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Effect of CPPU application on fruit development and quality of different Actinidia genotypes

Authors:
  • Warsaw University of Life Sciences - SGGW, Institute of Horticulture Sciences
60 Piotr Latocha, Tomasz Krupa
EFFECT OF CPPU APPLICATION ON FRUIT
DEVELOPMENT AND QUALITY OF DIFFERENT
ACTINIDIA GENOTYPES
Piotr Latocha, Tomasz Krupa
Warsaw University of Life Sciences  SGGW, Warsaw, Poland
INTRODUCTION
Actinidia arguta (Siebold et. Zucc.) Planch. ex Miq., known as hardy kiwifruit,
has recently become more popular on the market as baby kiwi or grape kiwi [Wi-
liams et al. 2003]. A. kolomikta (Maxim. et Rupr.) Maxim., known as super hardy
kiwi, is widely used as ornamental plant and in some Eastern European countries, such
as Russia or Lithuania, also as fruit plant. The fruits of both species have a smooth, edible
skin and are smaller than the popular kiwifruit (A. deliciosa). They are very aromatic,
contain large amounts of sugar and ascorbic acid (up to 182 mg per 100 g fresh weight
for A. arguta and up to 1164 mg per 100 g fr.wt for A. kolomikta) [Chesoniene 2000,
Latocha 2006]. They also display an excellent flavour and have high antioxidant proper-
ties [Okamoto and Goto 2005, Krupa and Latocha 2007]. Hardy kiwifruit is just beginning
to be commercially grown in Canada, Chile, France, Japan, New Zealand and the USA
[Ferguson 1999, Wiliams et al. 2003]. The Oregon state (USA) currently produces the
largest crop in the world, with more than 40 ha of one cultivar, Ananasnaya [Strik and
Hummer 2006]. As the fruit weight is often smaller than 10 g, increasing fruit size should
be one of the strategies for increasing yield and marketability of this species.
The use of plant growth substances is one of possible methods of increasing fruit
size. Basically, fruit growth and its final size depend on the number of cells at the phases
of fruit set and following cell division that are under hormonal control and at early fruit
growth that is influenced by growth promoters, such as auxins, gibberellins and cytoki-
nins. CPPU (N-[2-chloro-4-pyridyl]-N`-phenylurea) is known to be effective in enhan-
cing fruit enlargement by stimulating cell division and cell expansion, mainly in outer
pericarp [Cruz-Castillo et al. 1997, Ben-Arie et al. 1998, Lewis et al. 1996] in many kinds
of fruits, including kiwifruit [Biasi et al. 1991, Costa et al. 1997, Cruz-Castillo et al. 2002].
So far, very little research has been carried out on A. arguta [Jo et al. 2003, Kim et al.
2006] and no research has been done on A. kolomikta or Actinidia hybrids.
MATERIALS AND METHODS
The experiment was carried out in the years 2006 and 2007, on four genotypes of
Actinidia  `Sentiabrskaya` (A. arguta), `Ken`s Red` (A. arguta `Cordifolia` × A. mela-
nandra), `Issai` (A. rufa × A. arguta) and `Dr Szymanowski` (A. kolomikta). Three of
61BIOSTIMULATORS IN MODERN AGRICULTURE  FRUIT CROPS
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SEICEPS kenutaG RAVITLUC anaimdO NIGIROTNALP niloreinezdohcoP EPAHSTIURF ucowot³atzsK HSELF
RUOLOC roloK uzs¿¹im
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these cultivars are also popular at the Polish horti-
cultural market. Description of the tested genoty-
pes, along with their flowering and ripening period
is shown in Table 1. All plants were 8-10-year-old,
trained for the T-bar system and were grown on
the experimental field of the Warsaw University of
Life Sciences, Central Poland. Six single-flowering
shoots (with minimum twenty flower buds) were
selected on each plant and marked just before flo-
wering. At the anthesis, all selected flowers were
hand-pollinated. Three of the marked shoots were
sprayed with the 15 mg·l-1 CPPU solution (Sigma-
Aldrich), fifteen days after petal fall (DAPF) ac-
cording to Jo et al. [2003]. The control shoots were
not subjected to any treatments. After the CPPU
treatment, the length and diameter (in two direc-
tions) of all fruits was measured at fortnightly inte-
rvals, until harvest. Berries were picked at the sta-
ge of commercial maturity. After harvest, they were
kept in a plastic box at 200C for 48 h, to enhance
ripening before physicochemical measurements.
After ripening, fruit weight, colour, seed number and
mass, firmness, the content of soluble solids (SSC)
and titratable acidity (TA) were determined. L* (li-
ghtness), a* (greenness to redness) and b* (blu-
eness to yellowness) were determined using spec-
trophotometer Minolta CM-508i. Calculated hue
angle (h) and chroma (c) were used for comparing
colour differences between samples.
The same fruits were used to measure the other
physicochemical parameters. Firmness was measu-
red by Instron 5542 with 5 mm tip and expressed in
Newtons (N). Each fruit was subjected to a com-
pression speed of 1 mm/s after contact and pene-
tration to 10 mm. Soluble solids content (SSC) and
titratable acidity (TA) in fruit juice were assayed
using an Abbe`s refractometer and by titration with
0.5 N NaOH to the endpoint 8.1 pH. Titratable aci-
dity was expressed as percentage of anhydrous ci-
tric acid equivalent. Three replications were com-
pleted for each parameter measured.
The obtained results were subjected to statisti-
cal analysis by the Statgraphics® Plus 4.1 program-
me, using the one-way analysis of variance.
62 Piotr Latocha, Tomasz Krupa
FIGURE 1. CHANGES OF FRUIT DIMENSIONS IN MOST SENSITIVE CULTIVARS AFTER CPPU
TREATMENT
Rysunek 1. Zmiany wymiarów owoców odmian najwra¿liwszych na CPPU, po traktowaniu
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RESULTS AND DISCUSSION
For all genotypes, the most intense fruit growth was noted within 40 days after petal
fall (Fig. 1). `Issai` and `Sentiabrskaya` increased rather in length than in diameter, whe-
reas `Ken`s Red` and `Dr Szymanowski` rather in the longer diameter than in length.
However, the proportion of the fruit did not change for most of the genotypes, except for
`Ken`s Red` in the first year. Similar results were obtained for kiwifruit by Antognozzi et
al. [1997]. According to Jo et al. [2003] significant enlargement of fruit size of A. arguta,
A. deliciosa and A. chinensis occurred in both longitudinal and cross section, whereas in
A. melanandra and A. polygama it happened mainly in the cross section. However,
LENGHT/d³ugoæ [mm]
LENGHT/d³ugoæ [mm]
LONGER DIAMETER/
wiêksza rednica [mm]
SHORTER DIAMETER/
mniejsza rednica [mm]
SHORTER DIAMETER/
mniejsza rednica [mm]
* DAPF  DAYS AFTER PETAL FAL L
* DAPF  dni po opadniêciu p³atków
LONGER DIAMETER/
wiêksza rednica [mm]
C.(165('C
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63BIOSTIMULATORS IN MODERN AGRICULTURE  FRUIT CROPS
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398.0 376.0 *
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7.72 2.52 *
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61.1 10.1 .s.n
6.81 6.71 *
609.0 357.0 **
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0.53 5.72 **
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9.91 7.51 **
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8.51 5.61 .s.n
556.1 629.1 *
29.3 36.3 .s.n
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34.1 54.1 .s.n
5.41 5.41 .s.n
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53.2 09.1 .s.n
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6.82 0.42 **
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52.1 53.1 .s.n
8.31 0.51 .s.n
480.1 561.1 .s.n
48.2 66.2 .s.n
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3.01 0.7 **
9.82 3.42 **
9.52 1.22 **
5.22 0.02 **
21.1 01.1 .s.n
2.71 5.61 .s.n
369.0 138.0 *
26.2 83.2 .s.n
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0.9 2.4 **
9.13 9.42 **
6.12 9.61 **
2.91 6.51 **
84.1 74.1 .s.n
3.31 2.41 *
011.1 521.1 .s.n
60.4 76.3 .s.n
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9.3 7.3 .s.n
2.52 1.62 .s.n
2.91 6.91 .s.n
6.21 7.21 .s.n
13.1 33.1 .s.n
9.21 2.21 .s.n
895.1 016.1 .s.n
95.3 60.3 .s.n
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fruits of A. macrosperma were not affected by CPPU at 16 mg·l-1. Some differences
within the species were also recorded.
Fruit weight at harvest was significantly higher after CPPU application for all geno-
types, except for `Dr Szymanowski` (A. kolomikta) (Tab. 2). The greatest increase of
weight was noted for Issai (104.5% on the average), and the lowest for `Dr Szyma-
nowski` (10%). For kiwifruit, the observed increase in mass was up to 40%, but better
results were obtained with CPPU and GA3 mixture  up to 66% [Cruz-Castillo et al.
1991]. Among the fruits from the treated shoots the largest were those of `Ken`s Red`
and the smallest those of `Dr Szymanowski` (on the average, 17.6 g and 3.6 g respective-
ly). According to Cruz-Castillo et al. [2002] also the fruit origin and effectiveness of
pollination may affect the final fruit mass.
The CPPU applications had very little influence on TA and SSC of the tested genoty-
pes. A significant increase of SSC at harvest was found only for `Sentiabrskaya` in the
first year and for `Issai` in the second year of testing (Tab. 3). In this study, significant
changes in TA were noted especially in the first year. TA of the fruits treated with CPPU
was slightly lower than that of the untreated ones for most genotypes. After the treat-
ment, only `Sentiabrskaya` showed significantly higher level of TA in both years of the
experiment. According to many studies carried out on `Hayward` kiwifruit, CPPU treat-
ments increased the SSC content and decreased the TA. As a result, the fruits ripened
earlier and their firmness decreased [Antognozzi et al. 1997, Famiani et al. 1998]. In our
study, a slight increase in firmness of the treated fruit was noted, it was not statistically
significant, however.
65BIOSTIMULATORS IN MODERN AGRICULTURE  FRUIT CROPS
PHOTO 1. EFFECT OF CPPU ON FRUIT SIZE IN CULTIVARS MOST SENSITIVE TO THIS
BIOSTIMULATOR
PHOTO: AUTHOR
Foto 1. Wp³yw CPPU na wielkoæ owoców odmian najbardziej wra¿liwych na jego dzia³anie
Fotografia: Autor
1 cm
Colour measurements of the tested Actinidia fruits are summarised in Table 3. In our
experiment, the colour changes appeared to be the most important, especially for `Ken`s
Red`, because this cultivar had a bronze-red fruit. A significant decrease in L* and chro-
ma parameters and an increase in a* and Hue angle was recorded. This suggests fruit
brightening, along with an increase of the red colour intensity, resulting from the CPPU
treatment. Lawes et al. [1991] indicated that CPPU significantly influenced the darkness
of fruit skin (L*) without any effect on the hue angle or chroma in the kiwifruit. In our
study, the most significant changes were observed in `Sentiabrskaya` (in all parameters)
and in `Dr Szymanowski` (the a* excepted), albeit they were not easy to notice. The
green fruit got darker and the colour became more vivid. There were no significant
changes in coloration of `Issai`.
The CPPU has not affected the seed number per fruit. This characteristic was,
however, related to the genotype. The species and cultivars, fruit mass of which incre-
ased the most (`Ken`s Red` and `Issai`) also showed a significant increase of the seed
mass.
`Sentiabrskaya` fruits developed the thicker petals after treatment (data not shown).
Skin damage at the side exposed to sunshine was also noted. This may suggest an incre-
ased sensitivity of the hairless skin due to the application of CPPU on this cultivar. Some
66 Piotr Latocha, Tomasz Krupa
deformations of flowers (mostly carpels and styles) and of fruit of `Dr Szymanowski`
were observed in the second year of study. It might be related to the application of CPPU
in the previous year. Other genotypes did not show any deformations. Other authors also
mentioned some deformations caused by CPPU application. Alina [1998] described the
deformation of apple fruit and decrease in the number of flowers in the second year after
treatment. As for kiwifruit, the possible deformation could appear in the distal part of a
fruit and it is believed that the high concentration of CPPU and early application could be
the main reasons of the deformations [Lawes et al. 1991]. According to these authors,
the best period for the applications is three weeks after anthesis. Similar results for A.
arguta `Mitsuko` were obtained by Kim et al. [2006], who also noticed that the fruit mass
was positively correlated with CPPU concentration only up to 5 mg·l-1. However, the
results obtained by Jo et al. [2003] on different Actinidia species did not indicate any
noticeable deformation of the treated fruits.
CONCLUSIONS
1. Different genotypes respond to CPPU application in a different way. This suggests a
genetic background of this reaction. CPPU may be effective for enhancing fruit
enlargement of A. arguta and some hybrids.
2. The greatest increase in fruit mass due to CPPU may be expected in `Issai` (A. rufa
× A. arguta) and `Ken`s Red` (A. arguta `Cordifolia` × A. melanandra), whereas
`Dr Szymanowski` (A. kolomikta) does not respond to the treatment Moreover, CPPU
tends to deformate flowers and fruits in this species. In case of `Sentiabrskaya`, the
biostimulator applied may cause the increase of skin sensitivity to sun scald.
3. The application of CPPU slightly modifies the other physicochemical characteristics
of fruit of the Actinidia genotypes, albeit- only some colour changes are notable
without any negative effects.
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commercialization of `Baby Kiwi` (Actinidia arguta Planch.). Acta Hortic., 610, 81-86.
... This enlargement was similar to those reported by Patterson et al. (1993) and Antognozzi et al. (2013), whereas it was smaller compared to those obtained when higher CPPU concentrations were used (Lawes et al., 1992;Antognozzi et al., 1997). There is evidence that the CPPU effect on kiwifruit depends mainly on CPPU concentration and application time (Biasi et al., 1992;Kim et al., 2006), but also on the Actinidia species and cultivars (Latocha and Krupa, 2007;Brown and Woolley, 2010) and on time of anthesis (Cruz-castillo et al., 2002). ...
... Moreover, CPPU did not affect the seed number of fruits which in fruits grown in absence of pollinators were at least 20 fold lower than of fruits of all other treatments. These findings are in accordance with Iwahori et al. (1988) and Latocha and Krupa (2007) who pointed out that in absence of pollinators, seed number reduced dramatically. All the above, indicate that good pollination conditions in field are required for kiwifruit production and that CPPU had no further effect on poorly pollinated kiwifruits -except fruit shape -when applied 15 days after fruit set. ...
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The influence of N-(2-chloro-4-pyridyl)-N'-phenylurea (CPPU) on 'Hayward' kiwifruits was evaluated in the field under different pollination conditions: (a) natural pollination, (b) assist pollination and (c) absence of pollinators. Residue dynamics of CPPU on kiwifruit revealed that half life values in all treatments did not exceed 6 days, although in absence of pollinators the dissipation rate was reduced. At harvest, CPPU treatment increased the weight and the size only of pollinated fruits compared to untreated pollinated ones. Absence of pollinators resulted to more soften fruits with lower seed number and Ca2+ content, while CPPU application resulted additionally to misshapen fruits. No differences were detected on fruit total soluble solids, titrable acidity and ascorbic acid. In contrast, phenolics were higher on kiwifruits grown in absence of pollinators irrespective of CPPU treatment and antioxidant capacity was significantly lower only on untreated with CPPU fruits under assist pollination. These results suggest that (i) under good pollination conditions, CPPU indeed contributes to superior product quality of kiwifruits, and (ii) a strong influence of seed number on Ca2+ accumulation and phenolic content probably exists as indicated in the case of fruits grown under limited pollination conditions. © 2015 Elsevier B.V.
... arguta), 'Ken's Red' (A. arguta 'Cordifolia' x A. melanandra), 'Issai' (A. rufa x A. arguta) and 'Dr Szymanowski' (A. kolomikta) were sprayed with CPPU (15 mg/l) solution (Sigma Aldrich), fifteen days after petal fall (DAPF) (Latocha and Krupa, 2008). 'Issai' and 'Sentiabrskaya' increased rather in length than in diameter, whereas 'Ken's Red' and 'Dr Szymanowski' rather in the longer diameter than in length. ...
Chapter
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Plant Growth Regulator in Kiwi fruit
... Some aspects of plant morphology and influence of central Poland climate conditions on the growth of different kiwiberry cultivars were also studied (Latocha, 2008). The possibility of fruit improvement by CPPU and Vapor Gard application and occurrence of Actinidia viruses in Poland were investigated (Latocha and Krupa, 2008b; Latocha et al., 2009; Sala-Rejczak et al., 2012). In 2006, Latocha published a book about growing kiwiberry (in Polish). ...
Conference Paper
Kiwiberry (Actinidia argute), a smaller cold hardy cousin of the well-known kiwifruit (A. deliciosa and A. chinensis), has always been considered of minor economic importance. Its twining, woody vines produce small, grape-size fruit with diversity of flesh and skin colours. With intense flavour, kiwiberries can be eaten whole, without peeling and therefore are a very convenient kind of food, let alone their health benefits. Since the 1980s, their cultivation has been introduced in several regions, and although some small plantations are still operating, they did not gain much commercial success. In recent years, the interest for this fruit crop has grown up in Europe, driven by the increase in scientific and technical knowledge of various aspects of the cultivation and the support of a new marketing approach. Scientific research in Belgium, Poland, as well as field experiences in Switzerland and commercial attempts in France and Italy focused on morphology, physiology and phenology, with practical results in various aspects of growing, pollination, harvesting, shelf life and storage. The decision of presenting this fruit to customers as a berry, with an identity completely different from the current commercial kiwifruit, is an important part of the fruit's commercial success. Very likely kiwiberry cultivation technology will be further developed in the near future, and new promising cultivars, obtained by breeding and selecting work recently conducted in various countries, will be introduced. Kiwiberry and kiwifruit follow two different paths, which creates new opportunities for a great future for kiwiberry.
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Terai region of northern India is having great potentiality for growing low chilling cultivars of different temperate fruits as this region receives sufficient amount of low temperature every year. However, the major problems of low chill cultivars are small fruit size, along with poor physico-chemical quality attributes. The present investigation was aimed at addressing these problems. For determining the influence of plant bio-regulators CPPU (5 and 10 ppm), NAA (10 and 20 ppm) and salicylic acid (50 and 100 ppm) were applied as once, twice and thrice spray at fortnightly interval after fruit set. CPPU @ 10 ppm was found most effective for increasing fruit length (6.79 cm), width (6.67 cm), fruit weight (182.84 g) and volume (179.11 ml), while TSS (13.38°Brix), total sugars (8.25%), acidity (0.55%), number of fruits per tree (285.00) and yield (47.66 kg) were found better with the application of CPPU @ 5 ppm. Higher per cent of marketable fruits (86.56) and ascorbic acid (7.08 mg/ 100 g) content was found in SA @ 100 ppm treatment.
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In 1993 and 1994, trials were carried out to optimize the use of CPPU on Actinidia deliciosa. CPPU treatments were carried out at different times (2, 3, 5 and 7 weeks after full bloom), using different concentrations (0, 0.5, 1, 2.5, 5, 10 and 20 ppm), utilizing both a manual sprayer and a tractor-mounted atomizer; also repeated CPPU applications.were tested. All CPPU treatments increased the fruit weight and consequently the yield per vine with respect to the control. The increase in fruit weight was positively correlated to CPPU concentration up to 5 ppm, thereafter higher CPPU concentrations did not produce any further significant increase in weight. CPPU application greatly influenced fruit dry weight which was negatively correlated to the time of CPPU application. The repeat application of CPPU caused an increase in fruit fresh weight similar to the single application made with a concentration equal to the sum of the two used for the repeat application, but it resulted in a lower fruit dry weight. CPPU increased the percentage of fruits with a protruding distal end, with an intensity positively correlated to CPPU concentration; moreover, at the higher concentrations it increased the maximum/minimum diameter ratio of the fruit. Elimination of dripping during CPPU application, by use of an atomizer, reduced the percentage of fruits having a protruding distal end. The earliest CPPU treatment caused a rapid increase in soluble solids and a decrease in flesh firmness with respect to the control, resulting in early fruit ripening. During cool storage in normal atmospheric conditions (0±0.5 °C, R.H. < 95%), the soluble solids in the fruits increased in all the treatments and after 5 months of storage they were consistently higher in the control and early treated fruits. At the same time flesh firmness decreased with late treated fruits having lower values. The results indicate that with a correct use of CPPU in terms of application time, amount and modality of distribution it is possible to combine the positive influence of CPPU on fruit size and yield per vine with lack of negative effects on shape, quality and storability of fruits, by treating early and using relatively low concentrations of CPPU.
Book
Opis rodzaju i poszczególnych gatunków aktinidii. Charakterystyka odmian. Opis walorów owoców. Opis metod uprawy, zbioru i przechowywania owoców.
Article
The time of phenological changes, growth characteristics, winter hardiness and fruiting potential of 1-2 and 3 year-old Actinidia kolomikta (Rupr.) Maxim. plants have been investigated in Kaunas Botanical Garden. The differences in the time of bud burst, blooming, shoot growth and leaf fall comprise only 2-3 days for all of the cultivars studied, while the harvest maturity differs more considerably. Evaluation of economically important plant properties (growth, length of shoot, number of fruiting shoots, winter hardiness) allows to characterise the cultivars. The biggest part of berry yield is produced on the short (up to 15 cm) flowering shoots. Negative correlation indicates that with an increasing number of long (mixed) shoots the berry yield decreases. The cultivar 'Paukshtes Shakarva' is distinguished by its berry size (25.4×14.5mm) and mass (3.35 g). However, the highest contents of ascorbic acid is observed in 'VIR-1' and 'Pavlovskaja' - 1164.8 and 955.7 mg%, respectively. Except for one-year-old plants of cultivar 'Lande', which showed rather medium winter hardiness, other investigated cultivars wintered poorly. However, the two-year-old plants were sufficiently hardy.
Article
Berry size of seedless grapes is generally increased by application of GA3 sprays at the time of fruit set. For certain cultivars a number of applications is required to obtain a commercially acceptable sized berry, and this may have deleterious effects on fruit bud initiation for the following year. It has been found that a similar increase in berry size can be obtained by a single application of CPPU to Perlette, Superior and Thompson Seedless cultivars As with GA3, fruit ripening and maturation were delayed following CPPU application, sometimes even more severely than with GA3. However, other aspects of development of the cluster were affected differently by each growth regulator. Such aspects include the growth of the rachis and pedicels, berry shatter after harvest and susceptibility of the berry to fungal attack and decay. These differences may be attributed to the anatomical effects of the regulators, in that CPPU increased the number and density of cells, whereas GA3 enhanced cellular expansion and decreased cell density. Each growth regulator also increased the size of seeded berries of cultivar Zeiny and fruit ripening and resistance to decay were similarly affected. When a combination of both compounds was applied, there were no additive effects on berry size, berry firmness and fruit ripening, but the relative resistance to decay conferred by CPPU, was eliminated by addition of GA3. Anatomical observations showed that the cell area and skin thickness of berries treated with both regulators were the same as for berries treated with GAR alone.
Article
There exist about 61 species of Actinidia around the world and most are originated from China. Among them, 3 species, A. deliciosa, A. chinensis and A. arguta, are commercially cultivated, although A. arguta at a minor extent. Most Actinidia fruits are relatively small but some have very impressive horticultural advantages for commercialization, i.e., vitamin content, excellent flavor, sweetness and other. Several reports were recently published on CPPU [N-(2-chloro-4-pyridyl)-N'- phenylurea, Fulmet], a diphenylurea-derivative cytokinin-like compound that stimulates fruit development. However, they all focused on A. deliciosa cv. 'Hayward'. We investigated several Actinidia species for their CPPU sensitivity . Fruit of A. macrosperma were not affected by 16 ppm CPPU and fruit of A. polygama and A. melanandra showed a little increase only in cross diameter, but not in length. Other species, like A. chinensis and A. deliciosa, were rather sensitive to CPPU, and differences within species were also recorded. More details are discussed on fruit development, fruit quality and other features in the paper.