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Effect of pollination on dropping of flowers and fruits in new quince (Cydonia oblonga Mill.) cultivar and promising genotypes
Scientia Horticulturae 231 (2018) 126–132
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Effect of pollination on dropping of flowers and fruits in new quince (Cydonia oblonga Mill.) cultivar and promising genotypes
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Maryam Tataria, , Hamid Abdollahib, Asghar Mousavic a Horticulture Crops Research Department, Isfahan Agricultural and Natural Resources Research and Education Center, Agricultural Research, Education and Extension organization (AREEO), Isfahan, Iran b Temperate Fruits Research Center, Horticultural Sciences Research Institute, Agricultural Research, Education and Extension Organization (AREEO), Karaj, Iran c Horticulture Crops Research Department, Chahar-Mahal va Bakhtiari Agricultural and Natural Resources Research and Education Center, Agricultural Research, Education and Extension organization (AREEO), Shahrekord, Iran
A R T I C L E I N F O
A B S T R A C T
Keywords: Quince (Cydonia oblonga) Fruit set Pollen source Pollination Self-incompatibility
Access to sufficient and good quality fruits and solving problems such as early abscission of the fruitlets depends on the status of self-fertility and self-incompatibility in cultivars and genotypes and determination of the best pollinizer for them. This research was carried out to study the best pollinizer for some quince cultivars and promising genotypes from the collection of germplasm at the Horticultural Research Station of Isfahan, Iran in 2015 and 2016. For this, the effects of the pollen donor parent, including ‘Vidoja’, KVD2, KVD4, ‘Isfahan’, ‘Torsh’ and also open pollination were studied on quantitative and qualitative characteristics of the pollen recipient parent, including ‘Vidoja’, KVD2 and KVD4 as a factorial experiment in a randomized complete block design with four replications. To determine the best pollinizer, flowers of pollen recipient parents were emasculated at the balloon stage and were covered with isolation bags. Emasculated flowers were crossed with the pollen of donor parents during receptability of stigmas as well as hand self-pollination for each female parent. Results showed that some traits such as percentage of fruit set, seed numbers and total soluble solids (TSS) were affected by the pollen source. Phenol and pectin were not affected by the foreign pollen sources. According to the results, self-pollination did not have acceptable fruit set. The best pollen source for the ‘Vidoja’ was KVD2, for KVD4 was ‘Torsh’ and ‘Vidoja’ and for KVD2 was KVD4 and ‘Vidoja’, therefore the cultivation of all three female parents together in the quince orchards is recommended.
1. Introduction Quince (Cydonia oblonga L.) is economically the third fruit of pome fruit group. This species is native of Iran and its distribution centers are the northern forests of this country (Sabeti, 1995). This fruit was cultivated for 4000 years B. C. in Western Asia and it was favorite for many people in Greece and Rome and was called Mala cydonia. The scientific name of this tree has been described to the Sidon city, that had a lot of quince trees (Sabeti, 1995; Maniei, 1995). Quince is cultivated as extensive orchards in different provinces of Iran and area harvested has increased compared to previous years (FAO, 2014). One of the main problems for quince growers is abscission of flowers and fruits in the early stages of growth that reduce the yield and economic efficiency. Several factors, including the quantity and quality of water, soil quality, pruning, nutrition and environmental conditions before and after flowering can affect on abscission of flowers and fruits and ultimately reduction of yield. In the same condition of these factors,
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Corresponding author. E-mail address: [email protected] (M. Tatari).
https://doi.org/10.1016/j.scienta.2017.10.045 Received 18 June 2017; Received in revised form 9 October 2017; Accepted 31 October 2017 0304-4238/ © 2017 Elsevier B.V. All rights reserved.
the most important factor in fruit bearing is compatibility and fertilization (Ortega and Dicenta, 2004). Self- incompatibility or incomplete pollination of flowers is as one of the main causes of the fruit dropping (Boskovic and Tobutt, 2001). Compatible pollen grain, sufficient pollen transfer at the right time on the stigma and pollen tube growth in the style and finally fertilization are essential for successful production of fruit. The absence of pollination vectors or inappropriate weather conditions also cause problems in pollen transfer. In self-incompatible cultivars, pollen tube was not able to grow in the style and reach to ovule, so suitable pollinizer cultivars are essential to achieve economic yield (Socias I Company et al., 2004). Pollinizer cultivars should be compatible with main cultivar and have overlapping in flowering time. Therefore, weak pollination is as a limiting factor in crop production in the various regions (Alizadeh et al., 2009). Quince is introduced as a self compatible tree in the most references (Maniei, 1995), but based on the presented evidence about self-
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‘Torsh’ and ‘Isfahan’ cultivars along with three female parent (‘Vidoja’, KVD2 and KVD4) were considered as the pollen donor parent for pollination studies. Characteristics of the recipient and donor parent is as follows. Vidoja is early-flowering with a high density of the flowers, spore type, dwarf and without alternate bearing. Its fruit size is medium and uniformity, juicy, fragrant with low astringency and desirable quantitative and qualitative characteristics. KVD2 is early-flowering and high flowers density, spore type, dwarf, high yielding, early fruiting and without alternate bearing, fruits with medium to large size, uniform, juicy, fragrant with good quality properties. Characteristics of KVD4 are included early flowering and fruiting and without alternate bearing, spore type, dwarf, high yielding, large fruit and almost juicy with great aroma and sour taste. Isfahan cultivar is Late flowering and fruiting, without alternate bearing, invigorating, intermediate yields, large fruit, juicy, fragrant with low astringency and good quality. Torsh cultivar is early-flowering and fruiting, intermediate vigor, intermediate yields. Its fruit size is medium to large and almost juicy. The opening of 10%, 80% and 95% of flowers were considered as the beginning of flowering, full bloom and the end of flowering stages, respectively (Gharaghani et al., 2009).
incompatibility alleles on quince, there is a question that the use of cross pollination, what can be effective in increasing fruit in different cultivars of this species (Akbari Bisheh et al., 2016). More species in Rosaceae family have self-incompatibility, that's controlled by a locus with multiple S alleles (Zhang and Xue, 2008). In self-pollination aspect, quince cultivars are classificated into four groups, including the self-compatible cultivars with more than 10% of fruit set, semi selfcompatible cultivars with fruit set 3–8%, self incompatible cultivars with 1–2% of fruit set and completely incompatible cultivars with fruit set less that 1% (Nagy-Deri et al., 2013). Based on this classification, if semi self-incompatible cultivars are cultured next to pollinizer cultivars, they will produce more yield. The semi-self incompatible and completely self-incompatible cultivars need to compatible pollinizer cultivars for the appropriate fruit set. Therefore, the self-incompatibility percent of cultivars should be investigated in order to have sufficient yield. There are several methods to study the compatibility or incompatibility of different cultivars and determination of suitable pollinizer for them. These methods include controlled pollination, observation of pollen tube growth with a fluorescence microscope, extraction of ribonuclease from style, specific PCR of S allele and nucleotides sequencing related to incompatibility (Ortega and Dicenta, 2004; Mousavi et al., 2014). As regards, the controlled pollination method allows to estimate the performance of several cultivars in the orchard, so this method is recommended to determine the appropriate pollinizer (Rasouli et al., 2009). Nuzzo and Rubbi (2004) studied 22 quince cultivars in Italy and reported that most of them were self incompatible that required cross pollination to yield production. A number of cultivars have also been compatible that use of pollinizer was preferably necessary to obtain better yield. In order to study of the self-incompatibility and determination of the best pollinizer for ‘Isfahan’ commercial cultivar, female parent was pollinated with four genotypes that named KM1, PK3, KVD2 and NB4. KM1 genotype was introduced as the best pollinizer for ‘Isfahan’ with 80% of overlapping in flowering time and 23.42% of fruit set (Akbari, 2014). According to Talaei et al. (2007) fruit set percentage after the self-pollination was less than cross pollination. In the study of pollination status in 23 quince cultivars in various regions of Yugoslavia, it was found that only five cultivars of them are self-pollinated (Ershov, 1989). Pollination and fertility are one of the important issues in the quince trees. Some quince cultivars and genotypes, like other relatives in the Rosaceae family, including apples and pears, have self-incompatibility S-alleles. Self- incompatibility or incomplete pollination of flowers leads to fruitlet dropping and reduce yield, Therefore, it's necessary to be examined self or cross compatibility percent of any new cultivar with the introduction of it. Fourteen genotypes were identified and collected during the collection program of Iranian quince cultivars and genotypes in Isfahan, Iran. KVD1, KVD2 and KVD4 genotypes have been proposed as promising genotypes after preliminary evaluation of them. KVD1 genotype was released in the name of ‘Vidoja’. The aims of this study were the determination of the pollination status and suitable pollinizer for these genotypes and cultivar with the same time flowering period to achieve high performance.
2.2. Collection and viability test of pollen grains In order to collection of pollen grains, some branches of donor parents contain a sufficient number of flowers that are at the balloon stage were cut and transported to the laboratory. Brunches were placed in the sugar solution (4%). After opening flowers, the anthers were separated from flowers and were placed on glossy paper at room temperature (26–24°C) for 24–48 hours to be losing their moisture and drying of anthers. Pollen grains were separated from anthers and were collected in a sterile glass container with cotton cap. In order to study of viability, pollen grains were distributed in a petri dish containing medium with 1% agar, 15% sucrose and 50 mg/L of boric acid (Dalkilic and Osman, 2011) and germination percentage of pollen grains were recorded using a light microscope after 24 h exposure to laboratory temperature. Pollens with more than 75% of germination percentage were selected and were used as a pollen source. Sterile glasses containing pollen grains were stored in cold and dry condition at 4°C until pollination time.
2.3. Pollination With swelling of buds, suitable branches containing flower buds and uniform were randomly selected in the four geographical directions of each tree. Flowers were emasculated at the balloon stage and weak, immature and opened flowers as well as flower buds were removed from selected branches and were covered with cloth bags. After receptibility of stigma to accept pollen grain, each isolated branch was brought out of the bag and emasculated flowers were pollinated with considered pollen using a glass rod for each crosses. Pollinated flowers were counted and labeled and again were covered with cloth bags. To ensure of pollination, flowers were again pollinated with considered pollen 24 h later. Open pollination was evaluated as control treatment, while natural (only covered flower) and hand self-pollination was used for the study of self-pollination treatment. Self-pollination was carried out after emasculation by manually as hand self-pollination treatment. For each crosses, 100 flowers were emasculated and pollinated. These flowers were counted and labeled. For self-pollination treatment also 100 flowers were counted at the balloon stage and were covered with cloth bags without emasculation or any crosses. Cloth bags were removed after petal fall to allow the flowers to place under normal conditions.
2. Material and methods 2.1. Plant materials and flowering period This research was carried out at the Horticultural Research Station of Isfahan, Iran in 2015 and 2016. For this purpose, two promising quince genotypes called KVD2 and KVD4 along with ‘Vidoja’ cultivar that recently has been introduced were considered as the female parent.
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genotypes was recorded to determine overlapping in flowering time of them in 2015 and 2016 (Table 1). According to the results in Table 1, ‘Torsh’ cultivar in both years has been the very early flowering. Then ‘Vidoja’, KVD2 and KVD4 were early-flowering. ‘Isfahan’ was the late flowering that had not overlapping in flowering time with ‘Torsh’. This cultivar had overlapping in flowering time from three to four days with the ‘Vidoja’, KVD2 and KVD4. Results showed that flowering in the second year (2016) was sooner than the first year (2015). As regards quince is one of the latest flowering species, spring frost occurs rarely in Isfahan, Iran.
2.4. Measurement of traits In this study, some traits were measured, including the percentage of fruit set at harvesting time, the number of seeds per fruit, fruit dimensions, fruit weight, total soluble solids (TSS), firmness, titrable acidity (TA), total phenol and pectin content of fruit. The fruit set percentage was obtained by comparing the number of fertilized flowers with the number of final fruits. Fruit length, weight and the most fruit diameter were measured using ruler, scales and caliper respectively. Fruit firmness was measured by a penetrometer (model EFFEGI, Italy, plunger diameter 11.1 mm, depth 7.9 mm), at opposite peeled sides and expressed as kg/cm2. Total soluble solids (TSS) were determined by extracting and mixing two drops of juice from the two cut ends of each fruit into a digital refractometer (ATAGO N-1α, Japan) at 22°C. Titrable acids were determined in 10 g of pulp samples by titration of extracted juice with sodium hydroxide (0.1 mol/L) up to pH=8.1 and expressed as a percent of malic acid. Pectin content was measured according to Thakur et al. (1996). Briefly, 100 g of fruit tissue was grated and 400 mL of distilled water was added and was boiled for an hour. After passing through the filter paper, 300 mL of distilled water and 10 mL NaOH was added and the resulting solution was kept overnight at room temperature, then 50 mL acetic acid (1 mol/L) and 25 mL of calcium chloride was added to the solution. The resulting solution was kept for one hour at room temperature and then was boiled for an hour. Boiled solution was passed through filter paper. Difference between initial and secondary weight of filter paper was reported as pectin weight based on grams per 100 g of fruit pulp. Total phenols were measured in fruit Juice using Folin-Ciocalteu (Singleton and Rossi, 1965). Absorbance of the samples was determined at 765 nm wavelength with spectrophotometer model T80 UV/Visible, then compared with the standard of gallic acid and expressed as mg gallic acid per 100 g of fresh weight.
3.2. Pollen grain germination The results of pollen grain culture and viewing them under a light microscope showed that the pollen grain germination of ‘Torsh’, KVD4, KVD2, ‘Vidoja’ and ‘Isfahan’ was 84, 84, 85, 88 and 80 percent, respectively. 3.3. Fruit set percentage and seed number Analysis of variance for effect of recipient and donor parents on measured traits showed in Table 2. The highest percentage of fruit set for ‘Vidoja’ was obtained with pollen grains of KVD2 genotype in the first year with an average of 18.75%. In KVD4 genotype, pollen of ‘Torsh’ and ‘Vidoja’ produced the highest percentage of fruit set with an average of 28.64 and 18.54%, respectively. KVD2 with pollen of KVD4 showed fruit set equal to 30.28%. This genotype in cross pollination with pollen of ‘Vidoja’ had 19.86% of fruit set. Pollen sources of ‘Isfahan’ and ‘Torsh’ had little fruit set, especially for ‘Vidoja’ and KVD2 female parents. Generally, the fruit set percentage in the second year was less than the first year (Table 3). The lowest percentage of fruit set in all female parents was belonged to self-pollination treatments. Natural self-pollination in KVD4 and ‘Vidoja’ did not produce any fruit in both years. Fruit set in self-pollination of KVD2 was 3.47%. and hand self-pollination had also a very low fruit set percentage. According to the results (Table 4) the highest seed number was observed in KVD2 that pollinated with ‘Vidoja’ and KVD4. The least seed number was belonged to self-pollination treatments and especially natural self-pollination.
2.5. Statistical analysis Obtained results were compared using factorial experiment based on a randomized complete block design with four replications. As regards, the experiment was carried out in two years, compound analysis was also carried out to analyze the effect of the year. Analysis of data was performed by ANOVA method using statistical software SAS (version 9.1) and means comparison using the LSD test. Data normalization was carried out for all traits with ASIN formula.
3.4. Fruit dimensions and weight In cross-pollination of ‘Vidoja’ and KVD4 with other pollen sources as well as open pollination, fruit length and diameter was more than hand self-pollination. Self-pollination in these two cultivar and genotype did not produce any fruits. Fruit length and diameter in KVD2 genotype when crossed with ‘Isfahan’, ‘Vidoja’ and KVD4 as well as open pollination was more than in cross with ‘Torsh’ cultivar and selfpollination or hand self- pollination (Table 4).
3. Results 3.1. Flowering period The beginning of flowering and flowering period of cultivars and
Table 1 Flowering time of quince cultivars and promising genotypes in 2015 and 2016.
March- April 27
28
29
30
31
1
2
3
4
5
6
7
Vidoja (2015) Vidoja (2016) KVD2 (2015) KVD2 (2016) KVD4 (2015) KVD4 (2016) Isfahan (2015) Isfahan (2016) Torsh (2015) Torsh (2016)
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8
9
10
11
12
13
14
15
16
17
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Table 2 ANOVA for effect of donor and recipient parents on measured characteristics Source of variation
Year Rep (year) Recipient parent Donor parent Donor×Recipient Recipient×Year Donor×Year Donor×Recipient×Year Error Total C.V.% SOV
df
1 4 2 6 12 2 6 12 80 125 – df
Mean square Fruit set (%)
Seed number
Fruit length (cm)
Fruit diameter (cm)
Fruit weight (g)
19.03** 0.16 1.92** 14.76** 4.58** 0.37ns 1.43** 0.71** 0.26 – 15.38
9.17ns 154.56 70.16ns 361.98** 429.5** 18.62ns 15.5ns 10.41ns 62.36 – 18.09
0.006ns 0.25 2.06** 4.48** 0.55* 0.05ns 0.04ns 0.02ns 0.24 – 15.22
0.002ns 0.19 1.12** 4.32** 0.53* 0.04ns 0.03ns 0.02ns 0.22 – 17.74
0.15ns 12.33 49.8** 163.15** 21.54* 1.34ns 1.13ns 0.84ns 9.15 – 19.24
Mean square TSS (%)
Year Rep (year) Recipient parent Donor parent Donor×Recipient Recipient×Year Donor×Year Donor×Recipient × Year Error Total C.V.% **,* and
1 4 2 6 12 2 6 12 80 125 -
10.82** 1.27 2.15ns 14.58** 2.42** 0.11ns 0.21ns 0.18ns 0.81 – 16.86
Firmness (kg/cm2)
TA (%)
ns
ns
1.82 0.02 0.33** 0.2** 0.06** 0.47** 0.01ns 0.02ns 0.01 – 11.77
0.003 0.14 0.21ns 2.36** 0.32** 0.05ns 0.01ns 0.04ns 0.12 – 16.31
Pectin (g/100g) ns
0.0004 0.003 0.03** 0.02** 0.008** 0.002ns 0.001ns 0.001ns 0.002 – 5.44
Total phenol (mg/100gFW) 0.01ns 9.42 36.67** 79.63** 23.72** 2.55ns 1.65ns 1.53ns 4.37 – 17.69
ns
: Significant at the 1% and 5% probability level and non-significant difference respectively
3.6. Fruit firmness, total phenol and pectin content
Hand self-pollination in ‘Vidoja’ and KVD4 produced the lowest fruit weight while other pollen sources produced higher levels of fruit weight. Also, self pollination did not produce any fruit. According to the results, Fruit weight of KVD2 by hand self-pollination, self-pollination and cross pollination with ‘Torsh’ cultivar was less than that of KVD2 pollinated with other pollen sources (Table 4).
According to the standard errors related to fruit firmness in the ‘Vidoja’, firmness was not influenced by the type of pollen. In KVD2 and KVD4 genotypes, self pollination had less fruit firmness for these genotypes (Fig. 2). Different sources of pollen had no significant effect on the pectin and total phenol content in fruits of all three female parents. Lack of fruit set in some treatments caused significant difference between the crosses in these two traits. Regardless of the pollen source, KVD2 and KVD4 genotypes had more pectin in their fruits. Most of phenol was observed in KVD4 genotype (Table 4).
3.5. TA and TSS According to the results in Table 4, the highest TA was found in KVD4 that it is characteristic of this genotype. Type of pollinizer had no significant effect on TA in this genotype and no fruit production in natural pollination led to significant differences for this trait. Similarly, for ‘Vidoja’ and KVD2 also the TA was not affected by pollinizer cultivar. Interaction effect of the year and female parent on the TA percentage (Fig. 3) showed that ‘Vidoja’ and KVD4 in the first and second year had not significant difference, while in KVD2 genotype, TA in the second year was more than the first year. Pollen source had no significant effect on the TSS of ‘Vidoja’ fruits and TSS in hand self-pollination fruits were almost similar to TSS in other treatments. In this cultivar, natural self-pollination did not produce any fruit. In KVD2 genotype, natural and manual self-pollination as well as pollen of ‘Torsh’ cultivar produced less TSS than other pollinizers. The use of ‘Vidoja’ and ‘Isfahan’ pollen was led to the production of fruits with higher TSS. In KVD4, hand self-pollination treatment produced fruits with less TSS (Fig. 1). Regardless of the cultivar, the effect of year on TSS was significant, so that TSS in the second year was more than the first year (Fig. 4).
4. Discussion 4.1. Fruit set percentage The fruit set percentage depends on pollinizer cultivar, year of experiment and used method of pollination (Denisow, 2003). The highest fruit set percentage was obtained in ‘Vidoja’, KVD2 and KVD4 with pollen of KVD2, ‘Torsh’ and KVD4, respectively. ‘Vidoja’ was also suitable pollinizer for KVD2 and KVD4 genotypes. The desirability of open pollination was less than other pollinizers (Table 3). The number of the transferred pollen grains to the stigma can have positive or negative effects on fruit set in open pollination compared with controlled crosses. Transferred pollen grains to the stigma is more than 50 in open pollination while, this would be a few hundred pollen grains at the controlled crosses (Stosser et al., 1996). Minimum of pollen grains requires for pollen germination and fruit set. When the concentration of pollen is high, the increase of inhibition in pollen tube growth can lead
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according to the density of the flowers on quince trees is less than other trees, so the conversion of bloom to fruit in the range of 20–25 percent is desirable. They studied fruit set and fruit dropping of quince cv. ‘Ekmek’ in Van, Turkey and reported that the final fruit set in this cultivar during two years was 12.3% and 8.6%, respectively. Akbari (2014) introduced KM1 genotype as the best pollinizer for ‘Isfahan’ cultvar with 23.42% of fruit set. The effect of the year on fruit set with changing in the temperature, humidity, insect activity and the fruit orchard management conditions can be changed the fruit set percentage. According to Ortega and Dicenta (2004) different factors affect the amount and percentage of fruit set in various years. Some of these factors include the amount and quality of water, soil quality, pruning, fertilization and environmental conditions before and after flowering. In this study, it appears that spring frost can involve in less fruit set in the second year.
Table 3 Mean comparison of year, donor and recipient parents on fruit set percentage ± SD Year
Recipient parent
Donor parent
Fruit set (%)
2015 2015 2015 2015 2015 2015 2015
Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja
Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
0.5uv ± 0.5 4.73o ± 1.41 0v ± 0 18.75d ± 2.64 10.1g ± 4.16 11.47f ± 3.51 6.51mn ± 2.79
2015 2015 2015 2015 2015 2015 2015
KVD2 KVD2 KVD2 KVD2 KVD2 KVD2 KVD2
Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
3.07pqr ± 2.99 3.83p ± 1.89 19.86c ± 3.96 3.47p ± 3.28 30.28a ± 2.63 2.32rs ± 1.09 2.5qrs ± 2.34
2015 2015 2015 2015 2015 2015 2015
KVD4 KVD4 KVD4 KVD4 KVD4 KVD4 KVD4
Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
0.58uv ± 0.52 9.66gh ± 3.5 18.54d ± 2.01 7.82k ± 2.49 0v ± 0 14.69e ± 3 28.64b ± 3.04
2016 2016 2016 2016 2016 2016 2016
Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja
Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
0.58uv ± 0.38 3.16pq ± 2.56 0v ± 0 8.73ij ± 3.37 5.69n ± 1.86 7.36kl ± 3.59 3.78p ± 1.41
2016 2016 2016 2016 2016 2016 2016
KVD2 KVD2 KVD2 KVD2 KVD2 KVD2 KVD2
Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
1.73st ± 1.49 3.48p ± 2.81 8.09jk ± 1.68 3.82p ± 3.15 6.71lm ± 1.12 1.16tu ± 0.45 0.86u ± 0.93
2016 2016 2016 2016 2016 2016 2016
KVD4 KVD4 KVD4 KVD4 KVD4 KVD4 KVD4
Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
0.64uv ± 0.55 3.29pq ± 2.56 9.04hi ± 077 6.73lm ± 1.62 0v ± 0 7.42kl ± 1.007 10.04g ± 2.26
4.2. Seed number and fruit dimension and weight The lowest of seed number was obtained from hand self-pollination treatment (Table 4). Fruits resulting from pollination with pollen sources that had less compatibility with female parent and fewer fruit set, had almost lower seed number. Similarly, Matsumoto et al. (2012) also reported that the number of seeds per fruit was reduced semicompatible crosses. All of the studied quince fruits that were resulting from controlled crosses had some seeds in their fruits at harvesting time. This indicates that there are not parthenocarpy in the quince and fertilization is necessary for the fruit set, growth and development. Not parthenocarpy in ‘Isfahan’ cultivar have also been reported by Akbari Bisheh et al. (2016). Fruit length, diameter and weight resulting from controlled crosses followed a similar pattern (Table 4). According to the results, pollination with other pollen sources produced larger and heavier fruits than natural or hand self-pollination. For KVD2 exceptionally, ‘Torsh’ cultivar as well as self-pollination treatment could not produce fruits with the appropriate dimensions and weight. Matsumoto et al. (2012) did not observe a positive correlation between fruit weight and number of seeds in the apple, but Keulemans et al. (1996) reported this relationship in apple cv. ‘Fuji’ that is consistent with our results. 4.3. Other traits
Similar letters in each column indicate no significant difference at the 5% level of LSD test
Although the pectin content, total phenolics and TA were varied among female parents, but these traits were not affected by cross pollination in all three female parents and the lack of fruit set in natural self-pollination treatment was leds to a significant effect of pollinizers on these traits that has been shown in Table 4. Pollen source had no any significant effect on the fruit TSS in ‘Vidoja’ cultivar. Self-pollination was produced less TSS in the fruits of KVD4 genotype. In KVD2 genotype, the highest TSS were obtained from pollination with pollen source of ‘Vidoja’ (19.5%), ‘Isfahan’ (18.75%), open pollination (16.91%) and KVD4 (16.73%), respectively (Fig. 1). Naturally,‘Vidoja’ and ‘Isfahan’ has higher TSS than other genotypes, so it can be concluded that self pollination in these cultivars maintains TSS at a high level, while in the other genotypes which have less TSS, cross pollination with other pollen sources leading to increase of TSS in comparison with self-pollination. KVD2 genotype produced fruits with higher TSS when pollinated with ‘Vidoja’. The effects of pollinizer on TSS have been reported in apple cv. ‘Golabe Kohanz’ and ‘Shafiabadi’ by Talaei et al. (2007) and in sweet cherry by Rasouli and Arzani (2011). According to the results (Fig. 3), more TSS was observed in the second year. More TA was also found in KVD2 genotype in this year (Fig. 4). Due to the annual decrease in rainfall and less irrigation water in the second year, it seems the fruit juice in this year was less than first year, so the concentration of cell sap increased in the second year and
to increase of competition among pollen tubes (Beyhan and Karakas, 2009), but in this study, it seems controlled cross had not a negative effect on pollen concentration and fruit set that were according to the results of Beyhan and Karakas (2009). Natural self-pollination did not lead to the final fruit set in ‘Vidoja’ and KVD4. Fruit set percentage in self-pollination of sweet cherry cv. ‘Zarde Daneshkadeh’ was also equal to zero, so this cultivar was introduced as self-incompatible (Rasouli et al., 2010). Similarly Sadat (2009) studied some European pear cultivars including ‘Spadona’, ‘Beurre Bosc’, ‘Louise Bonne’, ‘Coscia’, ‘Shah Miveh’, ‘Shahak’, ‘Sard Roud’ and a genotype of ‘khoj’ and reported that all of them were completely self-incompatible. Hand self-pollination led to low fruit set percentage. There was not difference between hand and natural selfpollination in fruit set of KVD2 genotype. Reduce of fruit set percentage in self- pollination treatment have also been reported by other researchers (Ershov, 1989; Deri, 2013; Talaei et al., 2007). The percentage of fruit set in cross pollination with ‘Isfahan’ cultivar was not remarkable compared to other pollen sources. The percentage of fruit set in ‘Vidoja’ and KVD2 was low with ‘Torsh’ pollen grains. Salleles can also be involved in self-incompatibility or semi self-compatibility of cultivars and genotypes. Benedek et al. (2001) stated that 130
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Table 4 Mean comparison of donor and recipient parents on measured characteristics ± SD Recipient
Donor
Seed Number
Fruit length (cm)
Fruit diameter (cm)
Fruit weight (g)
Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja KVD2 KVD2 KVD2 KVD2 KVD2 KVD2 KVD2 KVD4 KVD4 KVD4 KVD4 KVD4 KVD4 KVD4
Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
12.33de ± 3.7 11.33de ± 2.16 – 24.16abc ± 4.23 20.33abcd ± 3.56 15.33cde ± 2.32 23abc ± 4.87 11.83de ± 4.41 13de ± 2.6 25.33ab ± 3.11 10.16e ± 4.1 27.66a ± 5.96 23abc ± 5.58 12.83de ± 5.45 10.66e ± 4.66 17.66bcde ± 3.28 23.66abc ± 3.29 20abcd ± 2.86 – 23.5abc ± 4.26 24.16abc ± 3.16
3.61ij ± 1.86 4.65fgh ± 0.41 – 4.68fg ± 0.63 4.28gh ± 0.36 4.86fg ± 0.74 5.08f ± 0.84 4.22gh ± 0.77 7.18ab ± 0.7 7.71a ± 0.59 4.09hi ± 0.99 6.97bc ± 0.23 7.41ab ± 0.28 4.95f ± 0.85 3.06j ± 0.39 5.89e ± 0.35 5.89e ± 0.26 6.54cd ± 0.7 – 5.93e ± 0.39 6.1de ± 0.42
3.88f ± 0.96 4.94cde ± 0.44 – 5.07cd ± 0.51 4.52e ± 0.23 5.44c ± 0.63 5.11cd ± 0.48 3.46fg ± 0.72 6.7a ± 0.74 6.96a ± 0.48 3.85f ± 0.99 6.67a ± 0.24 6.61ab ± 0.53 4.61de ± 0.59 2.94g ± 0.31 5.43c ± 0.46 5.36c ± 0.31 6.12b ± 0.69 – 5.38c ± 0.27 5.41c ± 0.3
92.64k ± 11.92 118.07g ± 6.58 – 120.72g ± 5.97 114.09h ± 1.57 120.66g ± 3.82 121.33g ± 7.66 108.06i ± 10.74 168.08c ± 11 185a ± 10.17 103.87j ± 10.71 168.006c ± 11.6 171.66b ± 10.7 113.95h ± 12.67 72.58l ± 9.25 139.37e ± 8.78 141.3e ± 8.63 146.003d ± 11.03 – 132.57f ± 4.53 134.32f ± 4.49
Recipient Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja KVD2 KVD2 KVD2 KVD2 KVD2 KVD2 KVD2 KVD4 KVD4 KVD4 KVD4 KVD4 KVD4 KVD4
Donor Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
TA (%) 0.42ef ± 0.1 0.36e ± 0.11 – 0.46ef ± 0.12 0.32e ± 0.1 0.44ef ± 0.1 0.45ef ± 0.13 0.43ef ± 0.18 0.67cd ± 0.12 0.67cd ± 0.13 0.53c ± 0.13 0.61cd ± 0.16 0.67cd ± 0.11 0.55c ± 0.19 0.95ab ± 1.02 0.98ab ± 0.26 1.14a ± 0.36 1.12a ± 0.38 – 1.11a ± 0.2 0.99ab ± 0.2
Pectin (g/100gr) 0.15c ± 0.07 0.18c ± 0.01 – 0.18c ± 0.01 0.18c ± 0.01 0.18c ± 0.008 0.18c ± 0.02 0.23ab ± 0.04 0.28a ± 0.01 0.28a ± 0.01 0.24ab ± 0.02 0.28a ± 0.01 0.29a ± 0.005 0.23ab ± 0.05 0.26ab ± 0.05 0.28a ± 0.01 0.28a ± 0.01 0.29a ± 0.01 – 0.25ab ± 0.03 0.29a ± 0.008
Total phenol (mg/100grFW) 4.21e ± 0.19 2.19ef ± 0.52 – 5.63e ± 0.86 3.17ef ± 0.63 5.6e ± 0.62 4.51e ± 0.51 10.71c ± 0.72 11.94c ± 0.59 10.43c ± 0.92 9.88cd ± 0.95 8.91cd ± 0.58 10.22c ± 0.31 9.19cd ± 0.62 45.24a ± 0.25 41.45ab ± 0.99 40.34ab ± 0.71 46.26a ± 0.34 – 38.72ab ± 0.67 38.44ab ± 0.21
Similar letters in each column indicate no significant difference at the 5% level of LSD test.
Fig. 1. Effect of recipient and donor parents on TSS with standard errors.
Fig. 2. Effect of recipient and donor parents on fruit firmness with standard errors.
leading to an increase in the TSS or TA percentage. Different pollinizers had not any effects on the firmness of ‘Vidoja’ fruits, but in the other two female genotypes self-pollination treatment decreased fruit firmness (Fig. 2). The seed is a source of plant growth regulators that they play an important role in cell division, so it can be offered that in fruits with lower seeds, less cell division occurs and fruit
tissue has less firmness. There is metaxenia in some species of fruit trees. This means that the pollen grain effect on the physical and chemical characteristics of the fruit. In this case, the pollen can be effective in the formation and growth of the embryo and can cause small, large, completeness or incompleteness embryo and thereby it has influence on the growth and 131
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References Akbari Bisheh, H., Abdollahi, H., Torkashvand, M., Ghasemi, A., 2016. Pollen compatibility and determination of appropriate pollinizer for Isfahan quince cultivar (Cydonia oblonga Mill.). Seed Plant Improvement J. 32, 13–26 (In Persian). Akbari, H., 2014. Study of incompatibility and determination of the best pollinizer for Isfahan quince cultivar. J. Hortic. Sci. 28, 192–198 (In Persian). Alizadeh, S., Arzani, K., Imani, A., 2009. Determination of effective pollination period in late flowering almond (Prunus dulcis Mill.) Shahroud 12 in Karaj climate conditions. Iranian J. Hortic. Sci. 40, 1–7 (In Persian). Benedek, P., Szabo, T., Nyeki, J., 2001. New results on the bee pollination of quince (Cydonia oblongaMill.). Acta Hortic. 561, 243–248. Beyhan, N., Karakas, B., 2009. Investigation of the fertilization biology of some sweet cherry cultivars grown in the Central, Northern Anatolian Region of Turkey. Scientia Hortic 121, 320–326. Boskovic, R., Tobutt, K.R., 2001. Genotyping cherry cultivars assigned to incompatibility groups by analyzing stylar ribonucleases. Theor. Appl. Genet. 103, 475–485. Dalkilic, Z., Osman Mestav, H., 2011. Invitro pollen quantity, viability and germination tests in quince. Afr. J. Biotechnol. 10, 16516–16520. Denisow, B., 2003. Self pollination and self fertility in eight cultivars of black currant (Ribes nigrum L.). Acta Biologica Cracoviensia Series Botanica 45, 111–114. Deri, H.N., Farkas, A.H., 2013. Floral attractivity of Cydonia oblonga Mill cultivars. Acta Botany Hung. 48 (3–4), 279–290. Ershov, L.A., 1989. Biology of Quince pollination. Reports of the Soviet Scientists to the 17th International Congress on Horticulture. pp. 106–111. FAO, 2014. Food and Agriculture Organization of the United Nations. http://www.fao. org./. Gharaghani, A., Talaei, A.R., Zamani, Z., Fattahi Moghadam, M.R., Hajnajari, H., Gardiner, S., 2009. Necessity of covering in controlled pollination of apple by detection of microsatellite alleles in offspring of ‘Golden Smoothee’×'Shafi Abadi'. J. Sci. Technol. of Agric. and Nat. Resour. 48, 1–15. Keulemans, J., Brusselle, A., Eyssen, R., Vercammen, J., van Daele, G., 1996. Fruit weight in apple as influenced by seed number and pollinizer. Acta Hortic. 423, 201–210. Maniei, A., 1995. Planting to harvest of pear and quince. Technical Publication Iran 113p (In Persian). Matsumoto, S., Soejima, J., Mejima, T., 2012. Influence of repeated pollination on seed number and fruit shape of ‘Fuji’ apples. Sci. Hort. 137, 131–137. Mousavi, A., Babadaei, R., Fatahi, R., Zamani, Z., Dicenta, F., Ortega, E., 2014. Self-incompatibility in the iranian almond cultivar ‘Mamaei’ using pollen tube growth, fruit set and PCR technique. J. Nuts 5 (2), 1–10. Nagy-Deri, H., Orosz-Kovacs, Z., Farkas, A., 2013. Comparative studies on nectar from two self-fertile and two self-sterile cultivars of quince (Cydonia oblonga Mill.) and their attractiveness to honey bees. J. Hortic. Sci. Biotechnol. 88, 776–782. Nuzzo, V., Rubbi, G., 2004. Description and Use of Quince for Fruit Production. Department of Pomology University of Potenza, Italy 15pp. Ortega, E., Dicenta, F., 2004. Suitability of four different methods to identify self-compatible seedling in an almond breeding program. J. Hortic. Sci. Biotechnol. 79, 747–753. Rasouli, M., Arzani, K., 2011. Effect of pollen grain on the pollen tube growth and quantitative and qualitative characteristics of cherry fruits (Prunus avium L.) Zard Daneshkadeh cultivar. Iranian J. Hortic. Sci. 41, 309–318 (In Persian). Rasouli, M., Fattahi Moghadam, M.R., Zamani, Z., Imani, A., Ebadi, A., 2009. Study of compatibility and additional pollination of Supernova cultivar with pollen of different almond cultivars. Iranian J. Hortic. Sci. 40, 61–70 (In Persian). Rasouli, M., Arzani, K., Imani, A., Fattahi Moghadam, M.R., 2010. Study of pollination compatibility in some sweet cherry cultivars with Zard Daneshkadeh cultivar. Iranian J. Hortic. Sci. 41, 143–152 (In Persian). Sabeti, H., 1995. Iran's Forests, Trees and Shrubs. Publication of Yazd University 81p (In Persian). Sadat Mousavi, S., 2009. Identification of Self-incompatibility, Cross Incompatibility and Chilling Requirement of Some Asian Pear (Pyrus Serotina Rehd.) Genotypes. M.Sc. Thesis. Tarbiat Modares University, Tehran, Iran 115pp (In Persian). Singleton, V.L., Rossi, J.A., 1965. Colorimetry of total phenolics with phospho- molybdicphospho- tungstic acid reagents. Am. J. Enol. Viticult. 16, 144–158. Socias I Company, Alonso, J.M., Aparisi, J.G., 2004. Fruit set as an evaluation criterion in almond breeding. Acta Hortic. 663, 763–768. Stosser, R., Hartmann, W., Anvari, S.F., 1996. General aspects of pollination and fertilization of pome and stone fruit. Acta Hortic. 423, 15–22. Talaei, A., Vahedi, B., Askari, M.A., Ershadi, A., 2007. Selection of the best pollinizer cultivar for some Iranian commercial apple cultivars. Iranian J. Hortic. Sci. 38, 241–250 (In Persian). Thakur, B.R., Singh, R.K., Nelson, P.E., 1996. Quality attributes of processed tomato products. Food Rev. Int. 3, 357–401. Zhang, Y., Xue, Y., 2008. Molecular biology of S-RNase-based self-incompatibility. In: Franklin-Tong, V.E. (Ed.), Self-Incompatibility in Flowering Plants: Evolution, Diversity, and Mechanisms. Springer-Verlag, Berlin, Heidelberg, Germany, pp. 193–216.
Fig. 3. Effect of year and recipient parent on TA with standard errors.
Fig. 4. Effect of year on TSS with standard errors.
development of fruit. In this study, physical and chemical properties of the fruit are almost not affected by cross- pollination.
5. Conclusion The best source of pollen for the ‘Vidoja’ is KVD2 genotype. ‘Vidoja’ had the highest percentage of fruit set with this pollen donor parent. KVD4 and ‘Vidoja’ as pollen sources were favorable for KVD2 genotype and pollen of ‘Torsh’ and ‘Vidoja’ was able to had the highest percentage of fruit set for KVD4 genotype. These pollinizers in addition to the high percentage of fruit set had favorable effects on the female parent from aspects of the fruit size, weight and TSS. Flowering time will be different by climate, but the flowering transposition of genotypes to each other less affected by climate changes. From the results, it can be concluded that in order to have the suitable yield of quince is not logical to culture single cultivar. According to the results Vidoja', KVD2 and KVD4 have overlap of flowering (Table 2) and have other desirable traits, including flowering habit on spurs, non- alternate bearing, and desirable quantitative and qualitative characteristics of fruits (Table 1). Therefor It is recommended that these cultivars be used together in the establishment of new orchards so that effectively prevent dropping of flowers and fruitlets and subsequently increase yield in quince orchards.
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Effect of pollination on dropping of flowers and fruits in new quince (Cydonia oblonga Mill.) cultivar and promising genotypes
T
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Maryam Tataria, , Hamid Abdollahib, Asghar Mousavic a Horticulture Crops Research Department, Isfahan Agricultural and Natural Resources Research and Education Center, Agricultural Research, Education and Extension organization (AREEO), Isfahan, Iran b Temperate Fruits Research Center, Horticultural Sciences Research Institute, Agricultural Research, Education and Extension Organization (AREEO), Karaj, Iran c Horticulture Crops Research Department, Chahar-Mahal va Bakhtiari Agricultural and Natural Resources Research and Education Center, Agricultural Research, Education and Extension organization (AREEO), Shahrekord, Iran
A R T I C L E I N F O
A B S T R A C T
Keywords: Quince (Cydonia oblonga) Fruit set Pollen source Pollination Self-incompatibility
Access to sufficient and good quality fruits and solving problems such as early abscission of the fruitlets depends on the status of self-fertility and self-incompatibility in cultivars and genotypes and determination of the best pollinizer for them. This research was carried out to study the best pollinizer for some quince cultivars and promising genotypes from the collection of germplasm at the Horticultural Research Station of Isfahan, Iran in 2015 and 2016. For this, the effects of the pollen donor parent, including ‘Vidoja’, KVD2, KVD4, ‘Isfahan’, ‘Torsh’ and also open pollination were studied on quantitative and qualitative characteristics of the pollen recipient parent, including ‘Vidoja’, KVD2 and KVD4 as a factorial experiment in a randomized complete block design with four replications. To determine the best pollinizer, flowers of pollen recipient parents were emasculated at the balloon stage and were covered with isolation bags. Emasculated flowers were crossed with the pollen of donor parents during receptability of stigmas as well as hand self-pollination for each female parent. Results showed that some traits such as percentage of fruit set, seed numbers and total soluble solids (TSS) were affected by the pollen source. Phenol and pectin were not affected by the foreign pollen sources. According to the results, self-pollination did not have acceptable fruit set. The best pollen source for the ‘Vidoja’ was KVD2, for KVD4 was ‘Torsh’ and ‘Vidoja’ and for KVD2 was KVD4 and ‘Vidoja’, therefore the cultivation of all three female parents together in the quince orchards is recommended.
1. Introduction Quince (Cydonia oblonga L.) is economically the third fruit of pome fruit group. This species is native of Iran and its distribution centers are the northern forests of this country (Sabeti, 1995). This fruit was cultivated for 4000 years B. C. in Western Asia and it was favorite for many people in Greece and Rome and was called Mala cydonia. The scientific name of this tree has been described to the Sidon city, that had a lot of quince trees (Sabeti, 1995; Maniei, 1995). Quince is cultivated as extensive orchards in different provinces of Iran and area harvested has increased compared to previous years (FAO, 2014). One of the main problems for quince growers is abscission of flowers and fruits in the early stages of growth that reduce the yield and economic efficiency. Several factors, including the quantity and quality of water, soil quality, pruning, nutrition and environmental conditions before and after flowering can affect on abscission of flowers and fruits and ultimately reduction of yield. In the same condition of these factors,
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Corresponding author. E-mail address: [email protected] (M. Tatari).
https://doi.org/10.1016/j.scienta.2017.10.045 Received 18 June 2017; Received in revised form 9 October 2017; Accepted 31 October 2017 0304-4238/ © 2017 Elsevier B.V. All rights reserved.
the most important factor in fruit bearing is compatibility and fertilization (Ortega and Dicenta, 2004). Self- incompatibility or incomplete pollination of flowers is as one of the main causes of the fruit dropping (Boskovic and Tobutt, 2001). Compatible pollen grain, sufficient pollen transfer at the right time on the stigma and pollen tube growth in the style and finally fertilization are essential for successful production of fruit. The absence of pollination vectors or inappropriate weather conditions also cause problems in pollen transfer. In self-incompatible cultivars, pollen tube was not able to grow in the style and reach to ovule, so suitable pollinizer cultivars are essential to achieve economic yield (Socias I Company et al., 2004). Pollinizer cultivars should be compatible with main cultivar and have overlapping in flowering time. Therefore, weak pollination is as a limiting factor in crop production in the various regions (Alizadeh et al., 2009). Quince is introduced as a self compatible tree in the most references (Maniei, 1995), but based on the presented evidence about self-
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‘Torsh’ and ‘Isfahan’ cultivars along with three female parent (‘Vidoja’, KVD2 and KVD4) were considered as the pollen donor parent for pollination studies. Characteristics of the recipient and donor parent is as follows. Vidoja is early-flowering with a high density of the flowers, spore type, dwarf and without alternate bearing. Its fruit size is medium and uniformity, juicy, fragrant with low astringency and desirable quantitative and qualitative characteristics. KVD2 is early-flowering and high flowers density, spore type, dwarf, high yielding, early fruiting and without alternate bearing, fruits with medium to large size, uniform, juicy, fragrant with good quality properties. Characteristics of KVD4 are included early flowering and fruiting and without alternate bearing, spore type, dwarf, high yielding, large fruit and almost juicy with great aroma and sour taste. Isfahan cultivar is Late flowering and fruiting, without alternate bearing, invigorating, intermediate yields, large fruit, juicy, fragrant with low astringency and good quality. Torsh cultivar is early-flowering and fruiting, intermediate vigor, intermediate yields. Its fruit size is medium to large and almost juicy. The opening of 10%, 80% and 95% of flowers were considered as the beginning of flowering, full bloom and the end of flowering stages, respectively (Gharaghani et al., 2009).
incompatibility alleles on quince, there is a question that the use of cross pollination, what can be effective in increasing fruit in different cultivars of this species (Akbari Bisheh et al., 2016). More species in Rosaceae family have self-incompatibility, that's controlled by a locus with multiple S alleles (Zhang and Xue, 2008). In self-pollination aspect, quince cultivars are classificated into four groups, including the self-compatible cultivars with more than 10% of fruit set, semi selfcompatible cultivars with fruit set 3–8%, self incompatible cultivars with 1–2% of fruit set and completely incompatible cultivars with fruit set less that 1% (Nagy-Deri et al., 2013). Based on this classification, if semi self-incompatible cultivars are cultured next to pollinizer cultivars, they will produce more yield. The semi-self incompatible and completely self-incompatible cultivars need to compatible pollinizer cultivars for the appropriate fruit set. Therefore, the self-incompatibility percent of cultivars should be investigated in order to have sufficient yield. There are several methods to study the compatibility or incompatibility of different cultivars and determination of suitable pollinizer for them. These methods include controlled pollination, observation of pollen tube growth with a fluorescence microscope, extraction of ribonuclease from style, specific PCR of S allele and nucleotides sequencing related to incompatibility (Ortega and Dicenta, 2004; Mousavi et al., 2014). As regards, the controlled pollination method allows to estimate the performance of several cultivars in the orchard, so this method is recommended to determine the appropriate pollinizer (Rasouli et al., 2009). Nuzzo and Rubbi (2004) studied 22 quince cultivars in Italy and reported that most of them were self incompatible that required cross pollination to yield production. A number of cultivars have also been compatible that use of pollinizer was preferably necessary to obtain better yield. In order to study of the self-incompatibility and determination of the best pollinizer for ‘Isfahan’ commercial cultivar, female parent was pollinated with four genotypes that named KM1, PK3, KVD2 and NB4. KM1 genotype was introduced as the best pollinizer for ‘Isfahan’ with 80% of overlapping in flowering time and 23.42% of fruit set (Akbari, 2014). According to Talaei et al. (2007) fruit set percentage after the self-pollination was less than cross pollination. In the study of pollination status in 23 quince cultivars in various regions of Yugoslavia, it was found that only five cultivars of them are self-pollinated (Ershov, 1989). Pollination and fertility are one of the important issues in the quince trees. Some quince cultivars and genotypes, like other relatives in the Rosaceae family, including apples and pears, have self-incompatibility S-alleles. Self- incompatibility or incomplete pollination of flowers leads to fruitlet dropping and reduce yield, Therefore, it's necessary to be examined self or cross compatibility percent of any new cultivar with the introduction of it. Fourteen genotypes were identified and collected during the collection program of Iranian quince cultivars and genotypes in Isfahan, Iran. KVD1, KVD2 and KVD4 genotypes have been proposed as promising genotypes after preliminary evaluation of them. KVD1 genotype was released in the name of ‘Vidoja’. The aims of this study were the determination of the pollination status and suitable pollinizer for these genotypes and cultivar with the same time flowering period to achieve high performance.
2.2. Collection and viability test of pollen grains In order to collection of pollen grains, some branches of donor parents contain a sufficient number of flowers that are at the balloon stage were cut and transported to the laboratory. Brunches were placed in the sugar solution (4%). After opening flowers, the anthers were separated from flowers and were placed on glossy paper at room temperature (26–24°C) for 24–48 hours to be losing their moisture and drying of anthers. Pollen grains were separated from anthers and were collected in a sterile glass container with cotton cap. In order to study of viability, pollen grains were distributed in a petri dish containing medium with 1% agar, 15% sucrose and 50 mg/L of boric acid (Dalkilic and Osman, 2011) and germination percentage of pollen grains were recorded using a light microscope after 24 h exposure to laboratory temperature. Pollens with more than 75% of germination percentage were selected and were used as a pollen source. Sterile glasses containing pollen grains were stored in cold and dry condition at 4°C until pollination time.
2.3. Pollination With swelling of buds, suitable branches containing flower buds and uniform were randomly selected in the four geographical directions of each tree. Flowers were emasculated at the balloon stage and weak, immature and opened flowers as well as flower buds were removed from selected branches and were covered with cloth bags. After receptibility of stigma to accept pollen grain, each isolated branch was brought out of the bag and emasculated flowers were pollinated with considered pollen using a glass rod for each crosses. Pollinated flowers were counted and labeled and again were covered with cloth bags. To ensure of pollination, flowers were again pollinated with considered pollen 24 h later. Open pollination was evaluated as control treatment, while natural (only covered flower) and hand self-pollination was used for the study of self-pollination treatment. Self-pollination was carried out after emasculation by manually as hand self-pollination treatment. For each crosses, 100 flowers were emasculated and pollinated. These flowers were counted and labeled. For self-pollination treatment also 100 flowers were counted at the balloon stage and were covered with cloth bags without emasculation or any crosses. Cloth bags were removed after petal fall to allow the flowers to place under normal conditions.
2. Material and methods 2.1. Plant materials and flowering period This research was carried out at the Horticultural Research Station of Isfahan, Iran in 2015 and 2016. For this purpose, two promising quince genotypes called KVD2 and KVD4 along with ‘Vidoja’ cultivar that recently has been introduced were considered as the female parent.
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genotypes was recorded to determine overlapping in flowering time of them in 2015 and 2016 (Table 1). According to the results in Table 1, ‘Torsh’ cultivar in both years has been the very early flowering. Then ‘Vidoja’, KVD2 and KVD4 were early-flowering. ‘Isfahan’ was the late flowering that had not overlapping in flowering time with ‘Torsh’. This cultivar had overlapping in flowering time from three to four days with the ‘Vidoja’, KVD2 and KVD4. Results showed that flowering in the second year (2016) was sooner than the first year (2015). As regards quince is one of the latest flowering species, spring frost occurs rarely in Isfahan, Iran.
2.4. Measurement of traits In this study, some traits were measured, including the percentage of fruit set at harvesting time, the number of seeds per fruit, fruit dimensions, fruit weight, total soluble solids (TSS), firmness, titrable acidity (TA), total phenol and pectin content of fruit. The fruit set percentage was obtained by comparing the number of fertilized flowers with the number of final fruits. Fruit length, weight and the most fruit diameter were measured using ruler, scales and caliper respectively. Fruit firmness was measured by a penetrometer (model EFFEGI, Italy, plunger diameter 11.1 mm, depth 7.9 mm), at opposite peeled sides and expressed as kg/cm2. Total soluble solids (TSS) were determined by extracting and mixing two drops of juice from the two cut ends of each fruit into a digital refractometer (ATAGO N-1α, Japan) at 22°C. Titrable acids were determined in 10 g of pulp samples by titration of extracted juice with sodium hydroxide (0.1 mol/L) up to pH=8.1 and expressed as a percent of malic acid. Pectin content was measured according to Thakur et al. (1996). Briefly, 100 g of fruit tissue was grated and 400 mL of distilled water was added and was boiled for an hour. After passing through the filter paper, 300 mL of distilled water and 10 mL NaOH was added and the resulting solution was kept overnight at room temperature, then 50 mL acetic acid (1 mol/L) and 25 mL of calcium chloride was added to the solution. The resulting solution was kept for one hour at room temperature and then was boiled for an hour. Boiled solution was passed through filter paper. Difference between initial and secondary weight of filter paper was reported as pectin weight based on grams per 100 g of fruit pulp. Total phenols were measured in fruit Juice using Folin-Ciocalteu (Singleton and Rossi, 1965). Absorbance of the samples was determined at 765 nm wavelength with spectrophotometer model T80 UV/Visible, then compared with the standard of gallic acid and expressed as mg gallic acid per 100 g of fresh weight.
3.2. Pollen grain germination The results of pollen grain culture and viewing them under a light microscope showed that the pollen grain germination of ‘Torsh’, KVD4, KVD2, ‘Vidoja’ and ‘Isfahan’ was 84, 84, 85, 88 and 80 percent, respectively. 3.3. Fruit set percentage and seed number Analysis of variance for effect of recipient and donor parents on measured traits showed in Table 2. The highest percentage of fruit set for ‘Vidoja’ was obtained with pollen grains of KVD2 genotype in the first year with an average of 18.75%. In KVD4 genotype, pollen of ‘Torsh’ and ‘Vidoja’ produced the highest percentage of fruit set with an average of 28.64 and 18.54%, respectively. KVD2 with pollen of KVD4 showed fruit set equal to 30.28%. This genotype in cross pollination with pollen of ‘Vidoja’ had 19.86% of fruit set. Pollen sources of ‘Isfahan’ and ‘Torsh’ had little fruit set, especially for ‘Vidoja’ and KVD2 female parents. Generally, the fruit set percentage in the second year was less than the first year (Table 3). The lowest percentage of fruit set in all female parents was belonged to self-pollination treatments. Natural self-pollination in KVD4 and ‘Vidoja’ did not produce any fruit in both years. Fruit set in self-pollination of KVD2 was 3.47%. and hand self-pollination had also a very low fruit set percentage. According to the results (Table 4) the highest seed number was observed in KVD2 that pollinated with ‘Vidoja’ and KVD4. The least seed number was belonged to self-pollination treatments and especially natural self-pollination.
2.5. Statistical analysis Obtained results were compared using factorial experiment based on a randomized complete block design with four replications. As regards, the experiment was carried out in two years, compound analysis was also carried out to analyze the effect of the year. Analysis of data was performed by ANOVA method using statistical software SAS (version 9.1) and means comparison using the LSD test. Data normalization was carried out for all traits with ASIN formula.
3.4. Fruit dimensions and weight In cross-pollination of ‘Vidoja’ and KVD4 with other pollen sources as well as open pollination, fruit length and diameter was more than hand self-pollination. Self-pollination in these two cultivar and genotype did not produce any fruits. Fruit length and diameter in KVD2 genotype when crossed with ‘Isfahan’, ‘Vidoja’ and KVD4 as well as open pollination was more than in cross with ‘Torsh’ cultivar and selfpollination or hand self- pollination (Table 4).
3. Results 3.1. Flowering period The beginning of flowering and flowering period of cultivars and
Table 1 Flowering time of quince cultivars and promising genotypes in 2015 and 2016.
March- April 27
28
29
30
31
1
2
3
4
5
6
7
Vidoja (2015) Vidoja (2016) KVD2 (2015) KVD2 (2016) KVD4 (2015) KVD4 (2016) Isfahan (2015) Isfahan (2016) Torsh (2015) Torsh (2016)
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8
9
10
11
12
13
14
15
16
17
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Table 2 ANOVA for effect of donor and recipient parents on measured characteristics Source of variation
Year Rep (year) Recipient parent Donor parent Donor×Recipient Recipient×Year Donor×Year Donor×Recipient×Year Error Total C.V.% SOV
df
1 4 2 6 12 2 6 12 80 125 – df
Mean square Fruit set (%)
Seed number
Fruit length (cm)
Fruit diameter (cm)
Fruit weight (g)
19.03** 0.16 1.92** 14.76** 4.58** 0.37ns 1.43** 0.71** 0.26 – 15.38
9.17ns 154.56 70.16ns 361.98** 429.5** 18.62ns 15.5ns 10.41ns 62.36 – 18.09
0.006ns 0.25 2.06** 4.48** 0.55* 0.05ns 0.04ns 0.02ns 0.24 – 15.22
0.002ns 0.19 1.12** 4.32** 0.53* 0.04ns 0.03ns 0.02ns 0.22 – 17.74
0.15ns 12.33 49.8** 163.15** 21.54* 1.34ns 1.13ns 0.84ns 9.15 – 19.24
Mean square TSS (%)
Year Rep (year) Recipient parent Donor parent Donor×Recipient Recipient×Year Donor×Year Donor×Recipient × Year Error Total C.V.% **,* and
1 4 2 6 12 2 6 12 80 125 -
10.82** 1.27 2.15ns 14.58** 2.42** 0.11ns 0.21ns 0.18ns 0.81 – 16.86
Firmness (kg/cm2)
TA (%)
ns
ns
1.82 0.02 0.33** 0.2** 0.06** 0.47** 0.01ns 0.02ns 0.01 – 11.77
0.003 0.14 0.21ns 2.36** 0.32** 0.05ns 0.01ns 0.04ns 0.12 – 16.31
Pectin (g/100g) ns
0.0004 0.003 0.03** 0.02** 0.008** 0.002ns 0.001ns 0.001ns 0.002 – 5.44
Total phenol (mg/100gFW) 0.01ns 9.42 36.67** 79.63** 23.72** 2.55ns 1.65ns 1.53ns 4.37 – 17.69
ns
: Significant at the 1% and 5% probability level and non-significant difference respectively
3.6. Fruit firmness, total phenol and pectin content
Hand self-pollination in ‘Vidoja’ and KVD4 produced the lowest fruit weight while other pollen sources produced higher levels of fruit weight. Also, self pollination did not produce any fruit. According to the results, Fruit weight of KVD2 by hand self-pollination, self-pollination and cross pollination with ‘Torsh’ cultivar was less than that of KVD2 pollinated with other pollen sources (Table 4).
According to the standard errors related to fruit firmness in the ‘Vidoja’, firmness was not influenced by the type of pollen. In KVD2 and KVD4 genotypes, self pollination had less fruit firmness for these genotypes (Fig. 2). Different sources of pollen had no significant effect on the pectin and total phenol content in fruits of all three female parents. Lack of fruit set in some treatments caused significant difference between the crosses in these two traits. Regardless of the pollen source, KVD2 and KVD4 genotypes had more pectin in their fruits. Most of phenol was observed in KVD4 genotype (Table 4).
3.5. TA and TSS According to the results in Table 4, the highest TA was found in KVD4 that it is characteristic of this genotype. Type of pollinizer had no significant effect on TA in this genotype and no fruit production in natural pollination led to significant differences for this trait. Similarly, for ‘Vidoja’ and KVD2 also the TA was not affected by pollinizer cultivar. Interaction effect of the year and female parent on the TA percentage (Fig. 3) showed that ‘Vidoja’ and KVD4 in the first and second year had not significant difference, while in KVD2 genotype, TA in the second year was more than the first year. Pollen source had no significant effect on the TSS of ‘Vidoja’ fruits and TSS in hand self-pollination fruits were almost similar to TSS in other treatments. In this cultivar, natural self-pollination did not produce any fruit. In KVD2 genotype, natural and manual self-pollination as well as pollen of ‘Torsh’ cultivar produced less TSS than other pollinizers. The use of ‘Vidoja’ and ‘Isfahan’ pollen was led to the production of fruits with higher TSS. In KVD4, hand self-pollination treatment produced fruits with less TSS (Fig. 1). Regardless of the cultivar, the effect of year on TSS was significant, so that TSS in the second year was more than the first year (Fig. 4).
4. Discussion 4.1. Fruit set percentage The fruit set percentage depends on pollinizer cultivar, year of experiment and used method of pollination (Denisow, 2003). The highest fruit set percentage was obtained in ‘Vidoja’, KVD2 and KVD4 with pollen of KVD2, ‘Torsh’ and KVD4, respectively. ‘Vidoja’ was also suitable pollinizer for KVD2 and KVD4 genotypes. The desirability of open pollination was less than other pollinizers (Table 3). The number of the transferred pollen grains to the stigma can have positive or negative effects on fruit set in open pollination compared with controlled crosses. Transferred pollen grains to the stigma is more than 50 in open pollination while, this would be a few hundred pollen grains at the controlled crosses (Stosser et al., 1996). Minimum of pollen grains requires for pollen germination and fruit set. When the concentration of pollen is high, the increase of inhibition in pollen tube growth can lead
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according to the density of the flowers on quince trees is less than other trees, so the conversion of bloom to fruit in the range of 20–25 percent is desirable. They studied fruit set and fruit dropping of quince cv. ‘Ekmek’ in Van, Turkey and reported that the final fruit set in this cultivar during two years was 12.3% and 8.6%, respectively. Akbari (2014) introduced KM1 genotype as the best pollinizer for ‘Isfahan’ cultvar with 23.42% of fruit set. The effect of the year on fruit set with changing in the temperature, humidity, insect activity and the fruit orchard management conditions can be changed the fruit set percentage. According to Ortega and Dicenta (2004) different factors affect the amount and percentage of fruit set in various years. Some of these factors include the amount and quality of water, soil quality, pruning, fertilization and environmental conditions before and after flowering. In this study, it appears that spring frost can involve in less fruit set in the second year.
Table 3 Mean comparison of year, donor and recipient parents on fruit set percentage ± SD Year
Recipient parent
Donor parent
Fruit set (%)
2015 2015 2015 2015 2015 2015 2015
Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja
Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
0.5uv ± 0.5 4.73o ± 1.41 0v ± 0 18.75d ± 2.64 10.1g ± 4.16 11.47f ± 3.51 6.51mn ± 2.79
2015 2015 2015 2015 2015 2015 2015
KVD2 KVD2 KVD2 KVD2 KVD2 KVD2 KVD2
Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
3.07pqr ± 2.99 3.83p ± 1.89 19.86c ± 3.96 3.47p ± 3.28 30.28a ± 2.63 2.32rs ± 1.09 2.5qrs ± 2.34
2015 2015 2015 2015 2015 2015 2015
KVD4 KVD4 KVD4 KVD4 KVD4 KVD4 KVD4
Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
0.58uv ± 0.52 9.66gh ± 3.5 18.54d ± 2.01 7.82k ± 2.49 0v ± 0 14.69e ± 3 28.64b ± 3.04
2016 2016 2016 2016 2016 2016 2016
Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja
Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
0.58uv ± 0.38 3.16pq ± 2.56 0v ± 0 8.73ij ± 3.37 5.69n ± 1.86 7.36kl ± 3.59 3.78p ± 1.41
2016 2016 2016 2016 2016 2016 2016
KVD2 KVD2 KVD2 KVD2 KVD2 KVD2 KVD2
Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
1.73st ± 1.49 3.48p ± 2.81 8.09jk ± 1.68 3.82p ± 3.15 6.71lm ± 1.12 1.16tu ± 0.45 0.86u ± 0.93
2016 2016 2016 2016 2016 2016 2016
KVD4 KVD4 KVD4 KVD4 KVD4 KVD4 KVD4
Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
0.64uv ± 0.55 3.29pq ± 2.56 9.04hi ± 077 6.73lm ± 1.62 0v ± 0 7.42kl ± 1.007 10.04g ± 2.26
4.2. Seed number and fruit dimension and weight The lowest of seed number was obtained from hand self-pollination treatment (Table 4). Fruits resulting from pollination with pollen sources that had less compatibility with female parent and fewer fruit set, had almost lower seed number. Similarly, Matsumoto et al. (2012) also reported that the number of seeds per fruit was reduced semicompatible crosses. All of the studied quince fruits that were resulting from controlled crosses had some seeds in their fruits at harvesting time. This indicates that there are not parthenocarpy in the quince and fertilization is necessary for the fruit set, growth and development. Not parthenocarpy in ‘Isfahan’ cultivar have also been reported by Akbari Bisheh et al. (2016). Fruit length, diameter and weight resulting from controlled crosses followed a similar pattern (Table 4). According to the results, pollination with other pollen sources produced larger and heavier fruits than natural or hand self-pollination. For KVD2 exceptionally, ‘Torsh’ cultivar as well as self-pollination treatment could not produce fruits with the appropriate dimensions and weight. Matsumoto et al. (2012) did not observe a positive correlation between fruit weight and number of seeds in the apple, but Keulemans et al. (1996) reported this relationship in apple cv. ‘Fuji’ that is consistent with our results. 4.3. Other traits
Similar letters in each column indicate no significant difference at the 5% level of LSD test
Although the pectin content, total phenolics and TA were varied among female parents, but these traits were not affected by cross pollination in all three female parents and the lack of fruit set in natural self-pollination treatment was leds to a significant effect of pollinizers on these traits that has been shown in Table 4. Pollen source had no any significant effect on the fruit TSS in ‘Vidoja’ cultivar. Self-pollination was produced less TSS in the fruits of KVD4 genotype. In KVD2 genotype, the highest TSS were obtained from pollination with pollen source of ‘Vidoja’ (19.5%), ‘Isfahan’ (18.75%), open pollination (16.91%) and KVD4 (16.73%), respectively (Fig. 1). Naturally,‘Vidoja’ and ‘Isfahan’ has higher TSS than other genotypes, so it can be concluded that self pollination in these cultivars maintains TSS at a high level, while in the other genotypes which have less TSS, cross pollination with other pollen sources leading to increase of TSS in comparison with self-pollination. KVD2 genotype produced fruits with higher TSS when pollinated with ‘Vidoja’. The effects of pollinizer on TSS have been reported in apple cv. ‘Golabe Kohanz’ and ‘Shafiabadi’ by Talaei et al. (2007) and in sweet cherry by Rasouli and Arzani (2011). According to the results (Fig. 3), more TSS was observed in the second year. More TA was also found in KVD2 genotype in this year (Fig. 4). Due to the annual decrease in rainfall and less irrigation water in the second year, it seems the fruit juice in this year was less than first year, so the concentration of cell sap increased in the second year and
to increase of competition among pollen tubes (Beyhan and Karakas, 2009), but in this study, it seems controlled cross had not a negative effect on pollen concentration and fruit set that were according to the results of Beyhan and Karakas (2009). Natural self-pollination did not lead to the final fruit set in ‘Vidoja’ and KVD4. Fruit set percentage in self-pollination of sweet cherry cv. ‘Zarde Daneshkadeh’ was also equal to zero, so this cultivar was introduced as self-incompatible (Rasouli et al., 2010). Similarly Sadat (2009) studied some European pear cultivars including ‘Spadona’, ‘Beurre Bosc’, ‘Louise Bonne’, ‘Coscia’, ‘Shah Miveh’, ‘Shahak’, ‘Sard Roud’ and a genotype of ‘khoj’ and reported that all of them were completely self-incompatible. Hand self-pollination led to low fruit set percentage. There was not difference between hand and natural selfpollination in fruit set of KVD2 genotype. Reduce of fruit set percentage in self- pollination treatment have also been reported by other researchers (Ershov, 1989; Deri, 2013; Talaei et al., 2007). The percentage of fruit set in cross pollination with ‘Isfahan’ cultivar was not remarkable compared to other pollen sources. The percentage of fruit set in ‘Vidoja’ and KVD2 was low with ‘Torsh’ pollen grains. Salleles can also be involved in self-incompatibility or semi self-compatibility of cultivars and genotypes. Benedek et al. (2001) stated that 130
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Table 4 Mean comparison of donor and recipient parents on measured characteristics ± SD Recipient
Donor
Seed Number
Fruit length (cm)
Fruit diameter (cm)
Fruit weight (g)
Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja KVD2 KVD2 KVD2 KVD2 KVD2 KVD2 KVD2 KVD4 KVD4 KVD4 KVD4 KVD4 KVD4 KVD4
Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
12.33de ± 3.7 11.33de ± 2.16 – 24.16abc ± 4.23 20.33abcd ± 3.56 15.33cde ± 2.32 23abc ± 4.87 11.83de ± 4.41 13de ± 2.6 25.33ab ± 3.11 10.16e ± 4.1 27.66a ± 5.96 23abc ± 5.58 12.83de ± 5.45 10.66e ± 4.66 17.66bcde ± 3.28 23.66abc ± 3.29 20abcd ± 2.86 – 23.5abc ± 4.26 24.16abc ± 3.16
3.61ij ± 1.86 4.65fgh ± 0.41 – 4.68fg ± 0.63 4.28gh ± 0.36 4.86fg ± 0.74 5.08f ± 0.84 4.22gh ± 0.77 7.18ab ± 0.7 7.71a ± 0.59 4.09hi ± 0.99 6.97bc ± 0.23 7.41ab ± 0.28 4.95f ± 0.85 3.06j ± 0.39 5.89e ± 0.35 5.89e ± 0.26 6.54cd ± 0.7 – 5.93e ± 0.39 6.1de ± 0.42
3.88f ± 0.96 4.94cde ± 0.44 – 5.07cd ± 0.51 4.52e ± 0.23 5.44c ± 0.63 5.11cd ± 0.48 3.46fg ± 0.72 6.7a ± 0.74 6.96a ± 0.48 3.85f ± 0.99 6.67a ± 0.24 6.61ab ± 0.53 4.61de ± 0.59 2.94g ± 0.31 5.43c ± 0.46 5.36c ± 0.31 6.12b ± 0.69 – 5.38c ± 0.27 5.41c ± 0.3
92.64k ± 11.92 118.07g ± 6.58 – 120.72g ± 5.97 114.09h ± 1.57 120.66g ± 3.82 121.33g ± 7.66 108.06i ± 10.74 168.08c ± 11 185a ± 10.17 103.87j ± 10.71 168.006c ± 11.6 171.66b ± 10.7 113.95h ± 12.67 72.58l ± 9.25 139.37e ± 8.78 141.3e ± 8.63 146.003d ± 11.03 – 132.57f ± 4.53 134.32f ± 4.49
Recipient Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja Vidoja KVD2 KVD2 KVD2 KVD2 KVD2 KVD2 KVD2 KVD4 KVD4 KVD4 KVD4 KVD4 KVD4 KVD4
Donor Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh Hand self p. Isfahan Vidoja KVD2 KVD4 Open p. Torsh
TA (%) 0.42ef ± 0.1 0.36e ± 0.11 – 0.46ef ± 0.12 0.32e ± 0.1 0.44ef ± 0.1 0.45ef ± 0.13 0.43ef ± 0.18 0.67cd ± 0.12 0.67cd ± 0.13 0.53c ± 0.13 0.61cd ± 0.16 0.67cd ± 0.11 0.55c ± 0.19 0.95ab ± 1.02 0.98ab ± 0.26 1.14a ± 0.36 1.12a ± 0.38 – 1.11a ± 0.2 0.99ab ± 0.2
Pectin (g/100gr) 0.15c ± 0.07 0.18c ± 0.01 – 0.18c ± 0.01 0.18c ± 0.01 0.18c ± 0.008 0.18c ± 0.02 0.23ab ± 0.04 0.28a ± 0.01 0.28a ± 0.01 0.24ab ± 0.02 0.28a ± 0.01 0.29a ± 0.005 0.23ab ± 0.05 0.26ab ± 0.05 0.28a ± 0.01 0.28a ± 0.01 0.29a ± 0.01 – 0.25ab ± 0.03 0.29a ± 0.008
Total phenol (mg/100grFW) 4.21e ± 0.19 2.19ef ± 0.52 – 5.63e ± 0.86 3.17ef ± 0.63 5.6e ± 0.62 4.51e ± 0.51 10.71c ± 0.72 11.94c ± 0.59 10.43c ± 0.92 9.88cd ± 0.95 8.91cd ± 0.58 10.22c ± 0.31 9.19cd ± 0.62 45.24a ± 0.25 41.45ab ± 0.99 40.34ab ± 0.71 46.26a ± 0.34 – 38.72ab ± 0.67 38.44ab ± 0.21
Similar letters in each column indicate no significant difference at the 5% level of LSD test.
Fig. 1. Effect of recipient and donor parents on TSS with standard errors.
Fig. 2. Effect of recipient and donor parents on fruit firmness with standard errors.
leading to an increase in the TSS or TA percentage. Different pollinizers had not any effects on the firmness of ‘Vidoja’ fruits, but in the other two female genotypes self-pollination treatment decreased fruit firmness (Fig. 2). The seed is a source of plant growth regulators that they play an important role in cell division, so it can be offered that in fruits with lower seeds, less cell division occurs and fruit
tissue has less firmness. There is metaxenia in some species of fruit trees. This means that the pollen grain effect on the physical and chemical characteristics of the fruit. In this case, the pollen can be effective in the formation and growth of the embryo and can cause small, large, completeness or incompleteness embryo and thereby it has influence on the growth and 131
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References Akbari Bisheh, H., Abdollahi, H., Torkashvand, M., Ghasemi, A., 2016. Pollen compatibility and determination of appropriate pollinizer for Isfahan quince cultivar (Cydonia oblonga Mill.). Seed Plant Improvement J. 32, 13–26 (In Persian). Akbari, H., 2014. Study of incompatibility and determination of the best pollinizer for Isfahan quince cultivar. J. Hortic. Sci. 28, 192–198 (In Persian). Alizadeh, S., Arzani, K., Imani, A., 2009. Determination of effective pollination period in late flowering almond (Prunus dulcis Mill.) Shahroud 12 in Karaj climate conditions. Iranian J. Hortic. Sci. 40, 1–7 (In Persian). Benedek, P., Szabo, T., Nyeki, J., 2001. New results on the bee pollination of quince (Cydonia oblongaMill.). Acta Hortic. 561, 243–248. Beyhan, N., Karakas, B., 2009. Investigation of the fertilization biology of some sweet cherry cultivars grown in the Central, Northern Anatolian Region of Turkey. Scientia Hortic 121, 320–326. Boskovic, R., Tobutt, K.R., 2001. Genotyping cherry cultivars assigned to incompatibility groups by analyzing stylar ribonucleases. Theor. Appl. Genet. 103, 475–485. Dalkilic, Z., Osman Mestav, H., 2011. Invitro pollen quantity, viability and germination tests in quince. Afr. J. Biotechnol. 10, 16516–16520. Denisow, B., 2003. Self pollination and self fertility in eight cultivars of black currant (Ribes nigrum L.). Acta Biologica Cracoviensia Series Botanica 45, 111–114. Deri, H.N., Farkas, A.H., 2013. Floral attractivity of Cydonia oblonga Mill cultivars. Acta Botany Hung. 48 (3–4), 279–290. Ershov, L.A., 1989. Biology of Quince pollination. Reports of the Soviet Scientists to the 17th International Congress on Horticulture. pp. 106–111. FAO, 2014. Food and Agriculture Organization of the United Nations. http://www.fao. org./. Gharaghani, A., Talaei, A.R., Zamani, Z., Fattahi Moghadam, M.R., Hajnajari, H., Gardiner, S., 2009. Necessity of covering in controlled pollination of apple by detection of microsatellite alleles in offspring of ‘Golden Smoothee’×'Shafi Abadi'. J. Sci. Technol. of Agric. and Nat. Resour. 48, 1–15. Keulemans, J., Brusselle, A., Eyssen, R., Vercammen, J., van Daele, G., 1996. Fruit weight in apple as influenced by seed number and pollinizer. Acta Hortic. 423, 201–210. Maniei, A., 1995. Planting to harvest of pear and quince. Technical Publication Iran 113p (In Persian). Matsumoto, S., Soejima, J., Mejima, T., 2012. Influence of repeated pollination on seed number and fruit shape of ‘Fuji’ apples. Sci. Hort. 137, 131–137. Mousavi, A., Babadaei, R., Fatahi, R., Zamani, Z., Dicenta, F., Ortega, E., 2014. Self-incompatibility in the iranian almond cultivar ‘Mamaei’ using pollen tube growth, fruit set and PCR technique. J. Nuts 5 (2), 1–10. Nagy-Deri, H., Orosz-Kovacs, Z., Farkas, A., 2013. Comparative studies on nectar from two self-fertile and two self-sterile cultivars of quince (Cydonia oblonga Mill.) and their attractiveness to honey bees. J. Hortic. Sci. Biotechnol. 88, 776–782. Nuzzo, V., Rubbi, G., 2004. Description and Use of Quince for Fruit Production. Department of Pomology University of Potenza, Italy 15pp. Ortega, E., Dicenta, F., 2004. Suitability of four different methods to identify self-compatible seedling in an almond breeding program. J. Hortic. Sci. Biotechnol. 79, 747–753. Rasouli, M., Arzani, K., 2011. Effect of pollen grain on the pollen tube growth and quantitative and qualitative characteristics of cherry fruits (Prunus avium L.) Zard Daneshkadeh cultivar. Iranian J. Hortic. Sci. 41, 309–318 (In Persian). Rasouli, M., Fattahi Moghadam, M.R., Zamani, Z., Imani, A., Ebadi, A., 2009. Study of compatibility and additional pollination of Supernova cultivar with pollen of different almond cultivars. Iranian J. Hortic. Sci. 40, 61–70 (In Persian). Rasouli, M., Arzani, K., Imani, A., Fattahi Moghadam, M.R., 2010. Study of pollination compatibility in some sweet cherry cultivars with Zard Daneshkadeh cultivar. Iranian J. Hortic. Sci. 41, 143–152 (In Persian). Sabeti, H., 1995. Iran's Forests, Trees and Shrubs. Publication of Yazd University 81p (In Persian). Sadat Mousavi, S., 2009. Identification of Self-incompatibility, Cross Incompatibility and Chilling Requirement of Some Asian Pear (Pyrus Serotina Rehd.) Genotypes. M.Sc. Thesis. Tarbiat Modares University, Tehran, Iran 115pp (In Persian). Singleton, V.L., Rossi, J.A., 1965. Colorimetry of total phenolics with phospho- molybdicphospho- tungstic acid reagents. Am. J. Enol. Viticult. 16, 144–158. Socias I Company, Alonso, J.M., Aparisi, J.G., 2004. Fruit set as an evaluation criterion in almond breeding. Acta Hortic. 663, 763–768. Stosser, R., Hartmann, W., Anvari, S.F., 1996. General aspects of pollination and fertilization of pome and stone fruit. Acta Hortic. 423, 15–22. 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Fig. 3. Effect of year and recipient parent on TA with standard errors.
Fig. 4. Effect of year on TSS with standard errors.
development of fruit. In this study, physical and chemical properties of the fruit are almost not affected by cross- pollination.
5. Conclusion The best source of pollen for the ‘Vidoja’ is KVD2 genotype. ‘Vidoja’ had the highest percentage of fruit set with this pollen donor parent. KVD4 and ‘Vidoja’ as pollen sources were favorable for KVD2 genotype and pollen of ‘Torsh’ and ‘Vidoja’ was able to had the highest percentage of fruit set for KVD4 genotype. These pollinizers in addition to the high percentage of fruit set had favorable effects on the female parent from aspects of the fruit size, weight and TSS. Flowering time will be different by climate, but the flowering transposition of genotypes to each other less affected by climate changes. From the results, it can be concluded that in order to have the suitable yield of quince is not logical to culture single cultivar. According to the results Vidoja', KVD2 and KVD4 have overlap of flowering (Table 2) and have other desirable traits, including flowering habit on spurs, non- alternate bearing, and desirable quantitative and qualitative characteristics of fruits (Table 1). Therefor It is recommended that these cultivars be used together in the establishment of new orchards so that effectively prevent dropping of flowers and fruitlets and subsequently increase yield in quince orchards.
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