Crop Protection 28 (2009) 41–45
Contents lists available at ScienceDirect
Crop Protection
journal homepage: www.elsevier.com/locate/cropro
Sigatoka disease reduces the greenlife of bananas
M. Chillet a, *, C. Abadie b, O. Hubert c, Y. Chilin-Charles b, L. de Lapeyre de Bellaire d
a
CIRAD-Persyst, UMR Qualisud, Departamento de Alimentos e Nutriçao Experimental – Faculdade de Ciencias Farmaceuticas – Universidade de Sao Paulo,
Avenida Lineu Prestes, 580 – Bloco 14 – 05508-900 Sao Paulo, Brasil
b
CIRAD-Bios, UPR Multiplication végétative, Station de Neufchâteau, 97130, Capesterre-Belle-Eau, Guadeloupe (FWI), France
c
CIRAD-Persyst, UMR Qualisud, Station de Neufchâteau, 97130, Capesterre-Belle-Eau, Guadeloupe (FWI), France
d
CIRAD-Persyst, UR Systèmes de culture banane et ananas, CARBAP – BP 832 Douala, Cameroon
a r t i c l e i n f o
a b s t r a c t
Article history:
Received 30 May 2008
Received in revised form 12 August 2008
Accepted 15 August 2008
Sigatoka disease (SD) of bananas is caused by the pathogenic fungus Mycosphaerella musicola Leach. This
disease provokes necrotic lesions on leaves and serious infestations can lead to a substantial reduction in
the leaf area of infected plants and thus to yield losses. In addition to these effects on yield, SD was found
to have an impact on fruit quality, especially because exported bananas ripen prematurely. In the present
work, a plantation survey and experiments have been conducted in Guadeloupe (FWI) to assess the effect
of this disease on the greenlife of bananas harvested at a constant physiological age, as measured in
degree-days (dd). Our results revealed that bananas harvested at 900 dd from plants with high Sigatoka
disease severity had normal diameter growth, but a shorter greenlife (GL) than bananas harvested from
uninfected plants. These results indicate that SD is directly responsible for the reduction of banana
greenlife since the reduction of GL could not be attributed to the harvest of fruits at a more advanced
physiological age (dd). Furthermore, a correlation was noted between SD severity and GL. The potential
physiological mechanisms involved are also discussed.
Ó 2008 Elsevier Ltd. All rights reserved.
Keywords:
Banana
Sigatoka disease
Mycospharella musicola
Greenlife
Fruit quality
1. Introduction
Banana (Musa acuminata, triploid AAA, Cavendish cv Grande
Naine) is a climacteric fruit (Burg and Burg, 1965; Seymour, 1993;
Inaba et al., 2007) that is harvested at a green preclimacteric stage
in commercial plantations prior to being exported. It is essential
that the bananas reach their export market destination in this
physiological state. They must be marketed at a specific ripeness
stage to ensure a sufficient shelflife. Fruit ripening can be artificially induced by exogenous ethylene treatment in commercial
ripening sheds (Marriott, 1980). The time between harvest and
natural fruit ripening initiation via the synthesis of endogenous
ethylene (climacteric rise) is called the greenlife (GL) (Peacock
and Blake, 1970). The period between harvest and artificial
ripening initiation in a ripening shed is variable and depends on
the shipping time and the banana market liquidity. This period
ranges from around 15 d to 20 d for bananas grown in the West
Indies and marketed in Europe. Bananas must therefore have a GL
of around 25 d in order to be in a preclimacteric stage when they
* Corresponding author. Tel.: þ55 11 3091 1483; fax: þ55 11 3815 4410.
E-mail address: [email protected] (M. Chillet).
0261-2194/$ – see front matter Ó 2008 Elsevier Ltd. All rights reserved.
doi:10.1016/j.cropro.2008.08.008
reach the ripening rooms – otherwise they will ripen during
transport and thus not be marketable.
GL is highly temperature dependent (Blake and Peacock, 1971),
so bananas are quickly cooled to 13 C after harvest in order to
extend their conservation. GL is also closely related to the physiological age of the fruit, as calculated by the sum of mean daily
temperatures, according to a negative exponential model (Jullien,
2000). When fruit pulp filling is not hampered by any cropping
stress, bananas harvested at 900 dd (at 14 C, i.e. the zero growth
threshold for cv Grande Naine) reach a marketable diameter
(Ganry and Meyer, 1975) and a GL that is sufficient to withstand
maritime shipping (Jullien, 2000). It has, however, been shown
that bananas harvested from plants that have undergone stress
during the fruit pulp filling phase have lower diameter growth.
This is the case when there is a marked decrease in the number of
leaves at flowering (Turner and Hunt, 1987; Chillet et al., 2006;
Daniells et al., 1987), and also when fruit pulp filling occurs in the
presence of excess moisture (Chillet et al., 2006). In these conditions, it has also been shown that when bananas are harvested at
a constant physiological age (900 dd thermal sum), these cropping
stresses have no impact on GL (Chillet et al., 2006), but they do
have a very marked effect on the fruit diameter. Such bananas
have to be harvested after 900 dd so that they will have enough
time to reach the commercial grade, so they will be harvested at
42
M. Chillet et al. / Crop Protection 28 (2009) 41–45
a more advanced physiological age and will have a much shorter
GL (Chillet et al., 2006).
Sigatoka disease (SD), caused by the pathogenic fungus Mycosphaerella musicola Leach (Stover, 1972), is an additional cropping
stress that can reduce banana GL. Heavy infestations of this leaf
spot disease can lead to a considerable reduction in the photosynthetic leaf area of the plant, and thus in the bunch weight
(Ramsey et al., 1990). Dessert banana cultivars grown for export
(Cavendish group) are all highly susceptible to this disease in most
banana-growing countries (Jones, 2000). SD is generally controlled
by frequent aerial sprays of fungicides and the use of tailored
cropping techniques, i.e. optimizing banana growth and eliminating SD-infected leaf parts. In Guadeloupe conditions, an average
of 10 fungicide applications is commonly applied throughout
the year.
SD control in export market-oriented banana plantations is
essential because this disease is highly detrimental to fruit quality.
It has been shown that severe SD infestations have a marked effect
on banana ripeness at harvest (Meredith, 1970; Stover, 1972;
Wardlaw, 1972). Bananas harvested from severely SD-infected
plants ripened very quickly after harvest, and were sometimes
already ripe on the plants at harvest. Ramsey et al. (1990) found
that the GL of bananas harvested from highly infected plants was
substantially shortened. However, in these experiments, bananas
were harvested at the commercial grade (3⁄4 full) and their physiological age was not specified. Because fruit growth is generally
delayed when the photosynthetic area is reduced, these bananas
were probably harvested at an advanced physiological stage, which
could therefore possibly explain the reduction in GL noted by these
authors, rather than being due to a direct effect of the SD. It is thus
essential to adopt an experimental approach that integrates the
notion of physiological age of bananas at harvest in order to be able
to differentiate a trophic effect, i.e. delayed growth leading to fruit
harvesting at an advanced physiological age, from a direct effect of
SD on fruit physiology.
This was the purpose of the present study through (i) a survey in
commercial plantations and (ii) an experimental design where the
relationship between Sigatoka disease and greenlife was
investigated.
described by Chillet et al. (2008). The size (diameter) of each
banana was also measured using a slide gauge.
Sampling was carried out at three periods on both plots
between November 2004 and January 2005.
2. Materials and methods
The experiment was replicated through three consecutive
cropping cycles (harvests of September 2006, January 2007 and
August 2007).
The effect of Sigatoka disease on fruit quality has been assessed
under two sets of conditions.
2.1. A survey on the effect of Sigatoka disease on the greenlife of
bananas harvested at 900 dd in commercial plantations
Two commercial plots with the same soil-climate conditions
were selected on two farms in a banana-growing area of Guadeloupe (2500 mm year1 rainfall; ferralitic soil; 30 m elevation;
mean temperature 28 C). One plot had a high SD infestation rate
(plot H), while the other plot (plot L) was barely infested.
For each banana sampling period, 20 banana plants at the
horizontal finger flowering stage were selected per plot. They were
chosen on the basis of homogeneity in plant size and number of
hands per bunch The crop management strategies were similar for
all banana plants: blue polyethylene bunch sleeving, monthly
fertilization, and crop protection treatments. Bunches were harvested at a constant physiological age according to the method
described by Ganry and Meyer (1975), i.e. when the mean daily
temperature sum at the 14 C threshold (STbase 14) between flowering and harvest reached 900 dd. A banana was then cut from the
third hand (median internal fruit) of each bunch, washed, treated
with thiabendazole (500 mg ml1). GL was measured at 20 C as
2.2. Experimental approach: evaluation on the effect of Sigatoka
disease on the greenlife of bananas harvested at 900 dd
Two plots were selected in the Neufchâteau research station,
Guadeloupe (3500 mm year1 rainfall; Andosol; 250 m elevation;
mean temperature 26 C). One plot had a high SD infestation rate
and no fungicide applications were carried out (plot C), so
a maximum SD infestation level had been obtained in this plot;
whereas the other plot (plot T) had undergone a regular SD control
treatment (fungicide spraying and tailored cultivation practices), so
it had a very low SD infestation level. These plots corresponded to
a second cropping cycle.
Twelve banana plants at the horizontal finger flowering
stage were selected per plot. The crop management strategies
were similar for all banana plants: blue polyethylene bunch
sleeving, monthly fertilization, and crop protection treatments
(apart from fungicide sprays). Bunches were harvested at
a constant physiological age, as in the first survey, and fruits
were also sampled and treated in the same way as in the
survey.
For each banana plant of this experiment, SD was quantified at
two development stages, i.e. at flowering and harvest at 900 dd, on
the basis of the two following parameters:
- The youngest leaf spotted (YLS) was assessed according to the
method of Stover and Dickson (1970), which involves the
monitoring of the youngest leaf (from the top of the plant)
bearing at least 10 SD necrotic lesions.
- The severity index (SI) was assessed according to the
method of Stover (1971), and modified by Gauhl et al.
(1993). This method involves a visual estimation of the
necrotic area per plant, scored according to a six-disease
grades scale. The SI for one plant corresponds to the sum of
the scores per leaf.
2.3. Calculation of physiological age of the fruit
Temperature probes (Gemini Data Loggers Ltd., Chichester,
West Sussex, UK) were placed under a forecast shelter on the
experimental plots. Mean daily temperature was calculated on
the basis of hourly temperatures. The physiological age of the
fruit (expressed in degree-days (dd)) was calculated by the mean
daily temperature sums accumulated by fruit at the 14 C
threshold during the flowering-to-harvest period (Ganry and
Meyer, 1975).
2.4. Data analysis
The fruit grades and GLs were compared according to the
production conditions and SD level. Analyses of variance followed
by Newman-Keuls tests (5% threshold) were carried out to compare
the two types of plot (high or low SD infestation) in the two
experiments using the STAT-ITCF software package (1991). The
parameters were correlated via linear, polynomial or logarithmic
regressions using Excel software (2003 version).
M. Chillet et al. / Crop Protection 28 (2009) 41–45
43
45
3. Results
40
3.1. Survey on the effects of Sigatoka disease on the greenlife of
bananas harvested at 900 dd in commercial plantations
The table gives the fruit diameter and GL values measured on
bananas harvested in the two situations (H, high SD severity,
and L, low SD severity) at a constant physiological age of 900 dd,
for three sampling periods. The fruit diameters were similar
(mean 35 mm) for the two disease levels. SD therefore did not
have an impact on the fruit growth, as assessed by the fruit
diameter. However, bananas sampled on the plot with a high SD
infestation had a mean GL of 17.3 d which was significantly
lower than that of bananas harvested from plants that were
barely infested (mean GL of 28 d).
GL (days)
35
30
25
R2 = 0,8681
20
15
R2 = 0,7864
10
5
0
0
10
20
30
40
50
60
70
Severity at Flowering (%)
Fig. 2. Relationship between banana greenlife and SD severity measured at the
flowering stage for bananas harvested at 900 dd in the two experimental plots (A).
4. Discussion
3.2. Experimental approach: evaluation on the effect of Sigatoka
disease on the greenlife of bananas harvested at 900 dd
SD severity was found to range from 35% to 60% on the
untreated plot (assessed at the flowering stage) and from 0% to 20%
on the treated plot (data not shown). The table shows that the mean
harvested fruit diameter ranged from 34.2 mm to 34.7 mm,
respectively on the untreated plot (high SD severity) and on the
treated plot (low SD severity). As noted in the survey under
commercial cropping conditions, there were no significant differences in fruit diameters between the two treatments. However,
a highly significant difference was noticed in the GL of bananas
harvested in the untreated plot (GL of 6.1 d) and in the treated plot
(GL of 28 d).
The YLS evaluated at harvest stage turned out to be a poor
indicator of GL, because all the banana plants in the highly SDinfected plot were found to have the same level (YLSharvest ¼ 1)
(data not shown). For the three other indicators, however, there
was high data variability in the untreated plot (range 2–7 for
YLSflowering) and in the treated plot (range 7–10 for YLSflowering). This
enabled us to establish correlations between GL and YLSflowering
(Fig. 1), GL and severity at flowering (Fig. 2) and GL and severity at
harvest (Fig. 3). The three calculated linear correlation coefficients
were high (>0.7), with the highest being obtained for the SD
severity index at flowering (r2 z 0.86), whereas the lowest (0.7)
was obtained with the YLS parameter at flowering. The relations
are not necessarily linear. They can be logarithmic (r2 ¼ 0.6621 for
YLS at flowering and r2 ¼ 0.7864 for severity at flowering) or
polynomial (r2 ¼ 0.8253 for severity at harvest). Regardless of the
indicator used, the GL decreased as the SD pressure increased even
though the fruits have identical diameters (see Table 1).
The survey and experimental results obtained in this study
revealed that bananas harvested at a same physiological age from
plots with a high SD infestation rate had a much shorter GL than
those from healthy plots, whereas they had the same fruit diameter.
It is noteworthy that bananas harvested on plots with a low SD
infestation rate had an equivalent GL in the survey and in the
experimental design (28 d). This GL is sufficient with respect to the
shipping time and is close to that previously measured for bananas
harvested at 900 dd (Chillet et al., 2006). However, GL values were
lower for bananas from highly infected plots, even if this trend was
not as marked in the survey realized on commercial plots. The
difference in GL, i.e. 17 d (survey) and 6 d (experiment), could likely
be explained by the fact that the SD infestation rate in the survey
plot was less serious than in the experimental plot. Differences in
the plant physiological status due to different crop management
strategies applied in the commercial and experimental plots could
also explain this difference (Bugaud and Lassoudiere, 2005).
Our results highlighted that SD has a direct effect on the extent
of ripeness of bananas harvested at a constant physiological age.
The results of this study showed that the reduction in GL observed
for bananas harvested from highly SD-infected plants resulted from
a direct effect of this leaf spot disease on fruit physiology, and was
not associated with late harvesting of bananas whose growth
would have been slowed down. Bananas harvested at a constant
physiological age (900 dd) from plots with a high SD infestation
rate therefore did not have the GL predicted by the model described
by Jullien (2000), contrary to bananas harvested from plots with
a low SD infestation rate.
Moreover, during these experiments, the reduction in the
photosynthetic area as a result of heavy SD infestation did not lead
45
45
40
40
35
GL (days)
GL (days)
35
30
2
25
R = 0,6621
20
2
R = 0,7003
15
30
25
10
5
5
0
2
4
6
8
10
12
YLS at Flowering
Fig. 1. Relationship between banana greenlife and YLS measured at the flowering stage
for bananas harvested at 900 dd in the two experimental plots (A).
2
R = 0,8005
15
10
0
R2 = 0,8253
20
0
0
20
40
60
80
100
120
Severity at harvest (%)
Fig. 3. Relationship between banana greenlife and SD severity measured at harvest for
bananas harvested at 900 dd in the two experimental plots (A).
44
M. Chillet et al. / Crop Protection 28 (2009) 41–45
Table 1
Fruit diameter and green life of bananas from the two plots of the survey and from
the two plots of the experiment.
Survey
Fruit diameter
(mm)
Greenlife (days)
Experiment
Highly
SD-infected
plot (H)
Barely
SD-infected
plot (L)
Plot with a high
SD infestation
rate (C)
Plot with a low
SD infestation
rate (T)
35.0 1 NS
35.3 0.8 NS
34.2 1.9 NS
34.7 1.1 NS
17.3 1.9**
28.0 1.7**
6.1 3.2**
28.0 3.7**
Fruit diameter (mm) and greenlife (days) of bananas harvested from plots of the
survey and of the experiment. Each value is the mean of triplicate samples of 20
(survey) or 12 (experiment) fruits. Fruits were harvested at 900 dd. ** indicates that
the statistical test was significant at the 5 % probability level. NS indicates that there
was no significant difference at the 5 % level.
to a reduction in the diameter of fruit that had been harvested at
a constant physiological age (900 dd). This was in line with results
obtained previously in plantain infected by Black Leaf Streak
Disease (Mycosphaerella fijiensis), a disease that is related to SD
(Cohan et al., 2004). However, this effect was unexpected and
differed from the effects of other cropping stresses on banana fruit
growth. Stresses associated with mineral deficiency, poor water
supply (excess or shortage), and heavy defoliation, had a very clear
impact on the fruit diameter growth rate (Chillet et al., 2006;
Turner and Barkus, 1982; Goenaga and Irizarry, 1998; Hegde and
Srinivas, 1989). In our experiments, it seemed that the extent of SDinduced defoliation was not enough to slow down fruit growth, and
probably a much higher SD infestation rate (YLS at flowering of less
than 3–4 and/or SI at flowering over 60%) would be required to
impact fruit growth.
The diameter growth of bananas harvested at a constant physiological age was unaffected, whereas the GL measured in bananas
from plots with a high SD infestation rate was markedly reduced,
clearly showing that this was not the result of a trophic effect but
rather to a direct effect of this leaf spot disease on green physiological development stages and on the extent of fruit ripeness.
Our results seemed to indicate a close relation between the level
of SD infestation of banana plants at flowering and GL measured on
bananas harvested at a constant physiological age. This relation was
better for SI than for YLS. This could be explained by the fact that
the SI provides a better quantitative assessment of SD on the plant
than the YLS, which only enables a qualitative evaluation. The
correlation coefficients for these relationships were high as there
were two scattered plots representative of two extreme levels of
disease severity, then no final conclusion should be drawn on the
type of model of the relation between SI and GL. Further experiments are thus required to clearly determine the nature of this
relation, through an increasing disease gradient. It would be
especially important to determine whether this relationship is
linear or influenced by threshold effects.
As SD was clearly shown to have a direct effect on the GL of
bananas harvested at a constant physiological age, it would be now
interesting to investigate the situation with respect to Black Leaf
Streak Disease (BLSD), caused by M. fijiensis. This latter banana leaf
spot disease is considered to be a major production constraint and
is actually more virulent than SD. BLSD was described for the first
time in Latin America (Honduras) in 1972 (Stover and Dickson,
1976) and has been spreading along the Caribbean arch, with recent
introductions in Grenada and Trinidad (Fortune et al., 2005)
threatening West Indian banana crops. Nevertheless there is no
evidence to date that analogous mechanisms are involved in host/
pathogen interactions for SD and BLSD.
These results also question the physiological mechanisms
involved in this relationship between a banana leaf disease and its
direct effect on fruit physiology. Are metabolites synthesized by the
pathogenic fungus that will have an impact on fruit physiology? For
BLSD, it was shown that M. fijiensis secretes a toxin, the juglone,
that can induce necrotic lesions on leaves (El Hadrami et al., 2005;
Hoss et al., 2000). An equivalent toxin might also be secreted by
M. musicola since this toxin seems to be involved in BLSD infection
process (Harelimana et al., 1997; Molina and Krausz, 1989). It would
thus be interesting to determine whether this toxin is also involved
in mechanisms responsible for the reduction in GL in highly
infected banana plants. It is also possible that the development of
the necrosis caused by M. musicola may elicit the pathway of
ethylene biosynthesis, which will initiate changes in GL in mature
bananas.
Finally, secondary plant metabolites involved in resistance
mechanisms might also have an impact on this relationship. For
other pathosystems, it has been shown that intermediate metabolites such as methyl-jasmonate and salicyclic acid, which are
involved in the host–pathogen relationship (Hammerschimdt,
2006), also have an effect on fruit ripening (Fan et al., 1998). In
addition, some hormones such as AIA and GB3 may be involved
(Rossetto et al., 2003; Purgatto et al., 2002, 2001). Modification of
the hormonal balance following a pathogen attack could be
responsible for the effects noted on fruit quality. Many additional
experiments should now be carried out to gain further insight into
the relationship highlighted in the present study.
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