Crop load (CL) is a critical regulator of production and quality on apple. It affects leaf photosynthetic rate and usually an increase is detected in leaves close to developing sinks. The objective of this work was to test if 13C discrimination during photosynthesis could be an indicator of carbon sink limitation. The natural plant carbon isotope composition (13C/12C ratio: d13C) is an indicator of water use efficiency and it is an effective tool to study environmental stresses in plants. Seven-year-old trees of Imperial Gala/Bud 9 (n=30), field-grown at the Clarksville Horticultural Research Station in Michigan, were hand-thinned to five levels of CL manipulating the leaf to fruit ratio (LFR: 4, 8, 16, 32, 64) after June drop. Net photosynthetic rate (A) of leaves was monitored daily during the season and elevated rates were observed in low LFR. The A was inhibited in low CL trees (LFR 32 and 64) more in the afternoon (from 20% to 42% in relation to normal CL: LFR 16) than in the morning (from 5% to 20%), and stomatal conductance declined over the afternoon. Shoot and fruit growth were affected (fruit size –11/+11%, shoot length –13/+18% from normal CL; LFR 16). Variations of the stable carbon isotope composition of leaves show a significant reduction of 13C discrimination in low CL trees (–3.2%: d13C –25.82) and an increase of 13C discrimination (+2.7%: d13C –27.38) in relation to normal CL trees (LFR 15.63). The results were similar to those reported in 2004, which imply isotopic discrimination in relation to source limitation. This is the opposite of what you would expect under water stress conditions. Although trees were well-watered during the season, the effect of water stress on apple trees and its interaction with source limitation will be discussed.
Paolo Sabbatini and James Flore
Paolo Sabbatini and G. Stanley Howell
Viticulture in Michigan is limited by a cool and humid climate and as a result, there is a problem of harvest season cluster rot, especially in cultivars with compact cluster morphology. Economically important wine grape varieties in eastern North America possess varying susceptibility to harvest season cluster rot. Some important cultivars that are susceptible are Pinot gris, Pinot noir, Riesling (Vitis vinifera L.) as well as Seyval and Vignoles (French–American hybrids or interspecific hybrid cultivars). A common characteristic of these cultivars is the compactness of the berries held on the cluster rachis. The aim of this work was to determine whether a quantified amount of leaf removal or a temporary reduction in carbon assimilation at the beginning of bloom would reduce fruit set and cluster compactness. Vines subjected to removal of four or six basal leaves had an average fruit set reduction of ≈45% from a non-treated control. Cluster weight and berries per cluster were similarly reduced with a greater effect on the basal than the apical cluster of the shoot. Reduced fruit set was associated with a reduction in cluster compactness and harvest season rot. This was also reflected in yield and basic fruit chemistry parameters associated with the importance of basal leaves to the developing cluster. Multiple applications of stylet oil at different time intervals resulted in significant reduction in net photosynthesis (Pn). A single application had no significant impact on Pn, whereas multiple applications reduced leaf assimilation rates. However, this reduction in Pn did not reduce fruit set or improve cluster compactness. There was a strong negative effect of early leaf removal in Year 1 on vine performance in Year 2; this carryover effect increased shootless nodes per vine, reduced the number of clusters per shoot and per vine, and dramatically reduced fruit set and consequently yield per vine.
Paolo Sabbatini and James A. Flore
The naturally occurring carbon isotope composition (or 13C: 12C ratio, expressed with the notation d13C) of plant tissue may be used as an indicator of water use efficiency during plant growth. d13C has been shown to be an effective tool to study physiological response of plant to environmental conditions, especially water stress. The objective of this work was to test if d13C could be an indicator of carbon limitations or a low source: sink ratio. Trees of `Imperial Gala'/Bud 9 (n = 12), 6-years-old, field grown at the Clarksville Horticultural Research Station (Clarksville, Miss.), were assessed with different crop load (LCL = Low Crop Load, 0.76 ± 0.44 fruit per trunk sectional area (TCA); NCL = Normal Crop Load, 7.25 ± 1.83 fruit/TCA; HCL = High Crop Load, 15.83 ± 1.76 fruit/TCA) and leaf: fruit ratio (LCL: 52.78 ± 8.55, NCL: 13.33 ± 3.06, HCL; 4.31 ± 0.68) immediately following June drop. Net photosynthetic rate of leaves were monitored during the season and elevated rates were observed in NCL and HCL and correlated with the fruiting process. Photosynthesis was inhibited in LCL more in the afternoon (from 20% to 42% in relation to NCL) than in the morning (from 5% to 20%) and this was positively correlated with crop sink strength. Variations of the stable carbon isotope composition of roots (fine and coarse), fruit, leaves, and current-year stems were examined. The d13C varied by tissue (fruit > shoot and leaf > root) and in relation to the level of crop load (d13C‰ in fruit: LCL –23.513 ± 0.248, NCL –24.891 ± 0.594; and HCL –24.935 ± 0.375). These results may have implications for analysis of isotopic signals in carbohydrate stress and fractionation steps will be discussed.
Letizia Tozzini, Paolo Sabbatini, and G. Stanley Howell
Viticulture in Michigan is often limited by cool and humid climate conditions that impact vine growth and the achievement of adequate fruit quality at harvest. Sugars, pH, acids, and yeast available nitrogen (YAN) are indices of quality and, as such, of suitability for wine production. The aim of this study was to evaluate the efficacy of foliar nitrogen (N) fertilization applied as a 1% w/v urea solution at veraison as a method to increase canopy N availability during the fruit ripening stage. To test the effect on different source sink conditions, we imposed three levels of defoliation (0%, 33%, and 66% of leaves removed per vine) and measured net photosynthetic rate (Pn), leaf efficiency parameters, yield components, and fruit quality parameters. Apical leaf Pn was increased by the 33% defoliation (+12% from the undefoliated control) and by the urea application (+6%) 2 weeks after veraison. In basal leaves we observed a reduction in chlorophyll content (SPAD) and maximum photochemical efficiency of PSII (Fv/Fm) as a result of the defoliation treatment and secondarily by the N application, which resulted in a reduction in Pn. Therefore, mean shoot Pn was unaffected by the treatments. Although neither main nor lateral shoot growth was increased by any defoliation treatment, both percent soluble solids (%SS) and berry weight were significantly reduced by the 66% defoliation treatment. Application of urea increased yeast available amino acids by 20% but did not impact %SS or other chemical parameters indicating a different accumulation pathway for sugars and amino acids in the berry.
Martin J. Bukovac, Jerry Hull, and Paolo Sabbatini
For studies on blossom/fruit thinning in apple, tree selection is often based on uniformity of bloom/crop load, assuming that such trees exhibit greater uniformity to treatment. However, the literature is replete with data showing marked variation for a given treatment. We followed variation in bloom/crop density of spur-type `Delicious'/MM.106 and effect of ethephon applied in high crop years on return bloom/yield. Uniform trees (n = 95), under identical cultural practices, were selected for varying crop load. Return bloom, yield and fruit size were monitored over six years. General mean (X) for yield was 94 ± 25 kg/tree and bloom density, rated 1 to 10 (highest), was 5.4 ± 1.7. Annual yield deviated from X by +56 to –40% and bloom density by +49 to –42%. All trees were ranked (decreasing yield) and assigned to five percentile (PCTL) groups (1st, 81-100; 2nd, 61-80; 3rd, 41-60; 4th, 21-40; 5th, 0-20 kg/tree). Trees in each group were reassigned annually to the five PCPL groups for the next five years. Of trees in 1st PCTL (n = 19, X = 187 ± 10 kg/tree) in year one, 5, 5, 24, 0 and 63% placed in PCPL 1, 2, 3, 4, and 5, respectively, in year two. Of trees in 1st PCTL (5%) in year two, all placed in PCTL 2 in year three. Effect of ethephon [200 mg·L-1 at 3, 3 + 6, 3 + 6 + 9 weeks after full bloom (WAFB)] applied in on years to `Redchief', with strong alternate bearing, were evaluated for six years. Ethephon at 3 WAFB had no effect. Yield from multiple applications differed from control (NTC) in off years, but not from each other. Total yield (3 on + 3 off years) for the NTC and ethephon at 3 + 6 WAFB was similar (479 vs. 471 kg/tree). However, 64% of the total yield was produced in the on years and 36% in the off years in NTC vs. 56 and 44% in 3 + 6 WAFB, respectively.
Martin J. Bukovac, Paolo Sabbatini, Philip G. Schwallier, and Michael Schroeder
NAA and BA are important compounds for regulating crop load in apples (Malus domestica Borkh.). When used for fruit thinning, both induce abscission, but at an equivalent crop load NAA tends to reduce and BA to increase fruit size. There is a strong interaction between NAA and BA when used together on ‘Delicious’ and ‘Fuji’, leading to excessive development of pygmy and small fruit (<65 mm diameter). The combination of BA (as Promalin, 1:1 BA + GA4+7) applied at king bloom (KB) and NAAm (amide) at petal fall increased the percentage of small fruit by 3.3- or 5.1-fold compared with BA or NAAm alone. Similar results were obtained with BA (Promalin) at KB oversprayed with NAA at 10 to 12 mm king fruit diameter (KFD). When NAA was oversprayed with BA during fruitlet development, i.e., 5 to 6 mm, 10 to 12 mm, and ≈18 mm KFD, the greatest inhibition of fruit growth occurred at the 10- to 12-mm KFD stage, and there was no significant effect at 18 mm KFD. Inhibition by treatment at the 5- to 6-mm stage was intermediate and trees were overthinned. NAA + BA inhibition of fruit growth in ‘Delicious’ and ‘Fuji’ was not crop load-dependent. In all experiments, crop load (wt basis) of trees treated with NAA + BA was similar or less than of those treated with NAA or BA alone, but they produced 2.5- to 5-fold more small fruit. NAA + BA increased the number of fruit per cluster, many of which failed to fully develop. Increasing the ratio of BA to NAA from 25:15 to 125:15 mg·L−1 increased small fruit formation. The presence of GA4+7 in commercial formulations of BA (0:100, Maxcel; 10:100, Accel; 50:50, Promalin) did not significantly affect the NAA + BA response. Fruit growth was not inhibited by the NAA + BA combination in large-fruited ‘Golden Delicious’ and ‘Jonagold’ and was increased in small-fruited ‘Elstar’ and ‘Gala’ compared with the nontreated control.
Davide Neri, Gianpaolo Mascanzoni, Paolo Sabbatini, Franco Zucconi, and James Flore
To simulate soil sickness, 1-year-old trees of `Golden Delicious' (grafted on M9 and M106) were grown in rhizotrons (1 × 1-m and 0.5-m depth) with different plant residues content, at Ravenna, Italy. Sandy loam soil was used as a substrate. Fine-grounded wood from apple and peach residues (6 kg per rhizotron) was mixed to the substrate and considered as main treatment. Mature compost (1% and 2.5% in volume) was added or not to the substrate with the organic residues and considered as subtreatment. The application of residues was localized either near the soil surface (0–25 cm) or deeper in the soil profile (25–50 cm). In each rhizotron, four trees on the same rootstock were planted and each soil treatment was replicated twice. After 2 years, the roots were accurately excavated (washing off the soil with water), and growth was measured. The presence of apple residues near the soil surface induced a 5% to 20% reduction of shoot growth. The reduction per plant dry weight was higher when trees were grafted on M106. At root level, the presence of residues increased the root migration in the search for fresh niches, enhancing root crossing and anastomosis. Both these shoot and root conditions are typical of replant diseases symptoms. The localization of apple residues in the lower part of the profile reduced the symptoms and so did the addition of compost. The peach residues did not affect shoot growth when compared to the control, but the shoot-to-root ratio was reduced, indicating a tendency to increase root migration.
Philip G. Schwallier, Paolo Sabbatini, and Martin J. Bukovac
The `Honeycrisp' apple has unique characteristics favored by consumers that has provided exceptional return to growers. This cultivar also has some traits that challenge plant management. There appears to be a strong inhibitory effect of crop load on flower initiation and thus annual cropping. We studied the relationship between fruit load, established by post-bloom hand and chemical thinning, and effect of ethephon and gibberellin (GA4+7) on flower initiation and thereby annual cropping. Initially, return bloom (RB) was related to previous season's crop load in three thinning studies on 3- and 9-year-old `Honeycrisp'/M 9, Pajam 1 trees. The RB density was rated 1–10 on trees (n=172), which produced 0-60 kg of fruit/tree. Return bloom ratings (RBR) on the 3-year-old trees ranged from 0–9. Percentage of trees with RBR >5 for previous season's yield of <5 kg, 5–10, 10–15, and 15–20 was 70, 9, 2.5, and 0, respectively. There was dramatic inhibition of flowering at a crop load of >5 kg/tree. In the second study (9-year-old trees), crop load ranged from 15–60 kg/tree (n=24). RBR for trees in the 30–40 kg/tree class ranged from 0–8 with high variability. Thirty-one percent of trees with crop load between 20–30 kg had RBR of 5–8, and 26% between 0–5. Twenty-one percent of trees in the 30–40 kg/tree class flowered and all but one had a RBR of 5 or less. Yields ranged from 22–81 kg/tree in the third study (n=60); crop load was normally distributed among trees. Flower initiation was almost completely inhibited. Fifty percent of the trees did not flower; the remainder had a RBR of <1, i.e. <10%. In the ethephon/GA study, RBR ranged 8–10 on trees producing <12 kg/tree, then decreased rapidly to <2 for yields of 25–50; greatest variation at 20–40 kg.
Vania Lanari, Oriana Silvestroni, Alberto Palliotti, Alan Green, and Paolo Sabbatini
Several studies have investigated water relationships in grapevines, but the responses to water limitation on individual leaves developed in different shoot positions are scarce in the literature. To begin to fill-in this gap, we examined the adaptive responses of vines at the leaf level to varying amounts of water stress using young hybrid ‘Vignoles’ in a controlled growth chamber. We found that the reduction in water availability to 40% of daily evapotranspiration limited shoot and leaf growth, affecting leaf number, shoot elongation, and leaf area. After 2 days of water stress we observed young developing leaves (nodes six to eight from the shoot apex) to have drastically reduced stomatal conductance (g S, about 20 mmol H2O/m2/s) and net photosynthesis (Pn, 2 μmol CO2/m2/s). On the 4th day Pn in mature leaves (nodes 9 to 12 from the shoot apex) fell to values below 2 μmol CO2/m2/s. After 6 days, both Pn and g S stabilized at lower values with fluctuations related only to leaf position along the shoot axis. Young leaves revealed substantial enrichment of carbon-13 (13C) and high water-use efficiency suggesting a higher and faster adaptive capacity to water shortage conditions as compared with mature leaves.
Martin J. Bukovac, Paolo Sabbatini, Franco Zucconi, and Phillip G. Schwallier
Alternate cropping is a common physiological trait in several important apple cultivars, including spur-type ‘Delicious’. Alternate bearing usually develops slowly in apple trees as they mature, but may be dramatically induced by environmental or biotic stresses. We describe the native temporal and within-year variation of bloom and yield of a highly uniform spur-type ‘Delicious’/M.106 population (n = 95) over a period of 9 years. Crop load was adjusted by hand in the first year to establish a defined normal fruit population distribution. Thereafter, all trees received identical practices. Bloom density (BD) was rated (1 to 10) and yield and fruit size distribution were determined annually on an individual tree basis. Temporal profiles for bloom and yield had four periods above and below the general mean (GM). There were two 3-year periods of nonbienniality. Annual variation in BD ranged from 3.1 to 8.0 and in yield from 54 to 168 kg/tree. Variation in cropping was greater when expressed as percentage deviation from the GM. Annual mean fruit weight was inversely related to yield, but percentage of small- (51 to 64 mm) or large-diameter fruit (70 to 82 mm) was not consistently related to yield. The within-year cv ranged from ≈11% to 66% for BD and from 13% to 42% for yield. The degree of synchrony (within-year variation) was lowest in the year after crop adjustment (to normal distribution), became highly synchronized in ≈4 years, and then decreased. The relationship of native variation of individual trees to the population and to flower initiation and fruiting are discussed in relation to the alternate bearing cycle and significance in selecting trees for experimentation.