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Abstract
Application of exogenous ethylene, irrespective of the method of application, caused intensification of mesocarp
discoloration in avocado fruit (Persea americana Mill.) during cold storage of all cultivars tested. ‘Ettinger’ fruit
treated with Ethrel (2-chloroethyl phosphonic acid) prior to packing and storage developed severe chilling injury (CI)
symptoms, expressed as mesocarp discoloration after 3 weeks at 5 °C. ‘Fuerte’ fruit treated with ethylene gas (100 ml
l − 1) for 24 h at 20 °C prior to storage at 5 °C exhibited mesocarp discoloration, which increased dramatically during
shelf life at 20 °C. ‘Fuerte’ fruit treated in cold storage with a continuous low ethylene dose (4 ml l − 1) developed
severe browning in the fruit pulp after 3 weeks at 5 °C. ‘Hass’ fruit treated with 50 ml l − 1 ethylene, for 12, 24 or 48
h at 5 °C showed a gradual increase in mesocarp discoloration after 3 weeks in cold storage plus shelf life; the 48 h
ethylene-treated fruit exhibited the most severe pulp browning. Use of absorbent sachets that removed ethylene from
modified atmosphere (MA) packaging reduced mesocarp discoloration and decay development in ‘Hass’ fruit after 5
weeks storage at 5 °C. Application of 1-methylcyclopropene (1-MCP), reduced mesocarp discoloration, decay
development and polyphenol oxidase activity, whereas this enzyme activity was induced in ethylene-treated fruits that
were cold stored for 4 weeks. © 2002 Elsevier Science B.V. All rights reserved.
Keywords: Persea americana; Chilling injury; Ethrel; 1-Methylcyclopropene; Ethylene absorbent; Modified atmosphere
0925-5214/02/$ - see front matter © 2002 Elsevier Science B.V. All rights reserved.
PII: S 0 9 2 5 - 5 2 1 4 ( 0 1 ) 0 0 1 3 4 - X
172 E. Pesis et al. / Posthar6est Biology and Technology 24 (2002) 171–181
2.5. Fruit ripening and chilling injury indices 2.6. Statistical analysis
Fruit firmness was determined by destructive Each treatment was applied to five export car-
and non-destructive methods. In the destructive tons. A 4-kg carton contained 14–16 fruit of cvs.
method, firmness was measured on two cheeks of Ettinger and Fuerte or 16–18 fruit of cv. Hass.
each fruit, using an electronic penetrometer Most data were expressed as mean9 SE. The
(Chatillon, New York) with a 6.5-mm conical tip. results of gas analysis in MAP are means of five
174 E. Pesis et al. / Posthar6est Biology and Technology 24 (2002) 171–181
3. Results
Table 1
Effect of ethylene application (100 ml l−1, 24 h at 20 °C) on fruit firmness (index 10–1), CI exocarp (index 0–10) and CI mesocarp
discoloration (% of fruit) of ‘Fuerte’ fruit after 2 and 6 days at 20 °C following 3 weeks at 5 °C
Firmness (index) CI exocarp (index) Firmness (index) CI exocarp (index) CI mesocarp (%)
Table 2
Effect of two ethylene absorbent sachets (MA+abs.) in MA packaging on decay development (%) and mesocarp discoloration (%)
in ‘Hass’ fruit after 3 and 7 days at 20 °C following 5 weeks at 5 °C
Mean separation in columns in the various dates by Duncan’s multiple range test, 5% level.
centrations in the MA packaging without sachets reduction in mesocarp discoloration, without in-
reached around 3 nmol l − 1 (Fig. 3). terfering with the softening process after 5 weeks
Storage of ‘Hass’ fruit (without any fungicide at 5 °C (Table 3). Application of 1-MCP (300 nl
treatment) in MA for 5 weeks led to decay devel- l − 1 for 24 h at 5 °C) reduced mesocarp discol-
opment, especially stem-end rot, compared with oration significantly and did not prevent soften-
unwrapped fruit (Table 2). In MA packaging with ing. However, a second application of 1-MCP
ethylene absorbent sachets, fruit exhibited signifi- (300 nl l − 1) after 10 days impaired the softening
cantly less mesocarp discoloration and lower of ‘Hass’ fruit: the fruit failed to soften normally
stem-end rot levels. The levels of the side rots and remained firm during shelf life (Table 3).
were relatively low and did not differ between ‘Fuerte’ fruit suffered from a significant amount
fruit stored in MA and those in MA plus ab- (50%) of mesocarp discoloration after 6 days at
sorbent sachets (Table 2).
Treatment Day of Gas (concentrations) following 24 h treatment Firmness (N) days at shelf life at 20 °C Mesocarp discoloration
1-MCP (nl l−1) treatment at (% of fruit)
5 °C
CO2 (kPa) O2 (kPa) C2H4 (ml l−1) 0 4 7
0 1 5.69 0.12 13.7 9 0.04 0.11 90.01 128.9 921.6 11.692.8 7.8 9 1.4 7.1 90.0
100 1 6.1 9 0.07 13.1 90.12 0.38 9 0.16 127.9 910.8 13.99 2.7 8.1 9 1.3 2.4 9 2.1
300 1 5.8 90.10 13.6 90.07 0.20 9 0.01 135.6 94.3 12.693.8 7.6 9 0.7 0.5 9 0.0
100×2 1 and 10 3.5 9 0.08 15.1 9 0.06 1.09 90.29 138.2 99.2 10.592.9 7.59 1.3 0.59 0.0
300×2 1 and 10 3.0 90.05 15.5 90.11 0.84 90.18 130.7 97.1 61.6930.5 26.2 9 17.9 0.5 9 0.0
Fruit firmness (N) was determined during shelf life at 20 °C following 5 weeks at 5 °C, and mesocarp discoloration (% of fruit) was determined on the seventh day
of shelf life. Data are mean 9 SE.
E. Pesis et al. / Posthar6est Biology and Technology 24 (2002) 171–181
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178 E. Pesis et al. / Posthar6est Biology and Technology 24 (2002) 171–181
4. Discussion
ethylene application clearly demonstrated that 1972). The greater stem-end rot development un-
ethylene exposure prior to cold storage could der MA conditions with high ethylene concen-
cause increased damage. This agrees with previ- trations (3 ml l − 1) corresponds to the in vitro
ous findings of induction of stem-end rot caused induction of C. gloeosporioides by ethylene
by Diplodia natalensis in ethylene-treated citrus (Flaishman and Kolattukudy, 1994).
fruit (Brown and Lee, 1993) and induction of Recently, it has been shown that application
fungal invasion by Colletotrichum gloeosporioides of 1-MCP to avocado fruit at a very low con-
by ethylene (Flaishman and Kolattukudy, 1994). centration of 30–70 nl l − 1 for 24 h was capable
The main stem-end rot fungus in avocado is C. of extending shelf life at 20 °C by delaying eth-
gloeosporioides, which causes great damage to ylene production (Feng et al., 2000). A similar
the avocado industry in Israel (Prusky, 1996). low concentration (100 nl l − 1) was also effective
There was no change in softening following a in delaying banana ripening (Jiang et al., 1999).
12-h ethylene application, and only a very slight In the present study, with application at 5 °C,
change in fruit softening after 24– 48 h. This and not 20 °C, a longer application or a second
agrees with the findings of Zauberman et al. application of 100 nl l − 1 for 24 or 48 h was
(1988), who showed that a 12 h ethylene appli- more effective in improving avocado quality
cation did not trigger softening. The appearance than only one application of 100 nl l − 1 for 24 h
of CI is not related to softening, and it has (Fig. 4). In banana, Jiang et al. (1999) found
been found in a more severe form in firm precli- that application of 500 nl l − 1 of ethylene in PE
macteric fruit (Paull, 1990). bags was more effective than 100 nl l − 1, while
There was more mesocarp discoloration and application of 1000 nl l − 1 was not more effec-
stem-end rot development after 3 weeks of cold tive, but did not cause damage. However, when
storage, in ‘Fuerte’ fruit treated with ethylene, 1-MCP was applied twice at 300 nl l − 1 for 24 h
stored at 3 °C than in that stored at 5 °C (Fig. during cold storage, avocado fruit failed to
2). This was probably because at the lower tem- ripen during shelf life, which indicated that 1-
perature there was more membrane damage, MCP, in higher concentrations, can be effective
which is the primary response of plants to chill- also at 5 °C (Table 3). Application of AVG (an
ing stress (Wang, 1982). inhibitor of ethylene synthesis) plus NBD (an
Use of MA enabled us to study the effects of inhibitor of ethylene action) to avocado slices
reduced ethylene concentrations in the atmo- completely inhibited ethylene production and ex-
sphere around the fruit, on mesocarp discol- pression of cellulase and polygalacuronase (PG)
oration and decay development. Ethylene genes (Starrett and Laties, 1993). It is possible
reduction was achieved by direct absorption of that applying excess 1-MCP can lead to irre-
ethylene and perhaps by elimination of autocat- versible inhibition of softening, because both the
alytic formation of new ethylene. Slowing the cellulase and PG enzymes are regulated by eth-
ripening process, by lowering the ethylene con- ylene and are initiated by the climacteric rise
centration in the atmosphere around fruit, led to (Starrett and Laties, 1993).
reduced CO2 and higher O2 concentrations (Fig. One aim of the present study was to evaluate
3). Avocados stored in MA with low ethylene the efficacy of 1-MCP in preventing CI in stored
exhibited lower levels of mesocarp discoloration avocado. It is obvious that 1-MCP reduced the
and stem-end rot development. This demon- harmful effect of ethylene in cold storage on
strated the role of ethylene in intensifying meso- mesocarp discoloration. Mesocarp discoloration
carp discoloration and stem-end rot was the lowest in ‘Fuerte’ fruit treated twice
development following 5 weeks of storage at with 1-MCP (100 nl l − 1) for 48 h during 4
5 °C. Our results are in agreement with others weeks at 5 °C, and this corresponded with the
that show reduction in CI in avocado expressed lowest PPO activity (Fig. 4). On the other hand,
as mesocarp discoloration, through removal of continuous application of ethylene (4 ml l − 1 at
ethylene by an absorbent (Hatton and Reeder, 5 °C) caused 100% pulp browning and the
180 E. Pesis et al. / Posthar6est Biology and Technology 24 (2002) 171–181
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