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Hiroshi Iwanami, Makoto Ishiguro, Nobuhiro Kotoda, Sae Takahashi and Junichi Soejima

The firmness of the flesh in 27 apple (Malus ×domestica Borkh.) cultivars and selections (genotypes) was measured as an indicator of storage potential at 20 days after harvest under 20 ± 2 °C, 80% ± 5%relative humidity storage conditions. Softening ranged from 9% to 58% of initial values among genotypes after 20 days of storage. In some genotypes, softening was not continuous, a minimum firmness being reached before day 20. After a period of rapid softening, firmness declined to at least 20% of that at harvest. For each genotype, linear regression analysis of firmness changes from harvest until when firmness decreased by 20% was carried out. In genotypes in which firmness did not drop >20% within 20 days of storage, the entire dates to 20 days were used for analysis. The homogeneity of the regression residual variances and their normal distribution was not rejected at P = 0.05, and the linear regression analysis was assumed to be applicable to the change in firmness for each genotype. Results of the regression analysis showed that the regression was significant for all genotypes except one. Therefore, storage potential could be evaluated by comparing the regression coefficient of each genotype.

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Hiroshi Iwanami, Shigeki Moriya, Nobuhiro Kotoda, Sae Takahashi and Kazuyuki Abe

Progenies from 38 unbalanced crosses using 20 apple (Malus ×domestica Borkh.) cultivars/selections as parents were evaluated for changes in flesh firmness after harvest in two seasons to determine the mechanism of inheritance of fruit softening. The change in firmness was fitted by linear regression, and the softening rate (N·d−1) expressed as the regression coefficient was used as the phenotypic value of softening after harvest. Fruit were stored under 20 °C and 85% relative humidity after harvest for up to 40 days. The softening rates in the progeny populations were distributed continuously around the softening rates of parents, despite a distinct segregation in the degree of mealiness at 30 days of storage. The narrow-sense heritability of the softening rate was estimated by parent-offspring regression, and the estimate was high (h2 = 0.93). Because the softening rate can be influenced by mealiness, an undesirable trait in the apple industry, the progenies were divided into individuals with and without mealiness, and the breeding values of the parents were estimated based on the softening rate of the nonmealy progeny. The softening rate of the nonmealy progeny was analyzed using a mixed linear model and the restricted maximum likelihood method, with general combining ability (GCA) as parental effects and specific combining ability (SCA) as parental interaction effects. The variance of GCA was significant, but the variance of SCA was small and nonsignificant. The narrow-sense heritability of the softening rate in the nonmealy progeny was estimated by sib analysis, and the estimate was moderately high (h2 = 0.55). A significant correlation was observed between the phenotypic value and the breeding value (twice the GCA effects) in nonmealy parents, but the phenotypic value did not significantly correlate with the breeding value in mealy parents. Therefore, contribution of a mealy parent to the softening rate of nonmealy progenies cannot be predicted by its phenotypic value.

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Hiroshi Iwanami, Shigeki Moriya, Nobuhiro Kotoda, Sae Takahashi and Kazuyuki Abe

Changes in flesh firmness and mealiness during storage were investigated in 24 apple [Malus ×sylvestris (L.) Mill. var. domestica (Borkh.) Mansf.] cultivars and selections (genotypes) up to 40 days after harvest under 20 ± 2 °C and 85% ± 5% relative humidity storage conditions. Flesh firmness was measured using a penetrometer, while mealiness was quantified by measuring the degree of cell separation in tissue induced by shaking discs of tissue in a sucrose solution. According to the relationship between the change in firmness and mealiness, the genotypes can be divided into four groups: those that did not soften and remained hard and nonmealy during storage; those that softened without mealiness; those that softened with slight mealiness; and those that softened with mealiness. Firmness decreased below 30 N in fruit that softened with mealiness, and the minimum firmness during storage was correlated with the degree of mealiness at 30 days of storage. The rate of softening was the highest in fruit that softened with mealiness. Therefore, it was concluded that, by measuring the firmness and changes in firmness that take place during storage, the genotypes resulting in softening with mealiness and those that result in softening without mealiness could be identified.

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Hiroshi Iwanami, Shigeki Moriya, Nobuhiro Kotoda and Kazuyuki Abe

Changes in turgor and flesh firmness during storage at 20 °C were investigated using 27 apple (Malus ×domestica Borkh.) cultivars for 2 years. Flesh firmness was measured using a penetrometer, and turgor was determined using a thermocouple psychrometer. Firmness and turgor of fruit decreased during storage. The cultivars with little softening during storage had low rates of reduction in turgor. The softening rates in mealy cultivars were high, but there were cultivars with low rates of turgor reduction. When the rates of reduction in turgor after harvest were low, the mealy cultivars of the fruit tended to develop severe mealiness during storage. Therefore, a low rate of reduction in turgor could contribute to cultivars with both good shelf life and severe mealiness. The reduction rates of turgor in progeny cultivars were nearly identical to the mean reduction rates of turgor of their parents. This suggests that a cultivar with a low reduction rate of turgor, although it can be mealy, has the potential to produce a progeny with a low reduction rate of turgor.

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Hiroshi Iwanami, Shigeki Moriya, Nobuhiro Kotoda, Sae Takahashi and Kazuyuki Abe

To compare changes in fruit quality during cold storage with those during shelf life conditions, flesh firmness and titratable acidity (TA) were measured during storage in 20 apple (Malus ×domestica Borkh.) cultivars. Fruit of each cultivar were divided into two groups and stored in chambers controlled at 20 ± 2 °C and 85 ± 5% relative humidity (RH) (shelf life conditions) or 0.5 ± 0.3 °C and 95 ± 5% RH (cold storage). Five of the stored fruit were removed for measurements at 5- or 10-d intervals for 40 d and at 1-month intervals until 10 months after harvest at 20 °C and 0.5 °C, respectively. Data for firmness and TA were subjected to a linear regression and a nonlinear regression, respectively. Moreover, to determine the advantages of 0.5 °C storage over 20 °C storage for retaining firmness and TA, the effect of storage type on extending the storage period was introduced as a parameter. The estimate of the effect of storage type showed that firmness and TA could be retained 8.9 and 3.7 times, respectively, longer at 0.5 °C than 20 °C, independently of the cultivar. Therefore, firmness and TA after cold storage could be predicted by the change in firmness and TA during shelf life conditions. Moreover, cultivar differences regarding quality change under cold storage could be determined in a short period after harvest because the cultivar differences under shelf life conditions were detected within 1 month after harvest.

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Nobuhiro Kotoda, Hiroshi Iwanami, Sae Takahashi and Kazuyuki Abe

Because fruit trees such as apple (Malus ×domestica Borkh.) flower and set fruit only after an extended juvenile phase lasting several years, efficient breeding of fruit trees is limited. We previously suggested that MdTFL1 (Malus ×domestica TFL1) functions analogously to TERMINAL FLOWER 1 (TFL1) and that MdTFL1 is involved in the maintenance of the juvenile/vegetative phase in apple. To clarify the function of MdTFL1 in apple, we produced transgenic `Orin' apple trees expressing MdTFL1 antisense RNA. One of them flowered only 8 months after the transfer to the greenhouse, whereas the nontransformed control plants have not flowered in nearly 6 years. As expected, the expression of endogenous MdTFL1 was suppressed in the transgenic lines that showed precocious flowering. In addition, the expression level of the transgene was correlated with the reduction of the juvenile phase. These findings confirm that MdTFL1 functions like TFL1 and that MdTFL1 maintains the juvenile and vegetative phase in apple. Flower organs of the transgenic apple trees were normal in appearance, and a precocious flowering transgenic line set fruit and seeds. Interestingly, some flowers of the transgenic apple trees developed without undergoing dormancy. The expression of MdTFL1 in apple may affect flower development as well as flower induction.

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Nobuhiro Kotoda, Masato Wada, Sadao Komori, Shin-ichiro Kidou, Kazuyuki Abe, Tetsuo Masuda and Junichi Soejima

Two apple [Malus sylvestris (L.) Mill. var. domestica (Borkh.) Mansf.] homologous fragments of FLO/LFY and SQUA/AP1 (AFL and MdAP1, respectively) were analyzed to determine the relationship between floral bud formation and floral gene expression in `Jonathan' apple. The AFL gene was expressed in reproductive and vegetative organs. By contrast, the MdAP1 gene, identified as MdMADS5, which is classified into the AP1 group, was expressed specifically in sepals concurrent with sepal formation. Based on these results, AFL may be involved in floral induction to a greater degree than MdAP1 since AFL transcription increased ≈2 months earlier than MdAP1. Characterization of AFL and MdAP1 should advance the understanding of the processes of floral initiation and flower development in woody plants, especially in fruit trees like apple.

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Monrudee Kittikorn, Katsuya Okawa, Hitoshi Ohara, Satoru Kondo, Nobuhiro Kotoda, Masato Wada, Mineyuki Yokoyama, Ohji Ifuku, Ariake Murata and Naoharu Watanabe

Changes of endogenous 9, 10-ketol-octadecadienoic acid (KODA) concentrations, which is synthesized from linolenic acid by 9-lipoxygenase, were analyzed in apple [Malus ×sylvestris (L.) Mill. var. domestica (Borkh.) Mansf.] buds. In addition, the effects of 15, 16-chloro, hydroxy-9-hydroxy-10-oxo-12(Z)-octadecenoic acid (CKODA) application, which is an analog of KODA, on flower bud formation and the expression of MdTFL1 (terminal flower 1) and MdFT1 (flowering locus t 1) genes in apple buds were investigated in heavy-crop treatment (HCT) and under shade. An increase of endogenous KODA in the buds in the fruit-thinning treatment, which resulted in a higher proportion of flower bud formation than in HCT, was observed at 63 days after full bloom, but no such increase was found in HCT. In the shade-treated and heavy-crop trees, the expression of MdTFL1 in the buds to which CKODA was applied was lower than that in untreated buds. In contrast, under shade, the expression of MdFT1 in the CKODA-treated buds was higher than that in untreated buds. These results suggest that endogenous KODA may be associated with flower bud formation, and its application may be effective at improving the proportion of flower bud formation through its effect on MdTFL1 and MdFT1.