Trans-jasmonic acid (JA), cis-JA, and trans-methyl jasmonate (MeJA) were quantified in pulp and seeds of `Tsugaru' apple [Malus sylvestris (L.) Mill. var. domestica (Borkh.) Mansf.] and `Satohnishiki' sweet cherry (Prunus avium L.). Trans-JA and cis-JA showed similar changes during development in both types of fruit. JA concentration was high in the early growth stages of apple pulp development, decreased with days after full bloom (DAFB), and then increased again during maturation. There was an initial decrease in concentration of MeJA in apple pulp, followed by a general increase towards harvest. Concentrations of JA and MeJA in the pulp of sweet cherry were high during early growth stages, then decreased towards harvest. PDJ treatment at 104 DAFB (preclimacteric stage) increased endogenous abscisic acid concentration and anthocyanin concentration at 122 and 131 DAFB (maturation stages) in apple. JA concentration in apple seeds was also high in the early growth stages, then decreased, and finally peaked at harvest. MeJA concentration in apple seeds increased towards harvest. In the seeds of sweet cherry, JA and MeJA concentrations generally increased until harvest. In both types of fruit, concentrations of JA and MeJA in the seeds were higher than those of pulp. On a dry weight basis, changes in concentration in the seeds preceded those in the pulp. These results demonstrate that relatively high amounts of JA and MeJA are associated with young developing fruit. These substances may have a role in regulation of fruit growth at early growth stages, though this has not been demonstrated. Chemical name used: n-propyl dihydrojasmonate (PDJ).
Satoru Kondo, Akihiro Tomiyama, and Hideharu Seto
Sunghee Guak, Michael Beulah, Norman E. Looney, and Leslie H. Fuchigami
Three experiments were conducted at two locations, two at Summerland, British Columbia, Canada and one at Corvallis, Ore., to evaluate synthetic auxins (MCPB-ethyl or NAA) and ethephon as blossom thinners for `Fuji' apple [Malus sylvestris (L.) Mill var. domestica (Borkh.) Mansf.]. These experiments also involved application of carbaryl at 1000 mg·L-1 in the postbloom period. All blossom thinners were sprayed at 85% full bloom while carbaryl was applied at 11-mm fruit diameter. Within these experiments, MCPB-ethyl at up to 20 mg·L-1 or NAA at up to 21 mg·L-1 increased whole flower cluster removal linearly with rate; however, with the Corvallis experiment MCPB-ethyl failed to result in any thinning. Neither auxin treatment consistently reduced fruit set on the remaining clusters, resulting in “clustering”. Bloom-time application of ethephon at 100 mg·L-1 with NAA further reduced crop load. Carbaryl reduced total crop load by increasing both whole cluster removal and number of sites with a single fruit. Return flowering was not improved by the auxin treatments except where there was very excessive crop reduction. Ethephon or carbaryl promoted return flowering with the carbaryl effect being more pronounced. However, this carbaryl effect was significantly countered by the bloom-time auxin whereas ethephon overcame the negative effects of the auxin treatments. The combined use of ethephon and carbaryl was effective in terms of both crop reduction and return flowering benefits. Chemical names used: 1-naphthyl N-methylcarbamate (carbaryl); 2-chloroethylphosphonic acid (ethephon); ethyl 4-(4-chloro-2-methylphenoxy) butanoate (MCPB-ethyl); and 2-(1-naphthyl) acetic acid (NAA).
Lailiang Cheng, Leslie H. Fuchigami, and Patrick J. Breen
Bench-grafted `Fuji' apple [Malus sylvestris (L.) Mill. var. domestica (Borkh.) Mansf.] trees on Malling 26 (M.26) rootstocks were fertigated for 6 weeks with N concentrations ranging from 0 to 20 mm. These treatments produced levels of leaf N ranging from 0.9 to 4.3 g·m-2. Over this range, leaf absorptance increased curvilinearly from 74.8% to 92.5%. The light saturation point for CO2 assimilation expressed on the basis of absorbed light increased linearly at first with increasing leaf N, then reached a plateau at a leaf N content of ≈3 g·m-2. Under high light conditions (photosynthetic photon flux of 1500 μmol·m-2·s-1), the amount of absorbed light in excess of that required to saturate CO2 assimilation decreased with increasing leaf N. Chlorophyll fluorescence measurements revealed that the maximum photosystem II (PSII) efficiency of dark-adapted leaves was relatively constant over the leaf N range, except for a slight decrease at the lower end. As leaf N increased, nonphotochemical quenching declined under high light, and there was an increase in the efficiency with which the absorbed photons were delivered to open PSII centers. The photochemical quenching coefficient remained high except for a decrease at the lower end of the leaf N range. Actual PSII efficiency increased curvilinearly with increasing leaf N, and was highly correlated with light-saturated CO2 assimilation. The fraction of absorbed light potentially going into singlet oxygen formation was estimated to be ≈10%, regardless of leaf N status. It was concluded that there was more excess absorbed light in low N leaves than in high N leaves under high light conditions. Nonphotochemical quenching was enhanced with decreasing leaf N to reduce both the PSII efficiency and the probability of damage from photooxidation by excess absorbed light.
Georgios Psarras, Ian A. Merwin, Alan N. Lakso, and John A. Ray
A 2-year field study of `Mutsu' apple [Malus sylvestris (L.) Mill. var. domestica (Borkh.) Mansf.] on `Malling 9' (M.9) rootstock was conducted to observe root growth in situ, and compare patterns of root growth, root maturation and turnover rates, and soil-root respiration. Rhizosphere respiration was monitored with a portable chamber connected to an infrared gas analyzer; root emergence, browning, and turnover rates were measured by direct observation through minirhizotron tubes inserted in the root zone. Negligible root growth was observed before the onset of shoot growth in mid-May. In both years, a main peak of new root emergence in late June and early July coincided partially with major phases of shoot and fruit growth. A smaller peak of root emergence during August to September 1997 consisted primarily of new roots at 20 to 45 cm soil depths. Most roots remained <1 mm in diameter and developed in the upper 25 cm soil profile; no roots were observed at any time below 50 cm, due to a compacted soil layer at that depth. The cumulative survivorship of new roots was 38% in 1996 and 64% in 1997, and 50% of emergent white roots turned brown or senesced within 26 days in 1996 and 19 days in 1997. Root turnover rates were highest in mid-August both years. Rhizosphere respiration was correlated (r 2 = 0.36 and 0.59, P = 0.01 and 0.004) with soil temperatures in 1996 and 1997, with Q10 values of 2.3 in both years. The Q10 for root-dependent respiration (the difference between soil only and combined soil-root respiration) in 1997 was 3.1, indicating that roots were more sensitive than soil microflora to soil temperature. The temporal overlap of high rates of shoot, root and fruit growth from late May to mid-July suggests this is a critical period for resource allocations and competition in temperate zone apple trees.
Georgios Psarras and Ian A. Merwin
One-year-old potted `Mutsu' apple [Malus sylvestris (L.) Mill. var. domestica (Borkh.) Mansf.] trees on scion invigorating Malling-Merton 111 (MM.111) and scion dwarfing Malling 9 (M.9) rootstocks were grown outdoors in containers under three levels of water availability (irrigated at -20, -80, and -200 kPa) to investigate the effects of soil water availability on combined soil/root (rhizosphere) respiration rates, and developmental morphology of root systems. Rhizosphere respiration was measured with a portable infrared gas analyzer, and root biomass was estimated by electrical capacitance. These nondestructive measurements were compared with final root dry weights of harvested trees, to determine their reliability for estimating relative differences in root biomass. Water stress reduced final biomass similarly for both rootstocks, but the relative reduction in shoot growth was greater for MM.111. Root to shoot ratios were higher and average specific root respiration was lower for M.9 rootstock compared with MM.111. M.9 appeared to be more tolerant of water stress then MM.111, due to reduced canopy transpiration relative to root system mass. Water stress increased root to shoot ratios, specific root length, and the carbohydrate costs of root maintenance as indicated by specific respiration rates. Root dry weight (DW) was better correlated to rhizosphere respiration than to root electric capacitance. The observed r 2 values between root capacitance and root DW were as high as 0.73, but capacitance measurements were also influenced by soil water content and rootstock type. Electrical capacitance estimated total root biomass more accurately for M.9 than for MM.111.
Robert A. Saftner, William S. Conway, and Carl E. Sams
Effects of postharvest pressure infiltration of distilled water, CaCl2 solutions at 0.14 or 0.27 mol·L-1 without and with subsequent fruit coating treatments of preclimacteric `Golden Delicious' [Malus sylvestris (L.) Mill. var. domestica (Borkh.) Mansf. `Golden Delicious'] apples on volatile levels, respiration, ethylene production, and internal atmospheres after storage at 0 °C for 1 to 6 months, and during subsequent shelf life at 20 °C were investigated. Over 30 volatiles were detected, most of the identified volatiles were esters; the rest were alcohols, aldehydes, ethers, a ketone, and a sesquiterpene. Pressure infiltration of water and increasing concentrations of CaCl2 resulted progressively in reduced total volatile levels, respiration, ethylene production, and internal O2 levels and increased CO2 levels in fruit following 2 to 4 months storage in air at 0 °C. Total volatile levels, respiration, ethylene production, and internal atmospheres of CaCl2-treated apples at 0.14 mol·L-1 gradually recovered to nontreated control levels following 2 weeks of shelf life at 20 °C and/or storage at 0 °C in air for more than 4 months. Following the calcium treatments with a shellac- or wax-based coating had similar but stronger and more persistent effects on volatile levels, respiration, ethylene production, and internal atmospheres than those found in fruit treated with CaCl2 alone. Calcium infiltration did not change the composition of volatile compounds found in fruit. Results suggest that pressure infiltration of `Golden Delicious' apples with CaCl2 solutions transiently inhibited volatile levels, respiration, and ethylene production, in part, by forming a more-or-less transient barrier to CO2 and O2 exchange between the fruit tissue and the surrounding atmosphere.
Susan Lurie, Amnon Lers, Zohar Shacham, Lilian Sonego, Shaul Burd, and Bruce Whitaker
Untreated control, 1-methylcyclopropene (1-MCP)-treated, and heated fruit of the superficial scald-susceptible `Granny Smith' cultivar of apple [Malus sylvestris (L.) Mill. var. domestica (Borkh.) Mansf.] were compared with respect to scald incidence, internal ethylene concentration (IEC), α-farnesene metabolism, expression of the genes AFS1, which encodes α-farnesene synthase, the final, rate-limiting enzyme in the α-farnesene biosynthetic pathway, and HMG2 and HMG3, which encode isozymes of 3-hydroxy-3-methylglutaryl-coenzyme A reductase, the proposed rate-limiting enzyme in the mevalonate pathway of isoprenoid synthesis. The incidence of scald in untreated `Granny Smith' apples after 16 weeks at 0 °C plus 1 week at 20 °C was 100%; 1-MCP treatment prevented scald development, whereas heat treatment delayed and reduced scald development. 1-MCP also inhibited both α-farnesene and IEC, suggesting that ethylene induces transcription of key genes involved in α-farnesene biosynthesis. Heat treatment reduced levels of α-farnesene and and its oxidation products, conjugated trienols (CTols), but not to the extent of 1-MCP. Internal ethylene concentrations in heated apples did not differ from those in the controls. In both control and heated fruit, a sharp increase in AFS1 mRNA during the first 4 weeks of storage preceded an increase in α-farnesene and a subsequent increase in CTols. AFS1 transcript was absent from 1-MCP-treated apples for the first 10 weeks of storage, and even at 16 weeks was lower than in heated and untreated control fruit. Levels of the HMG2 and HMG3 transcripts varied during storage and among treatments, and were not correlated with the incidence of scald. HMG2 mRNA transcript accumulation was low at harvest and increased in abundance during storage in all treatments, with the greatest increase occurring in 1-MCP-treated fruit. In contrast, HMG3 transcript was constitutively present at all storage times, although it too was slightly more abundant in 1-MCP-treated fruit.
Kathleen Delate, Andrea McKern, Robert Turnbull, James T.S. Walker, Richard Volz, Allan White, Vincent Bus, Dave Rogers, Lyn Cole, Natalie How, Sarah Guernsey, and Jason Johnston
By 2003, organic apple [Malus sylvestris (L.) Mill var. domestica (Borkh.) Mansf.] production had increased to 5626 ha in the United States and to 2964 ha in New Zealand by 2002. Common problems facing organic apple growers in the humid regions of New Zealand and the United States include effective management strategies for apple scab [Venturia inaequalis (Cooke)] and insect pests. Experiments conducted in Iowa in 2003–2004 demonstrated the effectiveness of a kaolin clay- and spinosad-based insecticide program in maintaining codling moth [Cydia pomonella (L.)] damage levels to less than 5% in the scab-resistant cultivars Enterprise, Liberty, Redfree, and Gold Rush. Similar pest management systems have been developed in New Zealand to comply with export standards and quarantines. The use of codling moth granulosis virus and a spinosad-based insecticide have led to reduced pest pressure and to an increase in organic exports with a 41% premium price over conventional apples. However, an association between spinosad use and woolly apple aphid [Eriosoma lanigerum (Hausmann)] population increase was observed in organic orchard surveys in 2006. An alternative to spinosad applications, insect disinfestation through controlled atmosphere (CA) treatment, was investigated to control quarantined pests and to extend the storage potential of scab-resistant cultivars. A CA treatment of 9 weeks of 2% O2 and 2% CO2 at 0.5 °C was determined to maintain firmness ratings to export standards in CA-stored, scab-resistant ‘Pinkie’ apples and to decrease internal ethylene concentration by 84% compared with apples stored in air. In addition, new scab-resistant cultivars with ‘Pinkie’ background under development in New Zealand show promise for organic production in humid regions. Few fruit quality differences were determined between ‘Pinkie’ fruits from integrated fruit production and organic production systems, although premium prices exist only for certified organic apples.
Elena de Castro, William V. Biasi, and Elizabeth J. Mitcham
Apple [Malus ×sylvestris (L.) Mill. var. domestica (Borkh.) Mansf. ‘Cripps Pink’] fruit were harvested yearly, at two or three maturity stages, from the same California orchard in 2002 through 2005. Fruit firmness, soluble solids, titratable acidity, background color, and percent blush were correlated with the starch pattern index at harvest. Fruit from each harvest were stored at 0.5 ºC in air or in a controlled atmosphere (CA) with 1.5 or 2 KPa O2 in combination with 1, 3, and 5 KPa CO2. Subsets of fruit were treated with 1 μL·L−1 1-methylcyclopropane for 24 hours at 0 ºC or 2200 μL·L−1 diphenylamine (DPA) for 5 minutes. Ethylene production was measured for 30 days after harvest. Ethylene concentration in the storage atmosphere was also monitored during storage. Fruit quality was evaluated after storage plus 5 days of ripening at 20 ºC. Fruit in a CA with 1 or 3 KPa CO2 maintained firmness and green background color, and produced less ethylene during ripening at 20 ºC than fruit stored in a CA with 5 KPa CO2; however, quality of all CA-stored fruit was better than air-stored fruit. Flesh browning developed only in CA storage, appearing by 2 months and not increasing in incidence with further storage periods. 1-Methylcyclopropane conserved fruit quality in air as well as CA during 4 months of storage, and DPA-treated fruit were firmer after CA storage, but similar after air storage, compared with untreated fruit. Diphenylamine did not control a stem-end scald disorder, which increased with time in storage and affected more than 80% of the fruit after 6 months of air storage.
Luiz C. Argenta, Xuetong Fan, and James P. Mattheis
The efficacy of the ethylene action inhibitor 1-methylcyclopropene (1-MCP) applied in water to slow ripening of ‘Golden Delicious’ [Malus sylvestris var. domestica (Borkh.) Mansf.] apples was evaluated in comparison with 1-MCP applied in air. The material was applied by dipping fruit in 1-MCP water solutions (0.03, 0.3, or 3 mmol·m−3) for 4 min or by exposing fruit to 1-MCP gas (0.42, 4.2, or 42 μmol·m−3) in air for 12 h. Fruit were held in air at 20 °C for 25 days after treatment or stored at 0.5 °C in air for up to 6 months followed by 7 days in air at 20 °C. Application of 1-MCP in water or air delayed the increase in respiration and ethylene production associated with fruit ripening and reduced the amount of fruit softening, loss of acidity, and change in peel color. Treatments applied in water required a 700-fold higher amount of active ingredient compared with treatments applied in air to induce similar physiological responses. Fruit responses to 1-MCP varied with treatment concentration, and the maximum effects were obtained at concentrations of 4.2 or 42 μmol·m−3 in air and 3 mmol·m−3 in water. Peel color change was impacted less than retention of firmness and titratable acidity for 1-MCP treatments applied at concentrations of 4.2 or 42 μmol·m−3 in air and 0.3 or 3 mmol·m−3 in water. Treatment with 1-MCP in air or water was less effective for slowing peel degreening when treated fruit were stored at 0.5 °C compared with storage at 20 °C. Fruit treated with 1-MCP and stored in air at 0.5 °C developed a peel disorder typified by a gray·brown discoloration that is unlike other disorders previously reported for this cultivar. Symptoms were present when fruit were removed from cold storage and no change in symptom appearance was observed during a 7-d holding period at 20 °C.