The Journal of Horticultural Science and Biotechnology

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The cytokinin CPPU delays maturity in litchi cv.

‘Mauritius’ and extends storage-life

A. Raphael Stern, Ohad Nerya & Ruth Ben-Arie

To cite this article: A. Raphael Stern, Ohad Nerya & Ruth Ben-Arie (2006) The cytokinin CPPU
delays maturity in litchi cv. ‘Mauritius’ and extends storage-life, The Journal of Horticultural
Science and Biotechnology, 81:1, 158-162, DOI: 10.1080/14620316.2006.11512043

To link to this article: http://dx.doi.org/10.1080/14620316.2006.11512043

Published online: 07 Nov 2015.

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 Journal of Horticultural Science & Biotechnology (2006) 81 (1) 158–162

The cytokinin CPPU delays maturity in litchi cv. ‘Mauritius’ and

extends storage-life

By RAPHAEL A. STERN1*, OHAD NERYA2 and RUTH BEN-ARIE2

1
MIGAL, Galilee Technology Center, P.O. Box 831, Kiryat-Shmona 11016, Israel
2
Fruit Storage Research Laboratory, Kiryat-Shmona, Israel
(e-mail: Raffi@migal.org.il) (Accepted 17 October 2005)

SUMMARY
The effects of the synthetic cytokinin N-(2-chloro-4-pyridyl);N’-phenylurea (CPPU) on fruit development and the
storage-life of litchi (Litchi chinensis Sonn.) cv. ‘Mauritius’ were examined in Israel over 5 years. CPPU at 5 – 10 mg l–1,
applied to green or slightly red fruitlets (25 or 30 mm in diameter), delayed harvesting by 2–3 weeks compared with
control trees. At harvest, CPPU-treated fruit that attained a red colour comparable to that of earlier harvested control
fruit, were 20–25% larger with total soluable solid contents:titratable acidity (SSC:TA) ratios more than 50% higher
than the controls. Despite their high SSC:TA ratios, CPPU-fruit stored well for 6 weeks at 1°C due to reduced
browning, lower decay development and less aril discolouration, and maintained an acceptable flavour. These results
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suggest that CPPU can be used to extend the harvest season for litchi fruit.

L itchi, cultivated mainly in South East Asia, has

become a popular exotic fruit for fresh consumption
in Europe and North America in recent years (Huang et
al., 2005). The supply from this region is limited from
May to July, due to a short harvesting period and limited
post-harvest life associated with rapid browning of the
attractive red pericarp, decay and discolouration of the
aril, and loss of flavour (Zee et al., 1998; Holcroft et al.,
2005; Huang et al., 2005).
Israel has no marketing advantage in early Summer,
when European markets are abundantly supplied by the
Far East, and therefore no effort has been made to
accelerate ripening. However, there is a marketing niche
from August to October, before litchi fruit begin to be
supplied from South Africa and Madagascar (Huang et
al., 2005). Therefore efforts have been made to delay
harvest in areas where high quality fruit are produced, by
introducing new cultivars (Goren et al., 1999; 2001) and
by developing methods to prolong the post-harvest life
of litchi cv. ‘Mauritius’ (Lichter et al., 2000; Pesis et al.,
2002). However, cold storage generally limits subsequent
shelf-life. Likewise, delaying harvest also shortens
storage-life, due to a range of processes that affect fruit
senescence (Pesis et al., 2001; 2002).
We proposed that senescence could be deferred by
cytokinins, which delay senescence in other fruit (Arteca,
1996). Dipping litchis in 100 mg l–1 benzyladenine (BA)
gave higher chlorophyll concentrations than in control
fruit through lower chlorophyllase activity and delayed
colouration (Wang et al., 2001; Huang, 2005). In grape, the
synthetic cytokinin N-(2-chloro-4-pyridyl);N’-phenylurea
(CPPU) applied at fruit set, or just after, at 5 – 10 mg l–1
delayed maturation by up to 2 weeks (Ben-Arie et al.,
1997). The same cytokinin, applied at 2 – 5 mg l–1 to
fruitlets delayed maturation in persimmon as indicated
by colour development (Prusky et al., 2004). A spray
application of 10 mg l–1 CPPU to blueberries 3 weeks
*Author for correspondence.
after full-bloom postponed harvest by 3 weeks, a
significant achievement (Yang, 2003). The objective of
our study was to extend the marketing season of litchi
fruit by pre-harvest application of the cytokinin, CPPU.
MATERIALS AND METHODS
In all experiments, the commercial product CipionTM
(Degussa Co. Ltd., Germany), containing 1% CPPU a.i.
was applied with 0.025% (v/v) Triton X-100 as a
surfactant.
Preliminary bunch trial (Almagor, 2000)
The trial was conducted in a 7 year-old orchard in
Eastern Galilee, 100 m below sea level. Tree spacings
were 6 m
6 m and the yield was about 15 tonnes ha–1.
Bunches with about 25 pink-to-green fruit were dipped
in CPPU at 0, 5 or 10 mg l–1 20 d, 10 d or 1 d before the
expected commercial harvest. The bunches (with leaves)
were immersed for 10 s in 4 l of water (control) or either
CPPU solution. The experiment was laid out in a
randomised block design with 10 single tree blocks and
one bunch per tree for each treatment.
Fruit were harvested when the majority of the controls
were at their optimum colour (see “Fruit assessment”).
Maturity at harvest was assessed on five fruit per
replicate, and the remainder were treated with SO2
according to commercial protocols (Holcroft et al., 2005).
Fruit from each replicate were packed in three ventilated
PVC punnets (10–14 litchis), and stored at 1°C for 3, 6 or
9 weeks. The external appearance of the fruit was
examined when they were removed from cold storage,
and the internal quality assessed after storage at 20°C
(see below).
Whole tree trial (Bet HaEmek, 2001 and 2002)
A 25 year-old orchard in the Western Galilee, 100 m
above sea level, was used in 2001 and 2002 with similar
spacings and yields as before. Individual trees were
 R. A. STERN, O. NERYA and R. BEN-AIRE
sprayed with 2 – 3 l of 5 or 10 mg l–1 CPPU, when fruits
were 25 – 30 mm wide, using a high presser hand-gun
sprayer (Kibbutz Degania, Israel) until run-off. The
direction of the rows in this orchard was North-South and,
in normal years, there was no difference in ripening rates
on both sides of the trees. However, a hot spell in early
June 2002, when fruits were 12 mm wide, accelerated
ripening on the East side compared to the West. Thus, on
the western side of the tree at the first application of
CPPU on July 2, fruit were light green on their East side
and dark green on their West side and, at the second
application on July 10, western fruit were light green and
eastern fruit pinkish. Therefore, in 2002 the experiment
consisted of two spray treatments at four stages of fruit
development, with an unsprayed control, in each of four
randomised blocks, with one tree per treatment. A similar
trial conducted in 2001 with two spray treatments at two
stages of fruit development, produced similar results. Only
the results from 2002 are presented.
Fruit maturation was assessed each week on five fruit
per replicate, from spraying until harvest, by measuring
fruit diameter, fruit weight, pericarp colour (using a scale
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of 1–5, see below), total soluble solids content (SSC) (%)
and titratable acidity (%). Fruit for storage were
harvested when 60% of the fruit on the tree was deemed
to be commercially acceptable. At the first harvest, only
fruit on the East side were harvested. The second harvest
occurred 10 d later and was the principal harvest, after
which few fruit remained on the controls. By the fourth
harvest no fruit remained on the controls.
Harvested fruit were treated, stored and examined as
described above, except that storage at 1°C was 4, 8 or 12
weeks and storage at 20°C was 3 d.
Commercial-scale trial (Bet HaEmek, 2003 and 2004)
Four randomised blocks, each consisting of three rows
of trees with 50 trees per row (150 per treatment), were
sprayed with 10 mg l–1 CPPU at 1,500 l ha–1, when fruit
were 25 mm-wide, using a commercial 1000 l air-blast
“Spidet” blower sprayer (Kibbutz Degania, Israel). Four
representative trees per treatment along the middle row
of each block were marked for sampling to assess fruit
maturation from spraying to harvest, and post-harvest
storage, as described previously. However, in these trials,
six punnets per replicate, each containing 250 g fruit,
were stored at 1.5°C. Fruit were examined after 3, 6 or 9
weeks of storage and then 3 d later at 20°C. A similar
trial, conducted in 2003, produced similar results, but
only the data for 2004 are presented.
Fruit assessment
Fruit colour was monitored visually on a five-point
scale from 1 = 100% green, to 5 = 100% red. The
diameter and weight of individual fruit were recorded.
The juice from five fruit was combined to assay SSC with
TABLE I
Stages of fruit development on the two spraying dates at Bet HaEmek in 2002.
Stage Exposure Spray date Colour grade (1–5)†
1 West 2 July 1.1d*
2 West 10 July 1.9c
3 East 2 July 2.5b
4 East 10 July 3.6a
*Column means followed by different lower-case letters are significantly different (P ≤ 0.05). n = 4.
†
Colour rated from 1 = 100% green, to 5 = 100% red.
159
a digital refractometer (PR-100 Atago Co. Ltd., Tokyo,
Japan) and titratable acid (TA) by automatic titration of
2 ml to pH 8.2 with 0.1 M NaOH. The percentage acidity
was calculated based on malic acid equivalent.
After storage weight loss was determined by calculating
the percentage change in weight of each punnet.
Browning was assessed using a five-grade scale: (1 = no
browning; 2 = brown pinnacles at the centre of the
tubercles; 3 = small brown patches; 4 = large brown
patches; 5 = > 75% of brown skin area), and fruit softness
on a three-grade scale (1 = soft; 2 = flexible; 3 = firm).
Fruit used after storage were also peeled and
discolouration of the aril was graded from 1 (= no
discolouration) to 4 (= severe discolouration). Juice was
extracted from five fruit per replicate for SSC and TA
measurements, and the remainder of the fruit used for
taste-testing by 8–10 people who were requested to
evaluate overall taste on a hedonic scale from 1 to 10.
The maximum number of samples per taste session was
six. Two-to-three fruit per sample were tested by each
panel number, with a mouthwash between samples.
Statistical analysis
All statistical analyses were performed using SPSS
12.1 for Windows. Data were submitted to one-way
analysis of variance (ANOVA), followed by Duncan’s
multiple range test to separate means (P ≤ 0.05).
RESULTS
Preliminary bunch trial (Almagor, 2000)
Treatments applied 1 or 10 d before harvest had no
significant effect on litchi fruit ripening or storage
potential. Therefore only data from the first application,
20 d before harvest, are presented. Colour development
was retarded by both 5 and 10 mg l–1 CPPU compared
with controls when commercial harvest began (colour
indices of 1.8, 0.5 and 3.2, respectively). Both CPPU
treatments inhibited browning after harvest and 3 weeks
in storage, but not after 6 weeks in storage (data not
shown). Although fruit at this stage were still marketable,
there was no advantage of the chemical treatment in
terms of external or internal appearance. The external
appearance of the fruit was still acceptable after 9 weeks,
but the arils were discoloured (data not shown).
Whole tree trial (Bet HaEmek, 2002)
In the preliminary trial, all fruit were harvested on the
same day, so that CPPU-treated fruit were not fully
matured. It was therefore of interest to examine how
long harvest could be delayed by CPPU treatment, as
well as to observe its effects in a different environment,
which naturally delayed harvesting. Unexpectedly, this
orchard offered an opportunity to examine CPPU
application at four stages of maturity (Table I). At the
Diameter (mm) SSC (%) TA (%) SSC/TA ratio
25.4c 12.0c 5.5a 2.2c
27.8b 15.9b 2.4c 7.1b
27.7b 15.3b 3.0b 5.3b
29.9a 17.0a 1.09d 16.1a
 160 Cytokinins delay litchi maturation

40 TABLE II
A a Effect of CPPU on the proportion (%) of fruit suitable for harvesting on
14 and 23 July at Bet HaEmek in 2002
Diameter (mm)

35 b b A
B
c C CPPU (mg l–1) Spray date 14 July 23 July
30 e d D 0 – 85.0a* 100.0
E 5 2 July 31.3bc 55.0a
25 F West 5 10 July 40.0b 52.5a
G
East 10 2 July 27.5bc 30.0b
20 10 10 July 17.5c 25.0b
1/7* 5/7* 9/7* 14/7* 23/7 28/7 4/8
*Column means followed by different lower-case letters are significantly
Date different (P ≤ 0.05). n = 4.

20 a control trees grew at a similar rate, although eastern fruit

B b
18 b b
were larger than western fruit from 1–14 July and had
A
c B C higher SSC. In contrast, TA was lower in eastern fruit on
SSC (%)

16 d
D D all dates. As the main criterion for commercial harvest is
14 fruit colour, the effect of CPPU on ripening was
E
West indicated by fewer suitable fruit at the first and second
12
F East
10
harvests (Table II). Harvesting finished on these trees in
1/7* 5/7* 9/7* 14/7* 23/7 28/7 4/8
early August. At the first harvest, CPPU delayed colour
Date
development only if applied early (2 July), whereas at the
second harvest, all applications were effective (except for
the early spray to stage 3 fruit, which had already
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7 coloured at this time; Table III).

6 C West CPPU increased fruit size when fruit from the
A East
5 B different treatments were compared when fully ripe or
4
close to fully ripe (e.g., CPPU-treated fruit on August 4
TA (%)

3 C
a
2 D
b E
1
c E
0 d e e
-1
1/7* 5/7* 9/7* 14/7* 23/7* 28/7*
Date
FIG. 1
Fruit diameter (mm; Panel A); total soluble solids content (SSC%; Panel
B); and titratable acidity (TA%; Panel C) on the East and West of the
trees, from time of CPPU-spray application to harvest. Means with
different upper-case (West) and lower-case (East) letters are
significantly different (P ≤ 0.05). Asterisks at a given sampling date
indicate significantly different (P ≤ 0.05) means (n = 4).
second application, untreated fruit on the West side were
similar to untreated fruit on the East side in terms of size
and SCC:TA ratio, but not in terms of colour or acidity.
For the latter two parameters, fruit were therefore
effectively treated at four stages of maturity (1 to 4).
Differences in ripening, measured on two sides of each
tree at the time of spraying, were less distinct at the
beginning of commercial harvest (Figure 1). Throughout
the 2-week harvesting period, fruit on both sides of
vs. control fruit on July 23).
E CPPU and control fruit had similar SSC:TA ratios
when compared on the same date (Table IV). However,
4/8
CPPU-treated fruit harvested on 28 July (except those
on the East treated on 2 July) had higher SCC:TA ratio
values than the respective controls, which were all
harvested by 23 July. CPPU-treated fruit (West, 2 July)
harvested on 4 August also had higher SSC:TA ratios
than controls harvested by 23 July.
The keeping quality of litchi deteriorates as the
harvest season progresses. Hence, a comparison of the
keeping quality of untreated and CPPU-treated fruit
harvested on the same day would be a combination of
the effects of CPPU on fruit quality and delayed
ripening. Accordingly, a comparison was made between
the storage potential of control fruit from the first
harvest (14 July) and CPPU-treated fruit from the third
harvest (28 July) when, in both cases, approx. 85% of
fruit were considered suitable for harvest on the basis of
colour. In other aspects (fruit size and SSC:TA ratio),
control and CPPU-treated fruit differed significantly
(Tables III and IV). After 4 weeks storage at 1°C, control

TABLE III
Effect of CPPU (East or West exposure, sprayed on 2 or 10 July) on average litchi fruit size and colour over four harvests at Bet HaEmek in 2004
Fruit colour index (1–5)† Fruit diameter (mm)
Harvest CPPU West West East East West West East East
date (mg l–1) 2 July 10 July 2 July 10 July 2 July 10 July 2 July 10 July
14 July 0 3.1c* 3.1de 4.5a 4.5abc 31.1d 31.1f 32.6bc 32.6bc
5 2.4d 2.6e 4.0b 4.3cd 31.1d 32.3ef 32.5bc 32.6bc
10 2.3d 2.6e 3.9b 4.0de 31.8cd 32.0ef 32.5bc 32.7bc
23 July 0 4.4a 4.4a 4.8a 4.8ab 32.8c 32.8def 31.3c 31.3c
5 3.6b 3.3d 4.7a 4.4bcd 32.8c 33.9cde 33.5bc 33.7abc
10 3.7b 3.1de 4.5a 4.2cde 32.8c 33.5de 32.7bc 33.8abc
28 July 5 4.4a 3.9c 3.9b 4.9a 35.2b 35.6bc 36.3a 35.5a
10 3.9b 4.1abc 3.8b 3.8e 35.7ab 34.8cd 34.8ab 35.7a
4 August 5 4.8a 4.5a ** ** 35.8ab 38.4a ** **
10 4.5a 4.0bc ** ** 36.8a 36.9ab ** **
*Column means followed by different lower-case letters are significantly different (P ≤ 0.01). n = 4.
**No fruit remaining on the tree.
†
Colour rated from 1 = 100% green, to 5 = 100% red.
 R. A. STERN, O. NERYA and R. BEN-AIRE 161

TABLE IV TABLE V
Effect of CPPU sprayed on 2 or 10 July and East or West exposure on Effect of CPPU (East or West exposure, sprayed on 2 or 10 July) on fruit
SSC:TA ratio at four harvests at Bet HaEmek in 2002 quality after 8 weeks of storage at 1°C, for fruit harvested at the peak of
harvest for each treatment at Bet HaEmek in 2002
Harvest West West East East
date Treatment 2 July 10 July 2 July 10 July Exposure/ SSC/TA
CPPU Date of ratio Decay Pericarp
14 July Control 14.3c* 14.3d 31.4b 31.4c (mg l–1) spraying at harvest (%) browning (1–5)†
CPPU† 11.6c 13.0d 27.1b 26.9c
23 July Control 36.9b 36.9c 46.6ab 46.6bc 0 – 22.9c* 90.0a 1.7a
CPPU 33.2b 36.2c 57.0a 51.0b 5 West, 2 July 26.5c 7.5bc 1.2c
28 July CPPU 55.5a 45.5b 61.0a 63.7a 5 West, 10 July 33.2bc 3.7c 1.3bc
4 August CPPU 58.6a 56.1a ** ** 5 East, 2 July 62.0a 15.0b 1.4b
5 East, 10 July 61.0a 20.0b 1.2c
*Column means of SSC:TA ratios followed by different lower-case 10 West, 2 July 36.9b 3.5c 1.5ab
letters are significantly different (P ≤ 0.01). 10 West, 10 July 36.2b 3.4c 1.6ab
†
Values for CPPU are the average for 5 and 10 mg l–1 applications. (n = 4). 10 East, 2 July 62.0a 20.0b 1.4b
**No fruit remaining on the tree. 10 East, 10 July 64.0a 20.0b 1.4b
* Column means followed by different letters are significantly different
(P ≤ 0.05).
and CPPU-treated fruit were still of satisfactory quality, †
Browning scored from 1 = no browning, to 5 = 75-100% brown. (n = 4).
without significant browning, decay or aril
discolouration, and were of acceptable taste (data not
shown). However, after an additional month of cold The keeping quality of litchi fruit, stored at 1.0°– 1.5°C
storage, fruit quality deteriorated in the controls due to for up to 9 weeks, was better after CPPU-treatment,
decay (Table V). Most CPPU-treated fruit also had less mainly due to less decay. This can probably be attributed
browning and aril discolouration (unrelated to SSC:TA to harvesting of fruit with younger pericarps, rather than
at harvest). Although riper CPPU-treated fruit were to a direct fungicidal effect of CPPU. CPPU was applied
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more susceptible to decay than less-ripe fruit, the
incidence of decay after storage remained significantly
less than in control fruit harvested 2 weeks earlier at a
more immature stage of ripening.
Commercial-scale trial (Bet HaEmek, 2004)
Fruit sprayed with 10 mg l–1 CPPU were harvested 5 d
later than those from control trees, but were significantly
greener (3.4 vs. 4.0), larger (34.4 mm wide and 25.1 g vs.
32.1 mm wide and 21.9 g, respectively) and had higher
SSC:TA ratios (49.5 vs. 33.7, respectively) and less decay
after 6 and 9 weeks storage (2.0% vs. 18% and 6.7% vs.
49.8%), respectively. Less aril discolouration also
occurred after 9 weeks in storage (70% vs. 92%). In spite
of some deterioration in quality, the taste of CPPU-
treated fruit remained acceptable (rated 6.6 out of 10)
compared to control fruit (rated 4.0). Browning, fruit
weight loss, fruit softening and changes in SSC and TA,
were not affected significantly (P ≤ 0.05) by CPPU-
treatment (data not shown).
DISCUSSION
The criteria commonly used for ripeness in litchis are
a red pericarp and an SSC:TA ratio of 30:1 (Underhill
and Wong, 1990; Zee et al., 1998; Holcroft et al., 2005).
When the SSC:TA ratio is 40:1, or higher, fruit ferment,
lose sugars and accumulate acids, acetaldehyde and
ethanol (Pesis et al., 2002; Holcroft et al., 2005). A similar
process has been described in other members of the
Sapindaceae, including longan and rambutan (Zee et al.,
1998). Therefore, to extend marketing, ripening needs to
be delayed while the litchis are still on the tree. Our
study has shown that ripening in litchi cv. ‘Mauritius’ can
be retarded by applying cytokinins. Spraying trees with
CPPU at 5 or 10 mg l–1, when the fruit were 25 – 30 mm
wide, delayed reddening and harvest by 2–3 weeks.
Similar results have been obtained with two other
synthetic cytokinins, TDZ (15 mg l–1 a.i.) and BA (50 mg
l–1 a.i.), without injuring the fruit or the tree (unpublished
results). However, SSC:TA ratios and fruit weights were
not always affected.
after cell division had terminated and it is unlikely that it
was re-initiated by the treatment. The delay in pericarp
maturation, shown by the slower loss in chlorophyll and
anthocyanin synthesis, might have affected the
development of the exocarp and the spongy tissue of the
mesocarp (Huang et al., 2004). Thus the elasticity of the
periderm could have been enhanced, and the extent of
micro-cracking reduced, which in turn would slow the
penetration of decay-causing organisms. As rotting is the
main factor that limits litchi storage under the high
relative humidity required to reduce browning, this
effect of CPPU-treatment is important.
The effects of cytokinins on aril ripening and
senescence are more complicated. On the one hand, the
younger pericarp with its greater elasticity enabled
continued aril growth with the increased SSC and
degradation of acids. As a result, the SSC:TA ratio in
CPPU-treated fruit at harvest exceeded the
recommended value of Underhill and Wong (1990). On
the other hand, despite the high SSC:TA ratio at harvest,
aril discolouration was somewhat inhibited and fruit
taste was not adversely affected during storage, in
response to CPPU-treatment. These findings suggest that
neither aril discolouration nor taste, which are indicative
of senescence, are directly connected to fruit SSC or
acidity at harvest. The onset of senescence in non-
climacteric litchi fruit is not associated with an outburst
of respiration in response to ethylene biosynthesis, but
rather to increased anaerobic respiration (Pesis et al.,
2002). The decline on aril appearance and taste have
been associated with the accumulation of acetaldehyde
and ethanol during storage (Holcroft et al., 2005).
Although these metabolites were not monitored in this
study, the results obtained from the taste panels suggest
that the onset of anaerobic respiration during storage
might have been delayed by CPPU (unpublished data).
Consequently, pre-harvest application of CPPU enabled
the marketing season of litchi to be extended due to
delayed harvest and better quality in storage.
The authors thank the growers Danny Stern, Roberto
Glussman, Hillel Miller, Moshe Harpaz and Oren
 162 Cytokinins delay litchi maturation
Yahalom for enabling use of their orchards. The
assistance of Alla Zvilling, Asia Gizis, Dan Gemarasni,
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