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Postharvest Physiology of Tropical Fruit

Jingtair Siriphanich
Department of Horticulture
Kasetsart University, Kampangsaen campus,
Nakhon Pathom 73140, Thailand
Keywords: Postharvest physiology, peel, durian, mango, mangosteen

Basic postharvest physiology of tropical fruit is similar to that of temperate
fruit. The only clear physiological difference is that tropical fruit are chilling
sensitive while most temperate fruit are chilling insensitive. Most of this knowledge
was derived from work with banana, mango, pineapple and papaya. However, little
work on chilling physiology has been reported in recent years. For other tropical
fruit, the studies have emphasized handling and storage, less so physiology. Lack of
physiological research will limit the expansion of tropical fruit in the world market.
Respiration and ethylene production rates are largely known but there is a need for
knowledge on sensitivity to ethylene is to improve postharvest handling and
management of these fruit. Furthermore, individual tropical fruit species are
unique and require detailed study. For example, durian fruit suffer from uneven
ripening. Basic work on growth and development of durian led to the suggestion
that the problem stemmed from the preharvest uneven development of the fruit
itself. Mangosteen fruit are affected by husk hardening and a translucent pulp
disorder. Studies on these two problems led to new knowledge on the biochemistry
of fruit texture.

There are numerous species of tropical fruit, but information on their
physiological behavior after harvest is quite limited, except for banana, mango and
pineapple. This paper reviews briefly the general postharvest physiology, and progress in
chilling injury research. It describes in more detail the postharvest physiology of the peel
and of individual fruit that behave atypically.


Once harvested, tropical fruit behave similarly to subtropical and temperate fruit.
They transpire and wilt, respire and lose food reserves, many produce ethylene and ripen,
senesce and become susceptible to microbial attack and finally die. Their ripening
processes are also similar and include color changes, usually from green to yellow or red,
loss of chlorophyll and the production of carotenoids and/or anthocyanins, textural
change, generally from firm to soft, due to the solubilization and degradation of pectin,
and changes in flavour, normally from astringent and tart to sweet due to the alteration of
their chemical composition. These changes included: conversion of starch to sugars,
although not all fruit accumulate starch during their development; an increase or decrease
in sugar content , and altered proportions of sucrose, fructose and glucose. Acid content
almost always declines except later in the senescence process in some fruit such as
banana, phenolic content decreases in association with the decline in astringency
presumably due to the polymerization of phenolic compounds, as reported in temperate
fruits and new volatiles are synthesized, giving the characteristic aroma of ripe fruit.
Although the patterns of change are similar among different fruit, the quantity, timing,
and chemical species are quite distinct. Soluble solids and tritatable acid contents of
some tropical fruits are shown in Table 1.
Respiration and ethylene production rates are largely known in most fruit, but
there are some differences among cultivars although the data were usually collected
under different conditions (Table 1). There are those with low rates comparable to many

Proc. IS on Trop. & Subtrop. Fruits

Ed. R. Drew 623
Acta Hort. 575, ISHS 2002
temperate fruit while others have very higher rates not seen in subtropical and temperate
fruit. Their patterns of respiration can also be divided into the climacteric and non-
climacteric types while some are still uncertain. Among the climacteric fruit the peak of
respiration can occur before, after or coincide with the peak of ethylene production
(Paull, 1993). This knowledge is important for commercial application such as for the
calculation of heat loads and cooling capacity, and the management of mixed loads.
For most tropical fruit, except those already well established in the world market
such as banana and pineapple, knowledge of their sensitivity to ethylene, such as that
reported by MaLauchlan et al. (1994), is essential. First, because they must be stored at
relatively higher temperature to avoid chilling injury. At these temperatures ripening
usually proceeds rapidly and is intensifed by ethylene. Second, because mixed loads are
often required to export these fruit into new markets where demand is not yet high or
consistent supplies of good quality produce cannot be obtained. Ethylene concentrations
must be kept low during shipment to ensure that the produce arrive at their destination in
optimum condition, without damaging other produce or requiring additional ripening
treatment. Unfortunately, knowledge on the sensitivity of tropical fruit to ethylene is
almost nil.

This physiological disorder is the most important obstacle to the expansion of
trade in tropical fruit in the world market. Despite the importance of this problem,
progress has been slow. There are more than 700 articles involving at least one aspect of
chilling injury recorded in the CAB abstracts from 1990 to early 2000. Only a quarter of
these articles were on physiology. Amongst these, less than ten were on tropical fruit.
Most were articles describing the symptoms or conditions under which the injury
developed or was alleviated.
Even at the present time the majority of the research still emphasizes the
prevention of injury by using various techniques. One approach involves reduced oxygen
concentrations during storage and handling by application of controlled atmospheres,
wrapping and surface coatings (Wang, 1993). Although this approach is widely used
commercially, eg. for the shipment of pineapples to distant markets, it is viewed as a
procedure to prevent symptom development rather than to prevent the injury or the
primary event itself. Other approaches involve temperature management (Wang, 1993)
including intermittent warming during storage, presumably to reverse the primary event
and eliminate toxic materials accumulated at chilling temperatures and heat treatment
before storage to acclimate the fruit to low temperature. The latter approach is rather
difficult to apply and commercial use is quite limited.
A newer approach is being developed for pineapple, probably because it is the
only commodity currently shipped at chilling temperatures. The current research
(Graham et al., 1998) uses a molecular approach aimed at manipulating the activity of
polyphenoloxidase, which is believed to be responsible for the development of brown
pigment visible after removal of pineapples to warm temperatures. Another molecular
approach that could also alleviate the injury is to enhance the production of ascorbic acid,
which is known to prevent internal browning if present in high concentrations in the pulp
of pineapples. However, this molecular approach can also be considered to merely
prevent the development of the symptoms, not the injury. If the enzyme(s) can be
controlled and browning is prevented, the injury still remains. Translucent areas that are
normally seen at the early stages of chilling injury in pineapples may still be observed.
A better approach should aim at preventing the injury itself. Unfortunately, the
mechanism of the injury especially the primary event when temperature drops into the
chilling range is not well understood. The hypothesis on lipid phase transition proposed
by Lyons (1973), and later modified by Raison and Lyons (1986) by redefining terms,
has not yet proved or disproved. A newer hypothesis was proposed in 1991 (Fig. 1).
According to this hypothesis membrane are also involved in the development of the
injury but are regarded not as a primary event but rather as a secondary event (Shewfelt,

2000). It was proposed that chilling stress induced reactive oxygen species. These active
oxygen species damage membranes and other molecules at a subcellular level. As a result
membranes breakdown causing further damage to cells and finally leading to the
appearance of symptoms. However, it is hard to comprehend how chilling temperatures
can induce the reactive oxygen species directly. Shewfelt also suggested that antioxidants
could act as a defense mechanism to prevent injury by destroying the reactive oxygen
species. These antioxidant species could be nonenzymic such as -carotene, lycopene, -
tocopherol, ascorbic acid and glutathione or enzymic such as catalase, peroxidase and
superoxide dismutase. Data reported by Sala and Lafuente (2000) supported this
hypothesis by showing that heat-conditioning reduced chilling injury in mandarins and
induced catalase activity. However, heat shock protein and polyamines, often found after
heat treatments, have not yet been shown to be associated with the proposed oxidative
process. Much more work is still needed especially with tropical fruit.


Many tropical fruit have a distinct peel that cannot be consumed and must be
separated from the pulp. Among the sixteen fruit listed in Table 1, half are those with
peel. Peel physiology is usually different to that of the pulp. Its physiology should be
studied seriously. Take banana as an example, most people thought that when banana
ripens the peel turns from green to yellow and coincides with the development of the
pulp, where starch is converted to sugar, cell walls are degraded and aroma volatiles are
produced. However, this is not the nature of all bananas. Cavendish, one of the main
cultivars in the world market, will ripen while still largely green if held under its natural
conditions. Lower temperatures are required during ripening to achieve degreening with
optimum eating quality (Lizada et al., 1990).
An contrary observation was made in mango. Certain cultivars, such as the Nam-
dok-mai cultivar in Thailand, do not develop full yellow or red color when ripened
naturally or even when treated with 1,000 ppm ethylene (Table 2). Lowering temperature
to about 20 C as recommended by Kader and Mitcham (1994) does not promote enough
degreening (Sangvanangkul, 1997). A bright yellow Nam-dok-mai mango may be
achieved by ripening with acetylene generated from calcium carbide at warmer than the
ambient temperatures. However, the exact conditions have not been determined.
Browning at the stylar end or black tip was often found under these conditions (Mitra and
Baldwin, 1997). It is worth studying the degradation of chlorophyll in mango peel.
In durian, which is a climacteric fruit, a study on respiration and ethylene
production of the husk and the pulp clearly showed that most carbon dioxide and
ethylene is evolved by the husk and only a fifth by the pulp (Siriphanich et al.,1994a).
Many attempts to ripen durian pulp separated from the husk have not been successful,
even with high concentrations of ethylene. Durian husk normally dehisces at the fully
ripe stage when the pulp is very soft and has a strong odor. Sriyook et al. (1994) showed
that the dehiscence process was induced by ethylene and physically promoted by the loss
of water from the peel (Fig. 2).
In mangosteen, the thick peel protects the pulp very well , but the peel itself is
very susceptible to mechanical damage. However, the response of mangosteen peel is
opposite to that found in other fruit. If the fruit is dropped on a hard floor the peel turns
hard in a few hours (Fig. 3). Further observations reveal that the hardening also
developed when mangosteen was stored at 4C. In addition, microbial attack also shows
the same response. The hardening process depends on oxygen indicating a enzymatic
reaction once the cells in the peel are damaged. Phenolic compounds decrease in the
affected area, while lignin accumulation was localized in the affected areas (Ketsa and
Attanee, 1998).
When mangosteen ripens the peel turns from green to patchy pink then pink, red,
purple, and finally black. The pulp is eaten during the purple and black stage. The rate of
color change is rapid. Normally it takes only 2 days after harvest at the pink stage to the
black stage at room temperature. This means that mangosteen on display shelves are

almost black and not very attractive to consumers. A detailed study into the changes in
peel pigment is lacking. Understanding pigment changes in mangosteen peel could lead
to a better understanding and a procedure to prolong the beautiful color of this fruit.
The attractive peel of rambutan fruit its contains several hundred spinterns that
are very susceptible to water loss and turn brown easily. Studies have shown that the
browning was due to both changes in anthocyanins as well as the development of brown
pigments (Landrigan et al.,1996). It is interesting to note that most of the commodities
that have hairs or trichomes on their surface usually develop skin browning, such as
lanzone, langsat as well as okra and vegetable soybean. It is obvious that these
commodities have a high surface area and are prone to water loss. However, why should
water loss cause browning in these commodities but not in many others under severe
water loss? The answer awaits further study.
Because of the natural separation between the peel and the pulp in many fruits,
the pulp also contains its own skin, usually as an epidermal tissue with a thick outer wall
eg. pummelo (Charoensub and Siriphanich, 1988). To some extent this epidermal tissue
may protect the pulp from microbes and water loss. Nevertheless, special care must be
taken to prevent damage to this natural skin layer when preparing minimally processed
fruit, even though the pulp of tropical fruit can normally be stored at a lower temperature
than the whole fruit. Temperature during handling and storage of minimally processed
tropical fruit must also be well managed to avoid contamination with pathogens and
subsequent catastrophe.

Durian is a minor fruit crop in the world market, but is quite important in South
East Asia. It is now gaining popularity in China and Taiwan. Thailand is the main
producing and exporting country. The export volume from Thailand has increased almost
10-fold from 15,600 tons in 1989 to 132,000 tons in 1999. However, the value that
farmers gain per kilogram is decreasing, due to over production and a limited market.
The expansion of this fruit to markets in other parts of the world is limited by the nature
of this fruit.
Being a tropical fruit, a mature but unripe durian suffers from chilling injury at
temperatures below 15 C. The ripe fruit can be kept at a lower temperature, while the
pulp can be stored at 4C for a month (Siriphanich et al., 1994b). The older
recommended storage temperature of 4C (Pantastico et al., 1975) was probably for the
fully ripe fruit. Physiological studies on durian are quite limited as in other tropical fruit.
We know that it is a climacteric fruit but do not know much about its changes during
ripening. The following is some physiological information gained in the past few years.
Besides chilling injury, durian also suffers from uneven ripening. Few studies
have focused on this problem. However, our study on the growth and development of
Monthong durian revealed that during pulp development, starch accumulation among
various locules in a fruit is not uniform. Even in the same locule or on the aril of the
same seed, starch accumulation was not evenly distributed, as shown by I2-KI staining in
Figure 4. Monthong, which is late cultivar with the highest pulp portion of about 50 % of
the whole fruit weight, suffers more from this problem than other earlier cultivars. In
Kradum, the earliest cultivar with a pulp portion of only about 25 %, ripens evenly.
Ethylene production in durian is low compared to other fruit and the pulp itself produces
very little ethylene (Siriphanich et al., 1994a). All this information suggests that the
uneven ripening problem in durian stems from a combination of factors starting with fruit
development and continuing into ripening. Being a large fruit with a high proportion of
storage tissue, the fruit forms a very large sink, which the leaves are unable to supply fast
enough to ensure that all locules can reach maturity at the same time. The uneven
ripening problem may become worse if the fruit is harvested immature. Once harvested,
if left to ripen naturally ethylene produced internally may not be distributed evenly to all
parts of the pulp, resulting in uneven ripening. This problem can be solved if some of the
fruit are thinned during their development, harvested at a mature stage and ripened with

exogenously ethylene.
Being large, durian poses another interesting phenomenon. Its respiration rate is
quite high. We measured rates up to 700 mg/kg-hr at 29C. The respiratory heat cannot
be easily released to the atmosphere. Normally the internal temperature is about 0.5 to
1.0 C higher that the surrounding atmosphere. Once ripened the internal temperature
could be up to 4 degrees higher. Those who handle durian must take this information into
account in designing a cooling procedure for the fruit. Cartons for holding durian must
have extra venting to aid heat exchange. Air circulation during cooling and storage must
be high enough to keep the entire durian fruit cool. This information also explains the
finding by Ketsa and Pangkool (1995) who showed that durian should be ripened at
relatively low humidity of about 70 %. At higher relative humidity, heat produced by the
respiration process might not be released fast enough, due to evaporative cooling effect at
lower relative humidity. Hence, ripening at higher relative humidity caused rapid
changes in the pulp and resulting in less satisfactorily pulp quality. However, it should be
kept in mind that consumer preferences for flavour and texture of durian differ among
ethnic groups or even among individuals. Consumers other than the Thais may prefer
durian ripened under higher relative humidity.

Mangosteen is a climacteric fruit. After harvest it has quite a long shelf life of
more than 10 days if not attacked by fungi. There is little change in the pulp after harvest
although soluble solids increase slightly and titratable acidity decreases (Dangcham,
Besides the problem of peel hardening in response to many external stresses, the
pulp also has two serious disorders. One is the translucent pulp disorder in which the
whitish pulp becomes translucent and three times as firm as normal. The other disorder is
gummosis, in which yellow latex is observed in the pulp as well as the peel. A recent
study showed that the translucent disorder was associated with the amount of rainfall and
irrigation (Laywisadkul, 1994) during the harvesting period. It was shown that
intercellular spaces in the normal pulp were filled with water. Soluble solids and
titratable acidity were lower than normal. About 30% of the cells in the translucent pulp
were damaged. Under these conditions ethylene production was limited, while carbon
dioxide production was enhanced, indicating an anaerobic condition. In addition, the
normal change in cell wall pectin from insoluble to soluble during ripening was
disrupted. A reverse process in cell wall pectin was observed. The water soluble pectin
fraction was converted into a NaCO3 soluble fraction. This was probably due to the
leakage of acids from the cytosol of damaged cells into the cell walls (Luckanatinvong,
1996). Further study showed that excess water that is associated with the disorder did not
come via the roots, because when water was forced directly into the fruit, the peel
cracked but no translucent disorder was observed. In addition, when comparing different
irrigation treatments for mangosteen plants either by sprinkler under the canopy or above
the canopy with or without covering the ground with Tyvex cloth (Luckanatinavong et al,
2000), it was found that more translucent fruit were recorded in the above-the-canopy
treatment than in the other treatments (Table 3).
The gummosis disorder is thought to be caused by excess water as well, since it
was observed that gummosis was more pronounced in the lower part, the north and the
east side of the canopy (Table 4), where less sunlight is received and less transpiration
occurs (Limpawiphagorn, 1998).
To avoid the translucent pulp and gummosis disorders in mangosteen, preharvest
practices must be improved. It is recommended that mangosteen should be grown in a
well-ventilated area. Tree spacing should be open and the tree should also be pruned to
allow transpiration especially during rain. Another suggestion is to promote early or off
season production of mangosteen to achieve higher prices and ensure better quality.

The basic postharvest physiology of tropical fruit is similar to subtropical and
temperate fruit. However, all of them suffer from chilling injury, but very little work has
been done on the physiology of this disorder in tropical fruit. Many of them have peels
that are physiologically distinct from the pulp and require special attention after harvest
in order to satisfy consumers. An important requirement is to synchronize changes in the
peel with those in the pulp. There are many phenomena in tropical fruit that have not
been observed in subtropical and temperate fruit. Unfortunately, there is much less
physiological research as compared to work on technology. To increase market share for
tropical fruit in the world market, a lot more attention should be given to their physiology
both before and after harvest.
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Table 1. Soluble solids content, titratable acidity, respiration and ethylene production
rates of some tropical fruits1/.
Fruit Type or Cultivar Soluble Acid Respiration3/ Ethylene
solid2/ content2/ (mgCO2/kg.hr.) production3/
(%) (%) l/kg.hr.)
Banana AA Khuai Khai 4-24 0.4-0.9 0.23-0.42 (18 C) 40-125 (18 C)
AAA Cavendish 3.5-24.3 0.04-0.27 55.7-450 (25 C) 10-48 (25 C)
Carambola mixed cultivar 7-10 0.2-7.0 20 < 0.5
Durian Chanee 14-26 0.06-0.10 200-750 (28 C) 2-40 (28 C)
Monthong 13-30 0.06-0.13 100-350 (28 C) 2.4-11 (28 C)
Guava mixed cultivar 7-9 0.2-0.4 70-150 1.0-10.0
Jackfruit 18-23 0.3-0.7 35-70 -
Java apple mixed cultivar 7-12 0.2-1.2
Longkong mixed cultivar 13-19 1.0-1.5 20-30 (RT)
Mango Nam Dok Mai 17-22 0.2-1.49 150-300 (28C) 5-23 (28 C)
Carabao 19.0 0.3 50-125 0.02-0.05
Mangosteen - 15-20 0.2-0.7 10-70 1-16
Papaya mixed cultivar 8-15 0.1-0.14 8.5-25 (25 C) 0.1-1.5 (25 C)
Pineapple Smooth Cayenne 11-18 0.4-1
< 35 0.1-1.0
Queen 15-17 0.47-0.7
Rambutan Rongrein 17-21 0.3-5.5 30-55 (25 C) 0.5-2.6 (25 C)
Salak mixed cultivar 18-19 0.4-0.9 75-150 24-13
Santol Local Thai 7-15 3.2 80-140 (28) 1-7 (28)
Sapodilla mixed cultivar 12-22 0.11-0.41 2-10.25
Sugar apple mixed cultivar 25-26 0.1-0.3 200-400 ~ 1.1
Data combined from various sources including Nakasone and Paull (1988), Mitra (1997),
Champ et al. (1994), Vijaysegaran et al. (1996), and data from the authors laboratory.
Amount during eating stages
Rate at pre and peak climacteric, or during eating stage.

Table 2. Quality of Nam Dok Mai mango treated with 200 ppm ethylene at different
temperatures (Sangvanangkul, 1997).

Temperature Peel color Pulp color Firmness SS TA

(C) score score (N) (%) (%)
20 3.18 a 3.22 a 5.88 a 15.6 a 0.26 b
25 3.44 a 4.37 b 5.39 a 15.4 a 0.09 a
30 3.15 a 4.85 c 5.49 a 16.6 b 0.09 a
: Means followed by the same letter are not significantly different by DMRT at Pr < 0.05

Table 3. Effect of method of application of water on physiological disorders of mangosteen

fruit (Luckanatinvong et al, 2000).
Physiological disorder ( number of fruit out of 400)
Crack Translucent Gummosis End-fruit
Ground sprinker (control) 15 b 60 b 72 a 156 a
Over head sprinker 66 a 121 a 97 a 76 c
Tyvex mulching and ground 59 a 78 b 79 a 103 bc
Tyvex mulching and over 56 a 114 a 58 a 122 b
head sprinker

Means followed by the same letter are not significantly different by DMRT at Pr 0.01
Table 4. Percentage of gummosis in mangosteen harvested from different parts of the
canopy (Limpawiphagorn, 1998).

Province Direction Lower Upper

North South East West Section Section
Trad 5.6 ab1/ 25.5 ab 25.0 ab 18.9 b 32.2 a 18.2 b
Chantaburi 54.2 xy 53.9 xy 51.2 xy 46.8 y 58.1 x 48.0 y
Means followed by the same letter are not significantly different by DMRT at Pr < 0.05


Reactive O2 Oxidative Membrane

Species Damage Breakdown
superoxide lipids
H2 O2 proteins
Chilling Visible Signs of
Stress Injury
alpha-tocopherol superoxide dismutase
ascorbic acid catalase
beta-carotene peroxidases

Fig. 1. A recent hypothesis on the mechanism of chilling injury (Shewfeft and del Rosario,







Total phenolics
(Abs 760 nm)





2.0 C
(Abs 280 nm)
Lignin content

0 1 2 3
Hours after Impact

Fig. 2. Effect of relative humidity (RH) or 24-h ethylene treatment on dehiscence score
of mature Chanee or Monthong durian (110 days post anthesis) (!) 65% RH,
(") 95% RH, (#) 0 ppm C2H4, ($) 100 ppm C2H4 (Sriyook et al., 1994) (n=5)


Dehiscence score

1 2 3 4 5

Days in storage

Fig. 3. Changes in (A) firmness, (B) phenolics (760 nm) and (C) lignin of undamaged ( )
and damaged (#, %) pericarp of mangosteen fruit in air (%) and nitrogen (#)
after impact. (Ketsa and Atantee, 1998)

Colorless area

Brown stained

Fig. 4. I2-KI (1.0%) staining of cross-sectioned Monthong durian 113 days after