Effects of naphthaleneacetic acid (NAA), aminoethoxyvinylglycine (AVG), and sprayable 1-methylcyclopropene (1-MCP) alone or in combination on fruit ethylene production, preharvest fruit drop, fruit quality, and fruit maturation were examined in ‘Delicious’ apples (Malus ×domestica Borkh.). 1-MCP and AVG + NAA, when applied 15 days before anticipated harvest (DBAH) for untreated control trees, more effectively delayed preharvest fruit drop than AVG or NAA used alone. However, there was no significant difference in ethylene production between fruit treated with 1-MCP or AVG + NAA and those treated by AVG. Two applications of NAA increased fruit ethylene production and fruit softening, whereas AVG inhibited NAA-enhanced fruit ethylene production and fruit softening. There was no significant difference in fruit ethylene production, fruit firmness, and fruit drop control between one and two applications of 1-MCP. The concentrations of 1-MCP did not affect the efficacy of 1-MCP when applied 15 DBAH, but high concentration of 1-MCP more effectively delayed preharvest fruit drop than low concentration of 1-MCP when applied 7 DBAH. Both AVG and 1-MCP suppressed expression of 1-aminocyclopropane-1-carboxylate (ACC) synthase gene MdACS1, ACC oxidase gene MdACO1, and polygalacturonase gene MdPG1 in fruit. Expression of ACS5A and MdACO1 but not MdACS1 in fruit abscission zones was decreased by AVG and 1-MCP. 1-MCP more effectively suppressed expression of MdPG2 in fruit abscission zones than AVG alone.
Rongcai Yuan and Duane W. Greene
BA applied at the 10-mm stage at 50 and 100 ppm thinned, increased fruit size, and seed abortion. Net photosynthesis was decreased and dark respiration was increased when temperature following BA application was high (30°C), whereas there was no effect when temperature was lower (20°C). The seed number in abscising fruit was greater in BA-treated fruit than in control fruit. The number of viable seeds in BA-treated fruit was reduced. Tipping the bourse shoot increased fruit set, regardless of BA treatment. BA did not thin fruit with 25 leaves or greater. The translocation of 14C-sorbital from leaves to fruit was promoted by BA application to the fruit, but not when BA was applied to the leaves. The thinning induced by BA will be discussed in relation to available carbohydrate.
Rongcai Yuan and Duane W. Greene
BA was applied at 50 or 100 mg·L-1 to `More-Spur McIntosh'/Malling 7 (M.7) apple trees [Malus sylvestris (L.) Mill var. domestica (Borkh.) Mansf.] at the 10 mm stage of fruit development. BA thinned fruit and increased fruit size. There were two distinguishable peaks of fruit abscission during `June drop'. BA accentuated the naturally occurring waves of fruit abscission, and enhanced translocation of 14C-sorbitol from leaves to fruit when applied directly to the fruit, but not when applied directly to the leaves. Net photosynthesis was decreased and dark respiration was increased when temperature following BA application was high (30 °C), whereas there was no effect when temperature was lower (20 °C). Total nonstructural carbohydrates, total soluble sugars, and starch in the leaves decreased dramatically over the 12- or 13-day observation period, regardless of BA treatment. These carbohydrate concentrations in the leaves were lowered further by BA application. Abscising fruit, based on specific reddening of the pedicel, had higher carbohydrate levels than persisting fruit, regardless of BA application. We conclude that BA thins fruit, at least in part, by increasing dark respiration and decreasing net photosynthesis. Chemical name used: N-(phenylmethyl)-1H-purine-6-amine [benzyladenine (BA)].
Rongcai Yuan and Duane W. Greene
Experiments were conducted to evaluate the effects of BA, removal of bourse shoot tips including only folded leaves and growing point, and different numbers of leaves per fruit on fruit retention and fruit development in `More-Spur McIntosh'/Malling 7 (M.7) apple trees [Malus sylvestris (L.) Mill var. domestica (Borkh.) Mansf.]. Removal of the bourse shoot tip increased fruit retention, whereas BA thinned fruit regardless of whether shoot tips were removed or not. There was no interaction between BA application and shoot tipping. BA thinned fruit only when one leaf per fruit was on a girdled small fruiting branch, but not when leaf number per fruit was two or greater. Fruit weight and soluble solids concentration increased dramatically with increasing leaf number per fruit. BA reduced fruit growth rate when <16 leaves per fruit were present on the girdled branches between 3 and 7 days after treatment, but it did not affect fruit growth rate when 32 leaves per fruit were on the girdled branches. Increasing leaf number also increased viable seed number per fruit while decreasing the number of aborted seeds, but it had no effect on the number of total seeds per fruit. BA reduced the number of viable seeds per fruit only when the number of leaves per fruit was less than four. Results suggest that BA thins apple fruit mainly by reducing carbohydrates available to developing fruitlets. Chemical name used: N-(phenylmethyl)-1H-purine-6-amine [benzyladenine (BA)].
Rongcai Yuan and Jacqueline K. Burns
The effect of temperature on the ability of 5-chloro-3-methyl-4-nitro-1H-pyrazole (CMNP) and ethephon to induce ethylene evolution and abscission of mature fruit and leaves was determined using 3-year-old potted `Hamlin' orange [Citrus sinensis (L.) Osb.] trees in environment-controlled growth rooms in seasons 2001-02 and 2002-03. Ethylene evolution and abscission of CMNP or ethephon-treated fruit and ethephon-treated leaves were highly temperature dependent. Fruit detachment force (FDF) and fruit ethylene evolution were not affected by application of ethephon at 200 mg·L-1 or CMNP at 200 mg·L-1 when air temperature was 10 °C for ethephon treatment or ≤15.6 °C for CMNP treatment. However, ethylene evolution of CMNP or ethephon-treated fruit increased sharply, and FDF decreased drastically as temperature increased from 10 to 26.7 °C for ethephon treatment or from 15.6 to 26.7 °C for CMNP treatment. Several 10 hour day/14 hour night temperature regimes were explored to determine the effect of varying daily and nightly temperatures on efficacy and ethylene evolution. At least 3 days of exposure to 21/10 °C were required for CMNP to effectively loosen fruit, whereas only one day of exposure to 26.7/15.6 °C was enough to induce similar changes. At 21/10 °C, CMNP significantly reduced FDF to<25 N and markedly enhanced fruit ethylene evolution, regardless of interruption by 1 day of low temperature at 10/10 °C in the first 5 d after application. Ethephon had no significant effect on leaf ethylene evolution and leaf abscission when temperature was 10 °C, but caused a marked increase in both leaf ethylene evolution and leaf abscission as temperature increased from 10 to 26.7 °C. CMNP did not stimulate leaf ethylene evolution and leaf abscission regardless of temperature. Chemical names used: 5-chloro-3-methyl-4-nitro-1 H-Pyrazole (CMNP); 2-chloroethylphosphonic acid (ethephon).
Rongcai Yuan and David H. Carbaugh
Effects of naphthaleneacetic acid (NAA), aminoethoxyvinylglycine (AVG), and 1-methylcyclopropene (1-MCP) alone or in combination on fruit ethylene production, preharvest fruit drop, fruit quality, and fruit maturation were examined in ‘Golden Supreme’ and ‘Golden Delicious’ apples (Malus ×domestica Borkh.). In ‘Golden Supreme’ apples, the combination of two applications of AVG and one application of NAA 3 and 1 week, respectively, before the anticipated optimum harvest date synergistically inhibited fruit ethylene production and delayed fruit drop and ripening. Compared with one or two applications of AVG, the combination of one application of AVG and two applications of NAA had much lower preharvest fruit drop, although there was no significant difference in fruit ethylene production among these treatments. In ‘Golden Delicious’ apples, 1-MCP at 396 mg·L−1 had a better effect in delaying fruit drop than did AVG at 125 mg·L−1 or NAA at 20 mg·L−1 when they were applied a week before the optimum harvest date. The combination of NAA and 1-MCP or AVG was more effective in delaying fruit drop than were NAA, 1-MCP, or AVG alone. Fruit ethylene production was inhibited by 1-MCP and AVG but not by NAA. 1-MCP and AVG delayed fruit ripening, whereas NAA increased fruit ripening as determined by fruit firmness and starch.
Jianguo Li, Hong Zhu and Rongcai Yuan
The expression of genes for ethylene biosynthesis, ethylene perception, and cell wall degradation in the fruit cortex and fruit abscission zone (FAZ) was examined in relation to preharvest fruit abscission (PFA) and fruit ripening in ‘Golden Delicious’ and ‘Fuji’ apple (Malus ×domestica Borkh.). PFA, fruit ethylene production, and fruit softening increased rapidly during fruit ripening in ‘Golden Delicious’ apples, whereas no PFA, little fruit ethylene, and gradual fruit softening were recorded in ‘Fuji’ apples. The transcript levels of 1-aminocyclopropane-1-carboxylate (ACC) synthase genes, MdACS1, MdACS3, and MdACS5A, increased rapidly in the fruit cortex of ‘Golden Delicious’ apples during ripening, but not in ‘Fuji’ apples. However, only the level of MdACS5A mRNA was up-regulated in the FAZ of ‘Golden Delicious’ apples. The transcript level of ACC oxidase gene, MdACO1, increased in the fruit cortex for both cultivars but increased only in the FAZ of ‘Golden Delicious’ apples. Expression of the ethylene receptor genes, MdETR1, MdETR2, MdERS1, and MdERS2, increased in the fruit cortex for both cultivars, but only MdETR2 and MdERS2 increased in the FAZ of ‘Golden Delicious’ apples. The transcript levels of MdPG2, a polygalacturonase gene (PG), and MdEG1, a β-1,4-glucanase gene, markedly increased only in the FAZ of ‘Golden Delicious’ apples, whereas only MdPG1 rapidly increased in the fruit cortex of ‘Golden Delicious’ apples. Our results suggested that MdACS5A, MdACO1, MdPG2, and MdEG1 in the FAZ might be related to the difference in PFA between these two cultivars, whereas MdACS1 and MdPG1 were associated with fruit softening.
Hong Zhu, Eric P. Beers and Rongcai Yuan
Effects of naphthaleneacetic acid (NAA) and aminoethoxyvinylglycine (AVG) on young fruit abscission, leaf and fruit ethylene production, and expression of genes related to ethylene biosynthesis and cell wall degradation were examined in ‘Delicious’ apples (Malus ×domestica Borkh.). NAA at 15 mg·L−1 increased fruit abscission and ethylene production of leaves and fruit when applied at the 11-mm stage of fruit development, whereas AVG, an inhibitor of ethylene biosynthesis, at 250 mg·L−1 reduced NAA-induced fruit abscission and ethylene production of leaves and fruit. NAA also increased expression of 1-aminocyclopropane-1-carboxylate (ACC) synthase genes (MdACS5A and MdACS5B), ACC oxidase gene (MdACO1), and ethylene receptor genes (MdETR1a, MdETR1b, MdETR2, MdERS1, and MdERS2) in fruit cortex and fruit abscission zones. However, AVG reduced NAA-induced expression of these genes except for MdERS2 in fruit abscission zones. NAA increased expression of the polygalacturonase gene MdPG2 in fruit abscission zones but not in fruit cortex, whereas AVG reduced NAA-enhanced expression of MdPG2 in fruit abscission zones. The expression of β-1,4-glucanase gene MdCel1 in fruit abscission zones was decreased by NAA but was unaffected by AVG. Our results suggest that ethylene biosynthesis, ethylene perception, and the MdPG2 gene are involved in young fruit abscission caused by NAA.
Rongcai Yuan, Ulrich Hartmond and Walter J. Kender
The seasonal abscission response of mature `Valencia' oranges [Citrus sinensis (L.)Osb.] to 5-chloro-3-methyl-4-nitro-1H-pyrazole (CMN-Pyrazole) was examined in relation to young fruit, shoot, and root growth. CMN-Pyrazole dramatically increased ethylene production in fruit and effectively reduced the fruit detachment force (FDF), except in a period of reduced response to CMN-Pyrazole in early May. Root growth was inhibited by trunk girdling, in combination with removal of spring vegetative flushes and flowers, but not by their removal alone. During the responsive period, there was no difference in both ethylene production and FDF of CMN-Pyrazole-treated mature oranges between 1) the unmanipulated trees and those manipulated by either 2) girdling, removal of spring flushes and flowers, or 3) removal of flushes and flowers alone. However, during the less-responsive period, ethylene production in CMN-Pyrazole-treated mature oranges was significantly lower while the FDF was higher from non-manipulated trees than from trees treated by either girdling and removal of flush, or only removal of flush. There was no difference in either ethylene production or FDF of CMN-Pyrazole-treated mature oranges between trees manipulated by girdling and removal of flush, and those by removal of flush alone. Flush growth terminated at least 2 weeks before the onset of the less responsive period. This suggests that the hormones from rapidly growing young fruit may be responsible for the less responsive period.
Rongcai Yuan, Ulrich Hartmond and Walter J. Kender
Effects of NAA, TIBA, ethephon, and CMN-Pyrazole on fruit detachment force (FDF) of mature `Valencia' and `Hamlin' orange [Citrus sinensis (L.) Osb.] fruit were examined in 2000 and 2001. NAA effectively inhibited the reduction in FDF or fruit abscission caused by ethephon when applied to the abscission zone 24 hours before ethephon application, but had no significant effect when applied to the fruit without contacting the abscission zone, or to the peduncle ≈4 cm above the abscission zone. TIBA, an auxin transport inhibitor, decreased FDF of mature fruit and promoted fruit abscission when applied alone as a spray to the canopy or directly to the fruit peduncle. This response was dependent on TIBA concentration. TIBA was more effective when applied in combination with ethephon or CMN-Pyrazole than alone. These results are consistent with our previous data that endogenous auxin concentration in the abscission zone of mature `Valencia' orange fruit is one of the factors controlling the sensitivity and thus the responsiveness of the abscission zone of mature fruit to abscission chemicals. Chemical names used: 5-chloro-3-methyl-4-nitro-pyrazole (CMN-Pyrazole); 2-chloroethylphosphonic acid (ethephon); naphthalene acetic acid (NAA); 2,3,5-triiodobenzoic acid (TIBA).