Fresh Produce ©2007 Global Science Books

Postharvest Biology and Handling of Banana Fruit

Xuewu Duan1 • Daryl C. Joyce2 • Yueming Jiang1*

1 South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, P. R. China
2 School of Land, Crop and Food Sciences, The University of Queensland, Gatton, Qld. 4343, Australia

Corresponding author: * ymjiang@scbg.ac.cn

ABSTRACT
Banana (Musa sp.) is one of the most economically important fruit crops in the world. The banana fruit is generally harvested and stored
in a mature green state. Fruit ripening involves a transient burst in ethylene production that coordinates ripening-associated process. These
processes include the respiratory climacteric, pulp softening, peel de-greening, and production of aroma compounds. Avoidance of
exposure to ethylene and control of endogenous ethylene synthesis are key measures for banana storage. Other important factors that
influence banana fruit storage life are pathogen development, mechanical damage, and variable maturity. The interaction of these factors
can lead to uneven and unpredictable ripening that has adverse implications for marketability. Low temperature storage is highly effective
in reducing decay and extending the storage life of harvested banana. However, banana fruit are chilling sensitive and storage at sub-
optimal temperatures results in injury symptoms that include peel discoloration and abnormal ripening. These symptoms are common
when banana fruit are stored at temperatures below about 13°C. Controlled atmosphere (CA) storage or modified atmosphere (MA)
packaging constitute adjunct or alternative technologies to extend the green life of harvested fruit. These technologies can be effective at
ambient temperatures, particularly in combination with the use of ethylene absorbing compounds and/or treatments that prevent ethylene
action or inhibit rots. However, if CO2 concentrations become too high, the fruit may fail to ripen normally. The relatively recently
introduced ethylene binding site blocker, 1-methylcyclopropene (1-MCP), can effectively inhibit ethylene action on banana fruit. Applied
as a gas, like ethylene, 1-MCP has demonstrated potential for the modulation of ripening and senescence processes in banana fruit.
Overall, postharvest research on banana fruit remains focused on control of ethylene synthesis and action and on suppression of disease
development, including by chemical-free means.
_____________________________________________________________________________________________________________

Keywords: disease, ethylene, physiology, ripening, storage

Abbreviations: 1-MCP, 1-methylcyclopropene; ABA, abscisic acid; ACC, 1-aminocyclopropane-1-carboxylic acid; ACO, 1-amino-
cyclopropane-1-carboxylic acid oxidase; ACS, 1-aminocyclopropane-1-carboxylic acid synthase; ATP, Adenosine triphosphate; CA,
controlled atmosphere; Chl, chlorophyll; CI, chilling injury; GA, gibberellic acid; IAA, indole-3-acetic acid; MA, modified atmosphere;
N2O, nitrous oxide; NO, nitric oxide; PAL, phenylalanine ammonia lyase; PEP, phosphoenolpyruvate; PG, polygalacturonase; PEL,
pectate lyase; PME, pectin methylesterase; PPO, polyphenol oxidase; RH, relative humidity; SPS, sucrose phosphate synthase and SuSy,
sucrose synthase

CONTENTS

INTRODUCTION...................................................................................................................................................................................... 141
POSTHARVEST BIOLOGY ..................................................................................................................................................................... 141
Respiration............................................................................................................................................................................................. 141
Ethylene................................................................................................................................................................................................. 141
Volatiles ................................................................................................................................................................................................. 142
Starch and sugar .................................................................................................................................................................................... 142
Texture................................................................................................................................................................................................... 143
Colour.................................................................................................................................................................................................... 143
Chilling injury ....................................................................................................................................................................................... 143
FACTORS AFFECTING STORAGE LIFE ............................................................................................................................................... 144
Fruit maturity......................................................................................................................................................................................... 144
Temperature........................................................................................................................................................................................... 144
Water loss and humidity ........................................................................................................................................................................ 144
Ethylene................................................................................................................................................................................................. 144
Mechanical damage ............................................................................................................................................................................... 145
Disease .................................................................................................................................................................................................. 145
Crown rots......................................................................................................................................................................................... 145
Anthracnose ...................................................................................................................................................................................... 145
Other postharvest diseases ................................................................................................................................................................ 145
POSTHARVEST TECHNOLOGY ............................................................................................................................................................ 145
Modified atmosphere (MA) and controlled atmosphere (CA) ............................................................................................................... 145
Coatings................................................................................................................................................................................................. 146
Anoxia or low oxygen ........................................................................................................................................................................... 146
1-Methylcyclopropene (1-MCP)............................................................................................................................................................ 146
Plant growth regulators.......................................................................................................................................................................... 147
Nitric oxide (NO) and nitrous oxide (N2O) ........................................................................................................................................... 147

Received: 4 August, 2007. Accepted: 6 September, 2007.

Invited Review
 Postharvest biology 1(2),
Fresh Produce and handling
140-152 of banana
©2007 fruit.
Global et al.
DuanBooks
Science

Control of postharvest disease ............................................................................................................................................................... 148

Chemicals ......................................................................................................................................................................................... 148
Natural extracts ................................................................................................................................................................................. 148
Heat treatment................................................................................................................................................................................... 148
Biological control.............................................................................................................................................................................. 148
CONCLUSIONS........................................................................................................................................................................................ 149
ACKNOWLEDGEMENTS ....................................................................................................................................................................... 149
REFERENCES........................................................................................................................................................................................... 149
_____________________________________________________________________________________________________________

INTRODUCTION during ripening is correlated with activation of cytosolic py-

ruvate kinase and/or PEP carboxylase, which control phos-
Banana is one of the most economically important fruit phoenolpyruvate (PEP) and pyruvate contents, respectively
crops. Banana production in the world was about 68 million (Beaudry et al. 1989; Ball et al. 1991). Complex allosteric
metric tons in 2005, with India, Brazil, China, the Philip- control of pyruvate kinase and PEP carboxylase is the
pines, Ecuador, and Indonesia contributing 63% of the total model for control of cytosolic glycolysis and PEP partition-
production (Table 1; FAO 2005). In the international fruit ing during banana fruit ripening (Law and Plaxton 1995,
trade, banana ranked 1st and 2nd among all fruits in terms of 1997; Turner and Plaxton 2000). Another important respira-
quantity and value, respectively. Ecuador was the biggest tion rate-controlling step in glycolysis in banana fruit is re-
banana exporter, followed by the Philippines, Costa Rica versible phosphorylation, i.e. fructose 6-phosphate to fruc-
and Columbia. These four countries accounted for 54% of tose 1,6-bisphosphate by fructose 1,6-bisphosphatase, ATP-
total export quantity in 2005 (FAO 2005). dependent phosphofructokinase and/or PPi-dependent phos-
Bananas are perennials grown and harvested year-round. phofructokinase (Beaudry et al. 1987, 1989; Ball et al.
Banana fruit have a short postharvest life of approximate 1991). Turner and Plaxton (2003) suggested that the pri-
10-15 days at ambient temperatures. This short postharvest mary and secondary control of glycolytic flux in banana
longevity limits trade and consumption, especially in deve- cultivar ‘Cavendish’ during ripening is exerted as the level
loping countries. In places such as China and India, there is of PEP and fructose 6-phosphate metabolism, respectively.
a shortage of cold chain facilities. The main factors affec- In banana fruit, there appears to be little or no contribution
ting banana fruit quality are rapid physiological deteriora- to electron transport by the alternate oxidase in the climac-
tion, physical damage, decay, chilling injury and uneven teric phase (Theologis and Laties 1978). The mechanisms
and unpredictable ripening. Postharvest technologies have of the initiation of the respiratory climacteric in harvested
been developed to maintain postharvest quality and extend bananas fruit, concerning respiration pathways, merits fur-
the postharvest life of banana fruit. Moreover, research on ther investigation.
postharvest biology and technology for banana fruit conti-
nues at a rapid pace. This overview considers the current Ethylene
status of postharvest research on banana fruit, with an em-
phasis on postharvest handling. Ethylene plays an important role in ripening and senescence
of harvested fruits. Ethylene, applied at as low as 0.1 Pl/l
POSTHARVEST BIOLOGY for 1 day is normally sufficient to initiate banana ripening
(Chang and Hwang 1990). Two ethylene production sys-
Respiration tems operate in harvested banana fruit. The low basal rate
of ethylene production is contributed to System 1 ethylene.
Banana is a typical climacteric fruit. It undergoes a rapid System 2 ethylene is responsible for the autocatalytic cli-
increase in ethylene synthesis, followed by a 4- to 5-fold macteric rise in ethylene production (McMurchie et al.
increase in respiration rate as indicated by CO2 production 1972). In higher plants, ethylene is biosynthesized from
during ripening (Barket and Solomos 1962; Pathak et al. methionine via a pathway in which 1-aminocyclopropane-
2003). CO2 production by banana fruit in the climacteric 1-carboxylic acid synthase (ACS) and 1-aminocyclopro-
phase is attributed to an increased flux of carbon through pane-1-carboxylic acid oxidase (ACO) catalyze the reac-
the glycolytic pathway to mitochondria, accompanied by tions of S-adenosylmethionine conversion to 1-aminocyclo-
conversion of starch to sucrose (Beaudry et al. 1987, 1989; propane-1-carboxylic acid (ACC) and ACC conversion to
Hubbard et al. 1990; Liu et al. 2004). The enhanced glyco- ethylene, respectively (Yang and Hoffman 1984; Wang et al.
lytic flux in association with mitochondrial respiration rise 2002). ACS and ACO have been isolated and purified from
in the climacteric period can generate adenosine triphos- banana fruit, and both enzymes appear to be encoded by
phate (ATP) for the conversion of starch to sucrose (Hill multigene families (Liu et al. 1999; Do et al. 2005; Huang
and Rees 1994). et al. 2006). In banana fruit cultivar ‘Grand Nain’, ethylene
Initiation of the respiratory climacteric in bananas fruit production is at least partially regulated by transcriptional

Table 1 Total production, area harvested, export quantity and export value of bananas in the major producing countries
in 2005 (data from FAO 2005).
Country Total production Area harvested Export quantity Export value
(1000 tonnes) (1000 Ha) (1000 tonnes) (million $)
India 11,710 404 13 4
Brazil 6,703 491 211 51
China 6,667 274 42 18
Philippines 6,298 417 1,964 735
Ecuador 6,118 221 4,085 1260
Indonesia 4,503 315 3 1
Costa Rica 2,353 48 1,597 513
Mexico 2,250 76 64 22
Thailand 1,865 140 25 7
Colombia 1,765 64 1381 453
Burundi 1,539 303 0 0

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levels of MA-ACS1 until the climacteric rise and by reduc- Starch and sugar
tion of ACC oxidase activity possibly through limited in
situ availability of its cofactors once ripening has com- Starch is the principal component of green banana fruit. It
menced, which in turn characterizes the sharp peak of ethy- constitutes approximately 20-25% of the fresh weight or
lene production (Liu et al. 1999). Golding et al. (1998) about 85% of the dry weight of the pulp tissues in mature
showed an involvement of a negative feedback regulatory unripe fruit. During the climacteric phase, starch is rapidly
mechanism in ethylene biosynthesis in banana fruit once degraded and most of the polysaccharide is converted into
ripening commenced. Inaba et al. (2007) found that expo- soluble sugars, mainly sucrose (Lii et al. 1982; Cordenunsi
sure of ‘Grande Naine’ banana fruit ripened previously by and Lajolo 1995).
propylene treatment to 1-MCP increased ethylene produc- Starch–sucrose transformation during banana fruit
tion concomitantly with an increase in ACS activity and ripening involves a series of enzymes and their related path-
ACC content, and prevented a transient decrease at the MA- ways (Zhang et al. 2005). It is generally accepted that -
ACS1 transcript in the pulp. In contrast, in the peel of amylases play a key role in starch degradation in banana
ripened fruit, 1-MCP treatment delayed the increase in fruit. These amylases hydrolyze -1,4-glucosidic linkages
ethylene production with subsequent ripening by reduction in starch and remove successive maltose units from the non-
of the increase in the MA-ACS1 and MA-ACO1 transcript reducing ends of the chains. Garcia and Lajolo (1988)
levels and inhibition of ACS and ACO activities. Therefore, found that -amylase activity increased before the initiation
it was suggested that ethylene biosynthesis in ripening ba- of the respiration climacteric, paralleling with the decrease
nana fruit may be negatively controlled in the pulp tissues, in starch, while activities of -amylase and glucosidase in-
but positively in the peel tissues at the transcriptional level. creased significantly only at the climacteric peak when the
As with the respiratory climacteric, the detailed mechanism starch had already been degraded. Nascimento et al. (2006)
of regulation of ethylene production in the banana fruit still reported that -amylase activity in ‘Nanicao’ banana fruit
needs to be fully elucidated. was well correlated to the decrease in starch, suggesting the
primary up-regulation by de novo synthesis. In 1-MCP-
Volatiles treated fruit, the amount of -amylase protein was almost
undetectable even though there was a strong induction of
The aroma compounds of banana fruit consist mainly of 3- transcription. Similar results were found by Purgatto et al.
methylbutyl esters, acetates, butanoates, pentan-2-one, (2001) and Rosecler et al. (2003) in ‘Nanicao’ banana fruit
esters, alcoholic compounds, esters, alcohols, aldehyde, treated with indole-3-acetic acid (IAA) and gibberellic acid
ketone, heptyl-acetate, isoamyl acetate, 2-methylbutyl (GA). These findings confirmed that activity of -amylases
acetate, and 2-heptyl acetate (Tressl et al. 1970; Shiota is essential for starch degradation in banana fruit during
1993; Nogueira et al. 2003; Pino et al. 2003). Perez et al. ripening.
(1997) found 25 free volatiles and 25 glycosidically bound Sucrose synthase (SuSy) and sucrose phosphate syn-
volatiles from ‘Valery’ and ‘Pequena Enana’ banana pulp thase (SPS) are key enzymes in sucrose metabolism. Corde-
using an Amberlite XAD-2 column. The free volatiles were nunsi and Lajolo (1995), Nascimento et al. (1997) and Nas-
identified as an ester, 14 alcohols, 2 aldehydes, 4 acids, 2 cimento et al. (2000) investigated the changes in SuSy and
ketones and 2 terpenes (Table 2). Aromatic compounds are SPS activities during development and ripening of harves-
synthesized during banana fruit ripening (Salmon et al. ted ‘Nanicao’ banana fruit and also the changes in carbo-
1996). For green ‘Grande Naine’ banana fruit, only two hydrates in fruit left to ripen on the plant. SPS was present
major peaks identified as isoamyl acetate and butyl acetate during fruit development, but at a very low activity level.
were found, whereas many peaks were detected for overripe SuSy activity was high and remained constant throughout
banana fruit (Boudhrioua et al. 2003). Constituent volatile the entire starch synthesis phase, followed by a reduction
compounds in banana fruit may vary with differences in during starch breakdown and disappearance in the post-
cultivars, geographic origin and postharvest handling proto- climacteric phase. Slower starch breakdown was related to
col. lower sucrose content and SPS activity, but higher SuSy

Table 2 Free and glycosidically bound volatile compounds in ‘Pequena Enana’ banana fruit (data from Perez et al. 1997).
Free volatile compounds ng/g (FW) Glycosidically bound volatiles ng/g (FW)
Methyl acetate 2.62 3-Methylbutanoic acid 12.2
Ethyl acetate 0.48 4-Methylhydroxypentan-2-one 2.74
Butan-2-one 0.17 Hexanol 1.18
Benzene 0.13 Hexanoic acid 10.52
Pentan-2-one 1.27 Hex-3-enoic acid 6.54
Butyl acetate 0.15 Hex-2-enoic acid
Toluene 0.10 2-(2-Ethoxyethoxy)ethanol 4.16
3-Methylbutyl acetate 0.76 Benzoic acid 5.42
2-Methylpropanol 0.22 2-Phenylethydecane 7.08
Pentyl acetate 0.03 2,5,6-Trimethyldecane 4.72
Pentan-2-ol 0.75 Phenylacetic acid 2.10
2-Methybutyl butanoate 0.01 Decan-1-ol 12.36
Butan-1-ol 0.18 3-Oxo-pentanoic acid 12.64
Hex-2-enal 0.04 Eugenol 8.00
2-Pentyl 2-methylpropanoate 0.04 -Decalactone 9.70
2-Butyl butanoate 0.01 9-Oxononanoic acid 1.68
3-Methylbutanol 0.19 Dodecanoic acid 1.92
3-Methylbutyl butanoate 0.01 Elimicine 5.42
2-Buthoxyethanol 0.10 3,4-Dimethoxyacetopenone 4.18
Acetic acid 0.38 Methyleugenol 0.94
Methyl decanoate 2-Furyloctanoic acid 13.38
Propanoic acid 0.12 Jasmonic acid 3.46
3-Methylbutanoic acid 0.08 3,4,5-Trimethoxyacetophenoone 2.36
Butanoic acid 0.14 Tetradecanoic acid 8.22
Pentanoic acid 0.19 Hexadecanoic acid

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 Postharvest biology and handling of banana fruit. Duan et al.

activity, for attached fruits as compared to detached fruits. pH-dependent manner, are required for fruit softening
Therefore, it seems that SPS is more important than SuSy (Brummell and Harpster 2001; Hayama et al. 2003; Sane et
for starch degradation during banana ripening. al. 2005). Expansins have been identified from ‘Harichhal’
During natural ripening of banana fruit, there is modu- and ‘Williams’ banana fruits and they are associated with
lation of the activity for starch-metabolized enzymes related fruit maturation and softening (Trivedi and Nath 2004;
to sucrose biosynthesis, such as SuSy and SPS (Cordenunsi Wang et al. 2006). However, as with most aspects of banana
and Lajolo 1995). The conversion of starch to sucrose is the ripening physiology and biochemistry, the detailed mecha-
result of the combined action of these enzymes. Purgatto et nistic role of expansins in fruit softening needs to be further
al. (2001) reported that GA treatment delayed starch degra- investigated.
dation and sucrose formation of harvested ‘Nanicao’ banana.
However, SuSy and SPS activities and their transcript levels Colour
were not affected, indicating no direct relation of these suc-
rose-metabolizing enzymes to prevention of sucrose ac- Peel colour is the most obvious character that changes
cumulation. Impairment of sucrose synthesis could be a during banana fruit ripening and is the major eating crite-
consequence of lack of substrate, since starch degradation rion for consumers. The typical change during ripening is
was inhibited. Nascimento et al. (1997, 2000) suggested loss of green and appearance of yellow. The loss of green
that substrate limitation could play an important role in the colour is due to chlorophyll (Chl) degradation from approx-
regulation of starch breakdown of harvested ‘Nanicao’ ba- imate 50-100 mg/g fresh weight (FW) to almost zero in ripe
nana because SPS and SuSy activities were shown to be up- fruit (Seymour 1993). The Chl degradation pathway in high-
and down-regulated, respectively. Thus, starch degradation er plants has been largely elucidated (Matile et al. 1999;
and sugar biosynthesis in banana fruit during ripening Hortensteiner 2006). There might be two Chl degradation
might be, at least partially, controlled at the transcriptional pathways in banana fruit, the chlorophyllase and chloro-
level. phyll oxidase pathways (Janave 1997). Janave and Sharma
(2004) confirmed the presence of chlorophyllase, magne-
Texture sium-dechelatase, pheophorbide a oxygenase, red fluores-
cent catabolite reductase and Chl oxidase in banana peel tis-
Texture is usually expressed as fruit firmness and is an im- sues.
portant quality attribute of harvested banana fruit. Textural Chl catabolism of banana fruit may differ somewhat
changes during ripening of banana fruit result from the when the fruit is held at high temperature, which inhibits
structural and compositional modification in cellular walls. Chl degradation (Seymour et al. 1987). Matile et al. (1996)
Decrease in fruit texture (softening) is mainly attributed to suggested that Chl should be solubilized from the thylakoid
solubilisation and depolymerisation of cell wall polysaccha- membranes prior to degradation and is apparently transpor-
rides, including pectins, hemicellulose and cellulose by a ted to the chloroplast envelope, where chlorophyllase is
series of activities of hydrolase, transglycosylase and pro- located. Ultrastructure studies of banana fruit peel tissues
teins, such as expansins. The major enzymes, involved in reveals that thylakoid membranes are retained to degree in
fruit firmness changes during ripening of banana fruit, are the fruit ripened at high temperature, which may hinder
polygalacturonase (PG), pectin methylesterase (PME), pec- release of Chl from thylakiod membranes to the chloroplast
tate lyase (PL), cellulose, -glucanase, and -galactosidase envelope, resulting in inhibition of the Chl degradation
(Prabha and Bhagyalakshmi 1998; Peumans et al. 2000; (Blackbourn et al. 1990). Drury et al. (1999) reported that
Pua et al. 2001; Marin-Rodriguez et al. 2003; Payasi and the colour-degraded products of ‘Grande Nain’ banana fruit,
Sanwal 2003; Ali et al. 2004; Lohani et al. 2004; Asif and chlorophyllide and pheophorbide, were not detected at any
Nath 2005; Imsabai et al. 2006). Ali et al. (2004) suggested stage of fruit ripening at 20 or 35°C. However, a non-fluo-
that PG, PME, (14)--glucanase and -galactosidase rescent Chl product accumulated to a higher concentration
might contribute importantly to ‘Mas’ banana fruit soften- at 20°C than at 35°C, indicating that the ‘stay-green’ effect
ing. However, Lohani et al. (2004) reported that the activi- of fruit ripening at 35°C was not due to inhibition of pheo-
ties of PG and cellulase increased sharply during softening phorbide a oxygenase. The biochemical mechanism of Chl
of ‘Harichhal’ banana fruit but PME activity increased grad- breakdown in banana fruit skin during ripening at high tem-
ually. Recent study indicated that fruit softening in ‘Harich- perature requires a full investigation.
hal’ banana results from the downstream effects of at least
four PG genes which are differentially expressed at various Chilling injury
ripening stages. MAPG3 and MAPG4 are believed to be
ripening-related and regulated by ethylene, whereas MAPG2 Chilling injury (CI) is a physiological disorder in banana
was associated more with senescence (Asif and Nath 2005). and other subtropical and tropical fruit, which occurs upon
Degradation of pectins requires the combined action of exposure to low, but non-freezing, temperatures. Storage of
methylesterases, which remove methoxy groups from pectin, banana below 13°C can lead to the development of CI
and depolymerases, which cleave the bonds between galac- (Wills et al. 1998). The CI symptoms depend on cultivar,
turonate units. Pectate lyase (PEL) catalyzes the cleavage of maturity, and temperature by time of low temperature ex-
(14) galacturonan linkages of pectate by a -elimination posure (Nguyen et al. 2003; Wang et al. 2006). The most
reaction, generating 4,5-unsaturated oligo-galacturonates. common visual symptoms of CI on banana fruit are a dull
Payasi and Sanwal (2003) reported that PEL activity was yellow skin, browning of the skin, failure to ripen, harden-
not detected in preclimacteric ‘Harichhal’ banana fruits, but ing of the central placenta, increased susceptibility to me-
increased progressively from the early climacteric phase, chanical injury, and in severe cases, flesh browning (Jiang
then reached maximum level at climacteric peak, and et al. 2004; Ratule et al. 2006).
finally declined in postclimacteric phase, indicating that Cellular membrane damage is typically an early CI res-
PEL might play a role in fruit softening. PELs have been ponse (Marangoni et al. 1996). Low temperature-induced
purified and characterized from banana fruit (Payasi and changes in the physical properties of cell membranes, due
Sanwal 2003). Pua et al. (2001) and Marin-Rodriguez et al. to modifications in the physical state of membrane lipids,
(2003) cloned PEL genes in ‘Williams’ and ‘Grande Naine’ cause imbalances in metabolism. Consequences of disrup-
banana pulp and found that transcripts of two PEL cDNAs tion to the various membranes are breakdown of cellular
were not detectable in unripe preclimacteric fruits. However, compartmentalization, loss of function of membrane-associ-
they began to accumulate as ripening progressed and re- ated proteins, death of the cells and the appearance of CI
mained at a high level in overripe fruit, which supports the symptoms (Marangoni et al. 1996; Wills et al. 1998). Ethy-
role of PEL in fruit softening. lene binding is intrinsic to cellular membranes and CI of
Apart from the hydrolases discussed above, expansins, a ‘Williams’ banana fruit appears to be associated with sup-
class of proteins inducing extension of cellular walls in a pressed ethylene binding capacity, resulting in a failure of

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fruit ripening and softening (Jiang et al. 2004). Wang et al. bananas at 15 and 18ºC instead of room temperature (26-
(2006) suggested that ethylene might alleviate CI of ‘Wil- 27°C) only temporarily reduced spotting. However, peel
liams’ banana fruit. Increased tolerance of banana fruit pre- spotting was completely prevented at 12°C.
treated with propylene to low temperature-induced chilling To delay ripening and reduce losses, green bananas are
was related to higher post-storage ethylene production rates shipped and stored at 13-14°C (Sommer and Arpaia 1992).
and enhanced expression of MaExp1 and MaExp2. Symptoms of CI generally occur at below 13°C (Hewage et
When subjected to chilling, browning of the banana al. 1996). Exposure to temperatures higher than 30°C cau-
fruit is associated with oxidation of phenolic compounds. ses heat injury. Some symptoms of heat injury are similar to
Nguyen et al. (2003) reported that CI development in those of CI, such as browning of the peel and increased
‘Kluai Khai’ and ‘Kluai Hom Thong’ banana fruit peel was moisture loss. Jiang et al. (2002) reported that exposure of
highly and inversely correlated with the level of free phe- ‘Williams’ banana fruit to 45°C for 45 min caused skin
nolic compounds, and highly and positively correlated with browning. High temperature treatment also resulted in in-
the activities of polyphenol oxidase (PPO) and phenylala- creased mass loss. For ‘Cavendish’ banana fruit, the time
nine ammonia lyase (PAL). Modified atmosphere condi- leading to a 70% mass loss (wet basis) at 30°C was about
tions reduced CI symptoms, which was related to the dec- twice at 40°C (Nguyen and Price 2007). Furthermore, the
reases of PAL and PPO activities in peel tissues (Nguyen et skin of heat-injured fruit may fail to turn yellow due to sup-
al. 2004). The increase of membrane permeability was res- pressed chlorophyll breakdown. The pulp becomes watery
ponsible for decompartmentalization between PPO (en- and translucent. The conversion of starch into sugars may
zyme) and phenolic compounds (substrates), and peel brow- not be triggered, and thus the pulp may not sweeten (Black-
ning (Nguyen et al. 2003; Ratule et al. 2006). Some pre- bourn et al. 1990; Zhang et al. 1993; Janave 1997).
treatments, including with abscisic acid (ABA), jasmonic
acid derivative and putrescine, help maintain partial mem- Water loss and humidity
brane integrity in banana fruit peel at low temperature and,
thereby, alleviate CI (Chaiprasart et al. 2002; Wang et al. Water loss can be highly problematical as it results not only
2003). The degree of CI may be mediated by the level of in direct quantitative losses in weight, but also in deteriora-
endogenous antioxidant activity. Wang et al. (2003) found tion in appearance, texture and nutritional quality in ad-
that infiltration treatment with 200 M ABA and 5 mM put- dition to accelerated and/or exacerbated expression of
rescine enhanced superoxide dismutase and peroxidase acti- symptoms of injuries (Akkaravessapong et al. 1992; Wills
vities, and delayed the appearance of CI of ‘Brazil’ banana et al. 1998). Losses of water reduce turgor (Banks and
fruit stored at 8°C. Kondo et al. (2005) suggested that the Joseph 1991), which contribute to decreased firmness and
concentrations of endogenous jasmonate, total phenolics may accelerate the ripening of banana fruit (Burdon et al.
and ascorbic acid, superoxide dismutase activity were 1994).
linked to the degree of CI of ‘Namwa’ banana. The rate of water loss depends on external and internal
factors. External factors include temperature, relative humi-
FACTORS AFFECTING STORAGE LIFE dity (RH), air movement and atmospheric pressure. At an
ambient RH of 95-100%, fruit lose little or no moisture and
Fruit maturity the ripening period is maximal. As humidity decreases, the
rate of water loss increases and the ripening period reduces.
Maturity at harvest determines the potential storage life and Ahmad et al. (2006) and Ullah et al. (2006) reported that
final banana fruit quality. Bananas are harvested at green high RH delayed ripening and produced good eating quality
mature stage and are ripened at or near the marketplace. of ‘Cavendish’ banana fruit compared with those held at
The more mature banana fruit is at harvest, the shorter it is low humidity. However, wetting (free moisture) can lead to
ripening period (Madamba 1977). Immature fruit are more peel splitting and encourage microbial decay (Ullah et al.
subject to shrivelling and mechanical damage, and are of 2006). Thus, the optimum RH for storage of banana fruit is
inferior flavour quality when ripe (Kader 1999; Ahmas et al. 90-95%.
2001). They also constitute reduced yield in terms of sale- Internal factors that modulate the rate of water loss
able weight. Moreover, fruit picked either too early or too include morphological and anatomical characteristics (e.g.
late in their season are more susceptible to postharvest phy- surface-to-volume ratio), and surface injury (Wills et al.
siological disorders than fruit picked at optimum maturity 1998). The greater the surface area to volume ratio, the
(Kader 1999). Within the harvest maturity band, the green shorter the postharvest life. Typically, large fruit lose less
mature stage at which bananas are harvested is determined water than small fruit. Fruit with thin skins lose more water.
by the time required to get them to market. Higher peel permeability leads to a higher rate of water loss.
As banana fruit matures, their cross-sectional diameter Also, a higher density of stomata may lead to a higher rate
increases. Fruit angularity also changes during growth and of water loss, which in turn can accelerate ripening (Shaun
maturation. As fruit approach full maturity, the angles and Ferris 1998).
become less acute. Fruit angularity can be used to predict
the optimum harvest date (Sommer and Arpaia 1992). Ethylene

Temperature As mentioned earlier, the postharvest physiology of banana

is characterized by a green life phase followed by a burst in
Temperature is generally the most important factor deter- ethylene production that signals the beginning of ripening-
mining the postharvest life of fruits and vegetables, and associated processes. These processes include the respira-
markedly affects their rates of respiration and general meta- tory burst, fruit softening, peel degreening and production
bolism. Typically, for every 10ºC increase, respiration rate of aroma compounds. Ethylene production is essential for
increases between 2- and 4-fold (Wills et al. 1998). ripening of banana fruit, and determines the time from
Bananas storage life decreased as external temperature harvest to the respiratory climacteric. That is, it determines
increased over the range 15-35ºC. The relationship was the green life or preclimacteric period (Clendennen and
logarithmic and described by the equation: log g = mT + c; May 1997). An atmospheric ethylene concentration of 0.1
where, g = preclimacteric period at temperature T, and m Pl/l can trigger internal ethylene production and the respira-
and c are constants. A 1ºC reduction increased storage pe- tory climacteric, and thereby shorten the pre-climacteric pe-
riod by 1-2 days (Marriott 1980). Trakulnaleumsai et al. riod (Chang and Hwang 1990).
(2006) reported that low temperature significantly reduced Exogenous ethylene application is routinely used to ini-
the senescence symptom of peel spotting, which starts with tiate uniform ripening of bananas. Conversely, commercial
small browned spots and then becomes large area with ad- strategies for banana fruit handling, transport and storage
vanced ripening, in ‘Sucrier’ banana fruit. Holding ripening are based on avoiding exposure to ethylene and/or attemp-

144
 Postharvest biology and handling of banana fruit. Duan et al.

ting to minimize ethylene production and action. These stra- peyre De Bellaire and Mourichon 1997b; De Lapeyre De
tegies include temperature and atmosphere control (Wills et Bellaire et al. 2000a). Appressoria send out infection pegs
al. 1998; Marchal 1998; Watkins 2002; Wang 2006). A rela- and form limited infection hyphae, giving rise to quiescent
tively new strategy for controlling ethylene binding and anthracnose (Swinburne and Brown 1983; de Lapeyre de
thus ripening and senescence of fruit is treatment with 1- Bellaire and Mourichon 1997b; Chillet et al. 2006). If the
methylcyclopropene (1-MCP) (Sisler and Serek 2003). 1- bananas are bruised, rot can develop in green banana fruit
MCP effectively delayed banana fruit ripening and thereby and then the lesions expand as the fruit ripens (Chillet et al.
extended their green life (Jiang et al. 1999b; Sisler and 2006). This wound anthracnose form can trigger an early
Serek 2003; Watkins 2006). However, the response of har- fruit ripening (Peacock 1973).
vested banana fruit to 1-MCP is dependent upon fruit matu- It has been reported that ethylene can trigger C. musae
rity, treatment temperature and concentration, and exposure infections, accelerate conidial germination and increase ap-
time (Jiang et al. 1999b; Harris et al. 2000; Bagnato et al. pressorial number in relation to banana anthracnose (Flaish-
2003; Pelayo et al. 2003). One hour exposure at 20ºC to man and Kolattukudy 1994; de Lapeyre de Bellaire et al.
1000 nl/l 1-MCP gas essentially eliminated ethylene-stimu- 2000b). However, Chillet et al. (2006) determined that ethy-
lated ripening effects of mature green ‘Cavendish’ banana lene was not directly involved in triggering rot development
while exposure for 12 h at 20ºC to just 50 nl/l 1-MCP had a as quiescent anthracnose symptoms appeared only after
similar effect (Jiang et al. 1999b). ‘Grande Naine’ banana fruit began ripening. In contrast,
wound anthracnose developed just as quickly in ‘green 1-
Mechanical damage MCP-treated bananas’ as in ‘yellow ripening bananas’. For
wound anthracnose, contrary to quiescent anthracnose, rot
Mechanical damage is a factor contributing to postharvest development was not dependent on the degree of peel ripe-
deterioration of banana fruit. Fruit damage during handling ness.
is beneficial for ethylene production. If ethylene production
is sufficient to initiate the climacteric response, fruit can Other postharvest diseases
ripen prematurely. Loss of fruit weight is another result of
mechanical injury, with consequences of lower market qua- Other commercially postharvest disease of banana fruit in-
lity and price. Accelerated weight loss is due to breakdown clude cigar-end rot (Trachysphaera fructigena and V. theo-
of cellular walls and increased permeability of the outer tis- bromae), finger or stem-end rot (L. theobromae), thielaviop-
sue layers to water vapour. Damage can also lead to secon- sis or ceratocystis rot (T. paradoxa), pitting disease (Pyricu-
dary infection, which further increases the rate of water loss laria grisea), squirter disease (N. sphaerica) and speckle
and reduces quality (Dadzie and Orchard 1997; Liado and and black tip (D. torulosa) (Sommer et al. 1992; Ploetz
Domingues 1998; Wills et al. 1998). 1998). All of these diseases can reduce the quality and post-
harvest longevity of the fruit.
Disease
POSTHARVEST TECHNOLOGY
Crown rots
Modified atmosphere (MA) and controlled
Crown rot is a severe postharvest disease in banana-grow- atmosphere (CA)
ing areas around the world. It is caused by a complex of
fungi, including Fusarium roseum, F. proliferatum, Lasio- The biochemical basis of MA/CA is to extend the longevity
diplodia theobromae, Thielaviopsis paradoxa, Verticillium of fruit by slowing metabolic processes at high CO2 con-
theobromae, Nigrospora sphaerica, Deightoniella torulosa, centrations and low O2 levels (Yahia 1998; Wang 2006).
and Colletotrichum musae (Sommer et al. 1992; Ploetz Optimal compositions of CA and MA storage for fresh pro-
1998; Ranasingle et al. 2002). Inoculations with various duce vary according to its genotype and maturity or ripe-
combinations of fungi show that the greatest damage results ness stage, and treatment temperature and duration (Wills et
from combinations of T. paradoxa, L. theobromae, C. mu- al. 1998).
sae and D. torulosa (Sommer et al. 1992). The disease is MA storage has been used for several decades for
characterized by darkening of the hand and adjacent marine shipment of bananas (Scott and Roberts 1966;
peduncle and loss of the ability of the hand to support the Woodruff 1969). Storage of bananas under the MA condi-
fruit (Sommer et al. 1992). Specific visual symptoms of this tion is primarily achieved using low-density polyethylene
disease include blackening of tissues at the cut crown sur- (LPDE) bags (Truter and Combrink 1990; Stiles 1991; Cha-
faces and a spreading grayish-white, pink or white mold on mara et al. 2000). Green mature ‘Cavendish’ bananas may
the cut crown surface (Ploetz 1998). Banana leaves, flowers, be stored in LDPE bags (0.05 mm thickness) for up to 30
bracts, and transitional leaves are commonly colonized by days at 14°C with ripening and sensory quality during shelf
the causal fungi. When the banana hands are harvested in not being adversely affected (Hewage et al. 1995). Poly-
the field, latex flows from the cut surface of the crown and vinyl chloride film (0.01 mm thickness) packaging also pro-
spores of the fungi may enter the wound and initiate disease longed the shelf life of ‘Sucrier’ banana fruit at peel stage 3
development. Once initiated, infection can progress from (more green than yellow) to 6-7 days at 20°C, compared
the crown into the pedicels and eventually into the fingers with 3-4 days in the control (Choehom et al. 2004; Rom-
(Krauss and Johanson 2000). phophak et al. 2004). Longevity may be extended using an
ethylene absorber, such as potassium permanganate in com-
Anthracnose bination with polymer films (Scott et al. 1970; Jiang et al.
1999a; Chamara et al. 2000). Ketsa et al. (2000) reported
Anthracnose is another important postharvest disease of ba- that bulk packaged ‘Sucrier’ bananas stored in non-perfo-
nana fruit that occurs in all producing areas. This disease is rated polyethylene (PE) bags with an ethylene absorbent
caused by Colletotrichum musae, which infects both green and carbon dioxide scrubber remained green for 6 weeks at
and ripe fruit. However, the symptom becomes evident as 14°C. Thus, MA has repeatedly been shown to be beneficial
the fruit ripen, especially in wounds (Ploetz 1998). for long-term storage of banana fruit at low temperature.
C. musae typically establishes a subcuticular latent in- Apart from conventional MA packaging for bananas, Ste-
fection. Both flower parts and the last bunch (proximal) mart et al. (2005) evaluated the potential of passive silicone
bract are potential sources of C. musae inoculum. Conidia membrane and diffusion channel systems to preserve the
of C. musae contaminate bananas during the month after quality and extend the shelf life of ‘Cavendish’ banana fruit.
flowering and may spread in rainwater trickling over the Banana fruit could be stored for 42 days at 15°C under MA
bunch. Conidia quickly germinate and may form a mela- conditions achieved using these novel systems.
nized appressorium (Muirhead and Deverall 1981; De La- Controlled atmospheres of 2-5% O2 and 2-5% CO2 are

145
 Fresh Produce 1(2), 140-152 ©2007 Global Science Books

considered effective for delaying banana ripening and 2.0

Ethylene production rate

reducing respiration and the effects of ethylene (Kader A
1997). The storage life of green banana fruit was up to 180
days at 20°C when they were ventilated continuously with 1.5

(Pl kg h )
-1
an atmosphere of 3% O2, 5% CO2 and 92% N2 (Wills et al.

-1
1998). However, care must be taken to ensure that sufficient
O2 and not too much CO2 is retained in the atmosphere. 1.0
Exposure of banana fruit to less than 1% O2 and/or more
than 7% CO2 may undesirably affect texture and flavour
(Kader 1997). Application of CA to delay ripening during .5
transport has facilitated the picking of bananas at the fully
green mature stage. 0.0
Coatings
70
Fruit coatings can modify internal fruit atmospheres and re- B
duce transpiration like MA films and thereby reduce respi- 60

(ml CO2 kg-1 h-1)

ratory activity and water loss without adversely affecting

Respiration rate
50
fruit taste. Kittur et al. (2001) investigated that the effects of
four different polysaccharide-based composite coating for- 40
mulations (chitosan, N,O-carboxymethyl chitosan, carboxy-
methyl derivatives of cellulose and starch, and hydroxyl- 30
methyl starch and hydroxypropyl starch) on banana quality
maintenance. The fruit treated with polysaccharide-based 20 control
coatings had retarded colour development, lower acidity 9 h N2
and greater firmness compared to the control. CO2 evolu- 10
tion and loss in weight were also reduced significantly. 0
Coating ‘Drawf Cavendish’ banana fruit with ‘PRO-LONG’
inhibited ethylene production in associated with reduction 0 5 10 15 20 25
of ACO activity under restricted O2 and thereby delayed Storage time (d)
fruit ripening (Dillon et al. 1989; Zhang et al. 1996). The
combination of calcium chloride infiltration with ‘Semper- Fig. 1 Changes in the rates of C2H4 production (A) and respiration (B)
fresh’ coating achieved more efficient effects than the coa- during storage of 9 h N2-treated and control ‘Brazil’ banana fruit. Each
ting alone (Chukwu et al. 1995). Coatings may also form a value is the mean ± standard error (n = 3). Vertical bars indicate the stan-
physical barrier against pathogenic infection, thereby redu- dard errors of the means where they exceed the symbol size (data from Yi
cing postharvest disease incidence (Ben-Yehoshua 1966; et al. 2006).
Amarante and Banks 2001).

Anoxia or low oxygen

1-Methylcyclopropene (1-MCP)
Anaerobic conditions may occur somewhat during post-
harvest handling, storage and transport of fruit, for example, The ethylene antagonist 1-MCP binds irreversibly to ethy-
during CA or MA storage, or after coating with various lene receptors in plant cells and prevents the ethylene mole-
waxes. Anaerobiosis can result in off-flavour and generally cule from binding. 1-MCP thereby inhibits ethylene signal
poor fruit quality. When bananas are held in sealed poly- transduction and downstream action, effectively delaying
ethylene bags for a relatively long time, a green ripe dis- ripening and senescence of ethylene sensitive fruits and
order may occur. This disorder is characterized by green re- vegetables. This gaseous compound has been approved for
maining in the skin along with pulp softening and off-fla- use commercially as a postharvest treatment for a range of
vour development (Satyan et al. 1992). However, short- climacteric fruits. Exposure to 1-MCP increases the longe-
term anaerobic conditions might be beneficial for posthar- vity of harvested banana fruit (Fig. 2). However, the res-
vest fruit quality under certain circumstances. Wills et al. ponse is dependent upon fruit maturity, treatment tempera-
(1982) reported that exposure of preclimacteric banana fruit
to low O2 for 2-3 days prior to storage in air extended the 45
time required for the fruit to ripen. 2% O2 stress for 48 h
was effective in preventing decay after shelf life. This low 40
O2 treatment also retarded ‘Ziv’ banana ripening processes
(viz. colour, firmness, respiration and ethylene production) 35
Shelf life (d)

and reduced CI symptoms, without impairing the taste 30

(Pesis et al. 2001). Exposure of banana fruit to <1% O2 or
100% N2 resulted in a significant reduction in the formation 25
of ethyl acetate, an important volatile compound (Wenda-
koon et al. 2004). Yi et al. (2006) found that exposure of 20 + Ethylene
‘Brazil’ banana fruit to pure N2 gas for 9 h reduced the rates 15 - Ethylene
of ethylene production and respiration, as well as the acti-
vities of polygalacturonase and pectin methyl esterase 10
during storage, and thereby effectively inhibited fruit ripen- 5
ing (Fig. 1). The ripening retardation may be related to
induction of acetaldehyde and ethanol production, which 0.0 .2 .4 .6 .8 1.0 1.2
could in turn inhibit ethylene production and action. This
process is known to alleviate CI of harvested fruits and 1-MCP concentration (Pl l-1)
vegetables (Duan et al. 2003; Pesis 2005). However, anae- Fig. 2 Changes in the shelf life at 20°C of harvested ‘Cavendish’ banana
robic treatments should be applied carefully and optimised fruit treated with 1-MCP for 24 h at various concentrations without (-) or
for individual species and temperature regimes. with (+) subsequent ethephon treatment. Each value is the mean for nine
fruit, and vertical bars indicate the standard error (data from Jiang et al.
1999a).

146
 Postharvest biology and handling of banana fruit. Duan et al.

ture and duration, and 1-MCP concentration (Jiang et al. Nitric oxide (NO) and nitrous oxide (N2O)
1999b; Harris et al. 2000; Bagnato et al. 2003; Pelayo et al.
2003). It is now well established that 1-MCP treatment N2O is a naturally occurring atmospheric gas, the primary
lowers ethylene production and respiration rates (Golding et source of which is soil containing aerobic denitrifying bac-
al. 1998, 1999; Pathak et al. 2003; Pelayo et al. 2003; Lo- teria (Firestone and Davidson 1989). NO is synthesized in
hani et al. 2004) and inhibits softening (Jiang et al. 1999a, animals, plants and microorganisms (Crawford 2006). Re-
1999b; Macnish et al. 2000; Pelayo et al. 2003; Lohani et al. cently, N2O and NO have been shown to have anti-senes-
2004) of harvested banana fruit. Colour changes are also cence and ripening properties. N2O and NO treatments have
delayed in 1-MCP treated banana fruit. A potential concern been found to extend the storage and marketing life of a
is that yellowing of banana fruit can be disrupted (i.e. in- range of fruits, vegetables and flowers (Gouble et al. 1995;
complete and uneven), even in the presence of the ethylene Bowyer et al. 2003; Badiyan et al. 2004; Soegiarto and
analogue propylene (Golding et al. 1998; Harris et al. 2000; Wills 2004).
Macnish et al. 2000). Moreover, total volatile production of Palomer et al. (2005) reported that ‘Cavendish’ banana
fruit may be inhibited by 1-MCP treatment. Quantitatively, fruit ripening was significantly delayed by N2O within the
ester concentrations were lower, while concentrations of al- concentration range from 20 to 80%. Delayed ripening was
cohols were higher in treated fruit (Golding et al. 1998). judged by effects on both ethylene synthesis and respiration
The sugar content was not affected by 1-MCP treatment rate in association with changes in fruit colour, acidity and
(Golding et al. 1998). 1-MCP treatment may increase the softening. This response to N2O was dose- and time-
susceptibility to CI of banana fruit. Jiang et al. (2004) con- dependent. Combinations of N2O with low O2 (8 and 12%)
cluded that the development of CI was associated with de- in controlled atmospheres had a synergistic effect on the
creased ethylene binding. ripening-delay capacity of N2O. The ability of N2O to slow
As noted, the action of 1-MCP is mediated through down fruit ripening is thought to be due to inhibition of
interacting with receptors and competing with ethylene for ethylene synthesis and action (Gouble et al. 1995).
these binding sites (Sisler and Serek 2003). The ripening
response of fruit treated with 1-MCP and subsequently trea-
ted with ethylene varies with the interval of time between 1- 8
MCP and ethylene treatments. As the time lag is increased, Ethylene production rate Control
new binding sites are synthesised and delayed ripening ef- SNP treatment
fect weakens (Jiang et al. 1999b). Synthesis of new binding 6
(Pl kg h )
sites can be affected by temperature. Temperature between
-1 -1

30 and 40°C result in faster recovery of ‘Williams’ banana

fruit ripening capacity. Also based on the temperature action, 4
application of 1-MCP at 2.5°C is less effective than at 15
and 20°C. This observation suggests that binding of 1-MCP
at low temperatures was incomplete (Jiang et al. 2002, 2
2004).
ACO and ACS are the rate-controlled enzymes of the
ethylene biosynthetic pathway. In banana fruit, inhibition of 0
ethylene production by 1-MCP was associated with both
lower expression and lower activities of ACO and ACS 12
(Pathak et al. 2003; Zhang et al. 2006). In addition, delayed
softening in 1-MCP treated banana is related to lower ex- 10
Firmness (kg cm )
-2

pression of an ethylene induced expansin (Maexp1) gene

(Trivedi and Nath 2004) and lower activities of pectin me- 8
thylesterase, polygalacturonase, endo--1,4-glucanase and
pectate lyase (Lohani et al. 2004). 6
Plant growth regulators 4
On the basis of ethylene evolution and respiration, it was 2
found that ripening of banana fruit was hastened by ABA
and 2,4-dichlorophenoxy acetic acid and delayed by IAA
0
and GA treatments (Pathak and Sanwal 1999; Jiang et al.
2000; Lohani et al. 2004). Climacteric respiration of ‘Nani- 14
Chlorophyll content (Pg cm )
-2

cao’ banana was reduced and starch degradation and suc-

rose formation were delayed by the action of IAA. However, 12
SuSy and SPS activities and transcript levels were not af- 10
fected, the increase in the activity and transcript level of E-
amylase was delayed by IAA treatment. Thus, prevention of 8
sucrose accumulation by IAA was not related to sucrose-
metabolizing enzymes, but was apparently a consequence 6
of lack of substrate due to inhibition of starch degradation
(Purgatto et al. 2001). GA delayed sucrose synthesis of ‘Na- 4
nicao’ banana by disturbance of sucrose-phosphate synthe-
sis, but it had no effect on sucrose synthase (Rosecler et al. 2
2003). Application of the secondary messenger compound
salicylic acid (SA) at 500 Pmol/l and 1000 Pmol/l also de- 0
layed ripening of ‘HariChhal’ banana fruit in association 0 4 8 10 11
with decreases in respiration rate, fruit softening and the ac-
tivities of major cell wall degrading enzymes (Srivastava Storage time (d)
and Dwivedi 2000). Fig. 3 Effect of the NO donor, sodium nitroprusside (SNP) on ethylene
production rate (A), firmness (B), and chlorophyll content (C) of ‘Brazil’
banana fruit during storage at 28qC. SNP was used in 5 mM concentration.
Each data point represents a mean ± standard error (n = 3) (data from
Duan et al., unpublished).

147
 Fresh Produce 1(2), 140-152 ©2007 Global Science Books

Evidence of interplay between NO and ethylene in the a standard benomyl (0.1%) fungicide treatment. Cymbopo-
maturation and senescence of fruit suggests an antagonistic gon nardus and Ocimum basilicum oils also had fungicidal
effect of these gases. Unripe green banana fruit contain high activity against C. musae and F. proliferatum when tested at
NO and low ethylene concentrations. The maturation pro- between 0.2-0.6% (v/v) in a poisoned food bioassay (An-
cess is accompanied by a marked decrease in NO conco- thony et al. 2004). Application of Ocimum basilicum essen-
mitant with an increase in ethylene (Leshem et al. 1998). tial oils (0.16% v/v) effectively suppressed crown rot and
Certainly, NO treatment can significantly decrease ethylene anthracnose diseases, enabling ‘Embul’ banana fruit to be
biosynthesis, delay pulp softening and peel degreening, and stored for up to 21 days at 13.5 ± 1°C without any detri-
prolong the shelf life of ‘Brazil’ banana fruit (Fig. 3). mental effect on their organoleptic properties. The efficacy
was comparable to treatment with benomyl (Anthony et al.
Control of postharvest disease 2003).

Chemicals Heat treatment

Control of most postharvest diseases in banana fruit is Hot water treatment (HWT) is an effective non-chemical
through application of fungicides as a dip or spray (Wills et method of postharvest pest and disease control if combina-
al. 1998; Krauss and Johanson 2000). In the past, the most tions of suitable temperatures and exposure times are selec-
commonly used fungicides were benzimidazoles, such as ted that prevent the loss of produce quality (Lurie 1998).
benomyl and thiabendazole (TBZ). This class of fungicides HWT has the potential to replace chemical fungicides to
inhibits spore germination, interferes with mycelial growth control crown rot of banana. HWT at 45°C for 20 min re-
and affects conidia formation by interrupting polymeriza- duced crown rot of ‘Santa Catarina Prata’ and ‘Williams’
tion of the tubulin protein (Wills et al. 1998). However, C. banana fruits inoculated with Chalara paradoxa spore sus-
musae has developed resistance to these fungicides (de La- pension from 100 to less than 15%. When fruit were ex-
peyre de Bellaire and Dubois 1997a). When control breaks posed to hot water at 50°C for 20 min, crown rot was re-
down, trizaoles such as imazalil and prochloraz may be duced to < 3% (Reyes et al. 1998). Hassan et al. (2004)
used, usually alternated with TBZ. Trizaoles inhibit deme- found that disease severity in banana fruit was significantly
thylation processes during ergosterol biosynthesis within reduced by HWT (50 ± 2°C for 5 min) and fungicide ap-
the fungus (Wills et al. 1998; Krauss and Johanson 2000; plication. Similarly, combining HWT with a bacterial anta-
Khan et al. 2001). gonist gave more effective control of anthracnose, crown
Considering potential adverse environmental and health rot and blossom end rot of ‘Emon’ and ‘Kolikuttu’ bananas
effects and also resistance development by pathogens to than using the two treatments individually (de Costa and
fungicides, it is desirable to develop alternatives to conven- Erabadupitiya 2005).
tional fungicides. Alvindia et al. (2004) reported that spore
germination of L. theobromae, T. paradoxa, C. musae, C. Biological control
gloeosporioides, F. verticillioides, and F. oxysporum was
completely inhibited by NaClO at 5 g/l and each of Biological control antagonists can be used as a single ap-
NaHCO3 and CaCl2 at 6 g/l. Dipping banana fruit for 10-15 plication using existing delivery systems (e.g. drenches, line
min in these concentrations reduced the incidence of crown sprayers and on-line dips) and can significantly reduce fruit
rot compared with the untreated fruits. Incidence at 17 days decay (Janisiewicz and Korsten 2002). Moreover, biocon-
after harvest was reduced with NaClO by 67%, NaHCO3 by trol agents may be applied directly to the targeted area (e.g.
62%, and CaCl2 by 33%. Postharvest acid (acetic acid and fruit wounds). Biocontrol systems for reducing decay have
citric acid) treatments may increase ‘Embul’ banana fruit re- been successfully established for pome and citrus fruits (Ja-
sistance to anthracnose, thereby reducing the benomyl con- nisiewicz and Korsten 2002).
centration needed to control this disease when acid and fun- For banana fruit, Brown and Swinburne (1980) found
gicide are combined (Perera and Karunaratne 2001). that culture filtrates or cell wall fragments of C. musae can
induce production of antifungal components in the peel of
Natural extracts green banana fruit, which inhibited conidial germination of
C. musae on the treated skin. Several mycoparasites of the
Plant extracts are emerging as alternatives to conventional crown rot complex and one antagonistic bacterium were
fungicides for the control of plant disease. They are gene- identified by Krauss et al. (1998). Some of these attacked
rally regarded as safe (GRAS) to humans and environmen- the whole range of fungi involved in this disease complex,
tally friendly. Some plant extracts have been shown to ef- including structures considered relatively inaccessible to
fectively control various plant diseases (Wilson et al. 1997; fungicidal attack (i.e. conidia and haustoria). Other control
Sarma et al. 1999; Bowers and Locke 2000). organisms showed tolerance to fungicides and thus could be
There are examples of successful control of postharvest combined with reduced concentrations of fungicide in an
banana diseases with plant extracts. Ranasinghe et al. integrated disease management system (Krauss and Johan-
(2002) reported that cinnamon and clove essential oils were son 2000). Furthermore, Krauss et al. (2001) suggested the
effective in vitro at low concentration against pathogenic or- existence of mycoparasite discrimination between different
ganisms isolated from banana, including C. musae, L. theo- strains of C. musae, the principal pathogen. The discrimina-
bromae and F. proliferatum. The major constituent of cin- tion was thought to be associated with different mechanisms
namon bark oil was cinnamaldehyde. Furthermore, treat- of action by different mycoparasites; viz. parasitism, antibi-
ments with cinnamon bark and leaf oils controlled crown osis, competition. Accordingly, combinations of different
rot. However, clove oil treatment did not affect develop- mycoparasite strains belonging to different species en-
ment of this disease. Treatment with emulsions of cinnamon hanced biocontrol efficacy against mixed infection.
oils combined with MA packaging can be synergistically A member of the Burkholderia cepacia complex, iso-
effective for extending the storage life of ‘Embul’ banana lated from the fructosphere of banana, has been shown to be
fruit. This combination gave a storage lives of up to 21 days effective as an antagonist of postharvest pathogens even
in a cold room and 14 days at 28 ± 2ºC without adversely after 5 years of storage in sterile distilled water at ambient
affecting the organoleptic and physico-chemical properties temperature. The most effective concentration of B. cepacia
of the fruit (Ranasinghe et al. 2005). Thangavelu et al. was determined to be 1010 CFU/ml for in vivo control of
(2004) noted that extracts of S. torvum at 25 and 50% con- anthracnose and crown rot (de Costa and Subasinghe 1998;
centration (w/v) completely inhibited mycelial growth of C. de Costa and Erabadupitiya 2005). Taechowisan and Lumy-
musae. The same extracts were found effective in reducing ong (2003) isolated a novel endophytic actinomycete from
the incidence of anthracnose disease on the three banana the root tissues of Zingiber officinale. The Streptomyces au-
cultivars (‘Robusta’, ‘Rasthali’, ‘Ney Poovan’) compared to reofaciens CMUAc130 isolate was determined to have

148
 Postharvest biology and handling of banana fruit. Duan et al.

potential against phytopathogenic fungi. Furthermore, both for the most part, these approaches have not yet been adop-
the culture filtrate and crude extract from this strain had ted commercially. The key to adoption may lay in develop-
inhibitory effects against C. musae from banana fruit. The ing high efficacy integrated disease management programs.
major active ingredients from the culture filtrate of S. au- In the future, it is expected that the use of novel approaches
reofaciens CMUAc130 were identified as 5,7-dimethoxy-4- will accelerate as more stringent international standards and
p-methoxylphenylcoumarin and 5,7-dimethoxy-4-phenyl- requirements concerning conventional fungicides becomes
coumarin (Taechowisan et al. 2005). Gunasinghe et al. the global norm.
(2004) reported that two local isolates of the biocontrol
agents Flavobacterium sp. W5481 and Pantoea agglome- ACKNOWLEDGEMENTS
rans W5482 reduced crown rot development on ‘Embul’
banana hands and, therefore, showed potential for the bio- Financial support from the National Natural Science Foundation of
control of banana pathogens. China (Grant Nos. 30500353 and 30430490), the International
Foundation for Science (Grant No. E/3656-1), the Natural Science
CONCLUSIONS Foundation of Guangdong Province (Grant No. 06200670 and
5300902) and the Science Foundation of South China Botanical
The published literature on mechanisms of quality mainte- Garden, CAS (Grant No. 2005-3357) is greatly appreciated.
nance of harvested fruit, including bananas is extensive.
Nonetheless, our understanding is still incomplete. Further REFERENCES
progress will be associated with even better understanding
of the postharvest biology of banana fruit and more compre- Adams-Phillips L, Barry C, Giovannoni J (2004) Signal transduction systems
hensive knowledge of factors that govern major rates of regulating fruit ripening. Trends in Plant Science 9, 331-338
quality maintenance. Particularly important advances have Ahmad S, Chatha ZA, Nasir MA, Aziz A, Mohan M (2006) Effect of humi-
dity on the ripening behaviour and quality of ripe fruit. Journal of Agricultu-
been made over the past decade or so in the field of ethy-
ral Society for Science 12, 54-57
lene perception, synthesis and transduction pathways (Sisler Ahmas S, Clarke B, Thompson AK (2001) Banana harvest maturity and fruit
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