it
signiÞcantly increased the levels of 3-carene and the
main ßavonoids content in the fruit, giving as a result
a greater resistance to infestation by A. obliqua. These
results favor the use of this rootstock because of the
beneÞcial effects observed in the resistance to infes-
tation by A. obliqua.
Acknowledgments
This work is part of Alma Va´zquez Luna Ph.D. disser-
tation. This research was funded by Universidad Vera
cruzana P/PIFI-2008-30MSU09408-21, Universidad Au-
to´noma Metropolitana, SEP-PROMEP (Grant UAM-I-CA-
26), BIOINNOVAL Co. for supplying the fruit. Alma
Va´zquez Luna holds a fellowship from CONACYT
(192870) during her studies in the Experimental Biology
Ph.D. Program.
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Received 11 July 2011; accepted 23 August 2011.
1998 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 104, no. 6
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