Auxin transport capacity in relation to the dwarfing effect of apple rootstocks

SUMMARY

Basipetal auxin transport in shoots obtained from dwarfing (M.9) and less-dwarfing (MM.Ill) apple rootstocks was compared by measuring the distribution of ¹⁴C-[carboxy]-labelled indole 3-acetic acid supplied to the ends of shoot segments. Auxin was transported at a greater velocity in MM.111 than in M.9 in both woody and non-woody segments, and was effectively blocked by 2,3,5-triiodobenzoic acid. The velocities of transport were approximately 8 mmh-¹ in M.9,and 13 mmh-¹ MM.111. There was relatively little transport in the acropetal direction, and there were no significant differences between the two rootstocks in total auxin uptake. Chromatographic analysis suggested that the metabolismof the auxin supplied followed similar patterns in both rootstocks. It is proposed that the slower auxin transport in M.9 results from a lower capacity for auxin efflux from transporting cells. A limiting auxin transport capacity could explain the reduced cambial activity and subsequent xylem formation in the dwarfing M.9 rootstock, and thus make an important contribution to the dwarfing effect.     

Effect of plant growth promoting rhizobacteria on young apple tree growth and fruit yield under orchard conditions

The effects of rootstocks (M9 and MM 106), cultivars (Granny Smith and Stark Spur Golden) and growth promoting rhizobacteria (OSU-142, OSU-7, BA-8 and M-3) on the tree growth and yield at apple (Malus domestica Borkh) trees were studied in a clay loam soil in the eastern Anatolia region of Turkey in 2002–2004. Plant growth promoting rhizobacteria (PGPR) were capable of producing indole acetic acid (IAA) and cytokinin, but three of them (OSU-7, BA-8 and M-3) were also able to dissolve phosphate. Maximum shoot number of apple trees was found after inoculation with BA-8 followed by OSU-7 and M-3. All the inoculated PGPR strains contributed to the increase in fruit yield of apple when compared to control but it was strongly depended on rootstocks, cultivars and treatments. Plant growth responses were variable and dependent on bacterial strains, rootstock and cultivar and growth parameters evaluated of young apple trees. Newly planted apple trees inoculated with OSU-142, OSU-7, BA-8 and M-3 PGPR increased average shoot length by 59.2, 18.3, 7.0 and 14.3% relative to the control and fruit yield by 116.4, 88.2, 137.5 and 73.7%, respectively. Bacterial inoculation increased shoot diameter from 7.0 to 16.3% when compared to control. The production of plant growth hormones has been suggested as one of the mechanisms by which PGPRs stimulate young apple sapling growth. The growth-promoting effect appears to be direct, with possible involvement of the plant growth regulators indole-3-acetic acid and cytokinin. In view of environmental pollution due to excessive use of fertilizers and high costs of the production of fertilizers, PGPR strains tested in our study have potential to be used for the sustainable and environmentally benign horticultural production.     

苹果紧凑型品种和矮化砧木内源激素的变化

 以不同矮化程度的苹果品种、砧木和嫁接植株为试材, 研究了春梢旺长期茎尖和春梢停长期叶片的内源激素变化。结果表明, 与普通型品种相比, 紧凑型品种茎尖和叶片中的赤霉素(GA₁₊₃) 含量显著降低而细胞分裂素(CTKs) 含量显著升高, 紧凑型品种的CTKs/ GA₁₊₃约是普通型品种的4. 78～6. 24 倍。不同矮化程度的砧木间相比, 春梢停长期叶片中, M9 的IAA 含量比平邑甜茶降低了53. 7 %。以M26作中间砧, 显著降低了接穗叶片和基砧新根中的IAA 含量; 但以新红星作中间砧则影响较小。将M26中间砧与紧凑型品种组合, 与乔砧普通型植株相比, 春梢停长期叶片和新根中IAA 含量分别下降了46. 1 %和33. 3 %,CTKs/ GA₁₊₃是其2. 8～3. 6 倍, 其变化幅度大于单一应用矮化砧、紧凑型品种的植株。因此, 紧凑型、矮化砧具有不同的激素调节机制, GA、CTK在紧凑型品种矮化中起重要作用, 而矮砧的矮化可能与IAA 密切相关。     

Relationships between sap flow,hydraulic conductivity,and the anatomical characteristics of stems and roots in apple rootstocks of different vigour

Summary

To find a useful parameter that could be related to the degree of rootstock vigour, for rootstock breeding programmes, sap flow, hydraulic conductivity, and the anatomical characteristics of stems and roots were measured in 1-year-old trees of apple rootstocks of different vigour. Sap flow was less active in dwarfing rootstocks than in invigorating rootstocks, when taking leaf area into account, under a controlled temperature of 20ºC with the root zone at 15ºC. The hydraulic conductivities of stems and roots were not consistent with the degree of dwarfing.The hydraulic conductivity of the stem appeared to be related to the diameter and density of the large vessels in the xylem. On the other hand, the phloem-to-xylem ratio in roots with diameters of 1.5 – 2.0 mm, was closely-related to sap flow and the degree of dwarfing. The phloem-to-xylem ratio decreased as the degree of vigour increased. Moreover, the phloem-to-xylem ratio was considered to be related to the ability of roots to absorb and transport water, which was expressed as the root-specific sap flow. Therefore, the phloem-to-xylem ratio could be a useful index for evaluating the potential of apple trees as dwarfing rootstocks.     

Stomatal Distribution as an Index for Predicting the Growth Potential of Apple Stocks

In the present study 27 apple rootstocks have been classified into five vigour groups, according to their stomatal distribution, which varied from 11·55 to 5·88 per unit area. Amongst the different stocks studied M. 25 had the maximum stomatal frequency, and M. 9 had the least. An indigenous apple clone M. boccata var. himalacia, very common in Kumaon hills, had 7·20 stomatal counts, i.e. between those for MM. 106 and M. 7. It is suggested that this may be an alternative dwarfing stock for apples.     

Rootstock effect on growth of apple scions with different growth habits

Apple scions with diverse growth habits were grafted on various size-controlling rootstocks and morphological characteristics were measured after 6 years of growth in the field. Scion had more influence than rootstock on monthly growth rate. Across all rootstocks, scions with spreading growth habits grew rapidly in April and May and achieved most seasonal growth earlier than scions with upright growth habits that grew slowly early in the season. In all growth habits and rootstocks, growth rate slowed appreciably but did not cease by August and growth did not terminate earlier for any one scion–rootstock combination. Across all scions, the dwarfing rootstock, M.9, consistently had the lowest and seedling rootstock had the greatest tree height and trunk diameter. However, no one size-controlling rootstock consistently influenced dates of bud break and full bloom, shoot elongation rate, or duration of growth. Tree growth form was not fundamentally affected by rootstock. Significant interactions indicated that effects of size-controlling rootstock on components of shoot growth will vary with apple tree growth habit. These effects on phenology and development can be significant to growers and may assist breeders in developing new apple cultivars.     

新型苹果矮化砧木——SH_1的选育

SH1苹果矮化砧是从国光×河南海棠的杂交实生苗中选育而成。经30 a的田间调查及区试鉴定表明,其做中间砧与基砧八棱海棠、山定子和红星、红富士等苹果品种亲和性好、树体矮化、结果早,尤其是果实着色、风味品质、耐贮性及抗逆性等主要经济性状均超过了M26。SH1矮化砧适宜在黄土高原或气候类似的苹果产区栽培。     

Rootstock and pruning effects on first season dry weight distribution in delicious apple trees

Redchief Delicious apple trees on MM. 106 and M.9 rootstocks were left unpruned, dormant headed at planting, or summer headed 12 weeks after planting, to investigate pruning and rootstock effects on dry weight distribution between roots and shoots during the first growing season. Dormant heading stimulated new shoot dry weight and reduced new root dry weight during the first six weeks after planting. However, by 12 weeks differences in new root dry weight and new shoot dry weight were insignificant. Summer heading resulted in a significant increase in new shoot growth and decrease in new root growth. This caused the root/shoot ratio in headed trees to return to the unpruned value within six weeks after summer heading. New root growth was reduced over a longer time by dormant heading trees on MM. 106 than those on M.9. New shoot growth was increased longer in trees on M.9. Allometric constants (K) of new shoot growth relative to new root growth were significantly changed by dormant heading with both rootstocks. Summer heading had the greatest effect on K values by shifting growth heavily toward new shoots. Root/shoot ratios in all treatments for both rootstocks showed no differences 24 weeks after planting.     

Endogenous hormone concentrations and bud-break response to exogenous benzyl adenine in shoots of apple trees with two growth habits grown on three rootstocks

Summary

Scions from two siblings of a ‘Goldspur Delicious’ × ‘Redspur Delicious’ cross were budded to three rootstocks with different levels of vigour [M.9, M.7, and Malus domestica ‘Antanovka’ seedling] and planted in the field in 1997. The scions had two contrasting growth habits: one with narrow crotch angles, numerous short branches and an upright narrow (UN) canopy; and the other with wide crotch angles, few short branches, and a spreading round (SR) canopy. Shoot tips were collected at the time of bud-break in April 2004 and analysed for auxin (AUX), cytokinins (CK), and abscisic acid (ABA) to determine relationships between scion growth habit, size-controlling rootstock, and shoot tip hormone concentrations. Although not statistically different, the UN growth habit had numerically higher AUX, lower ABA, and equivalent CK levels as the SR growth habit. These differences resulted in statistically higher AUX:CK ratios (ACR). It is possible that the higher ACR contributed to the UN growth habit, which had more anti-gravitrophic shoot growth and appeared to have greater apical dominance than the SR growth habit. Either growth habit, grown on seedling rootstock, had nearly twice the ACR than on M.7 or M.9 rootstocks. The synthetic CK, 6-benzyl adenine (BA), was applied to 30 cm shoot explants of both growth habits in a greenhouse in March 2006. An 8.7 mM BA concentration stimulated bud-break in both growth habits, compared with controls, and bud-break was increased more in the UN than the SR growth habit. The results indicate that the ACR may be a factor regulating bud-break and the development of growth habit in apple scions, and that rootstock modified the hormone concentrations in shoot tips.     

Rootstock influence on ‘delicious’ and ‘golden delicious’ apple fruit quality at harvest and after storage

The influence of 9 rootstocks (M2, M7, M25, M26, MM104, MM106, MM109, MM111 and seedling) on fruit quality at harvest and after storage of ‘Wellspur Delicious’ (WS) and ‘Goldspur’ (GS), and of 3 rootstocks (M7, M26 and MM106) on fruit quality of ‘Red King Delicious’ (RK) and ‘Golden Delicious’ (GD) apple (Malus domestica Borkh.) was evaluated during a 4-year period. Fruits from trees on M26 were larger, developed earlier color and soluble solids (SS), and maintained higher levels of acidity (at harvest and during storage) in comparison with other rootstocks. Fruit from trees on M2 tended to have high SS. Fruit color from trees on MM104, MM106 and MM109 tended to be comparatively poor. There were significant rootstock effects on SS, starch, acidity, color, circumference, weight and box size.     

The Relationship Between Growth Vigour of Rootstock and Phenolic Contents in Apple (Malus × domestica)

The aim of the study is to determine the effect of different growth vigorous rootstocks on phenolic compounds in leaves of apple. For this purpose it was used the leaves of cultivar ‘Red Chief’ grafted on dwarf (M9), semi-dwarf (M26) and semi-vigorous (MM106) rootstocks. During mid-July, the leaf samples were taken from the middle part of annual shoots. Phenolics of the leaves were determined by HPLC analysis. While significant differences among the rootstocks for p-hydroxybenzoic acid, eriodictyol, ferulic acid and p-coumaric acid were detected, these differences were insignificant for gallic acid and quercetin. It was shown that semi-vigorous rootstocks (MM106) had higher phenolic contents in total than the other two dwarf rootstocks. In addition, apigenin-7-glucoside, chlorogenic acid, caffeic acid, rosmarinic acid, epicatechin, syringic acid, catechin, rutin, resveratrol, hesperidin, naringenin, luteolin, apigenin and acacetin could not to be detected. Data showed that there is the relationship between growth vigour and phenolic contents of apple leaves. Especially, p-hydroxy benzoic acid and p-coumaric acid contents were higher in semi-vigorous rootstock than in dwarf rootstock.     

缺锌胁迫对苹果砧木幼苗抗氧化能力和激素含量的影响

 在人工气候室条件下,采用溶液培养法研究了缺锌胁迫对平邑甜茶（Malus hupehensis Rehd.）和小金海棠（M. xiaojinensis Cheng et Jiang）两种苹果砧木幼苗SOD、POD和CAT的活性,膜质过氧化水平及内源激素IAA、GA3、ZR和ABA含量的影响。结果表明,缺锌胁迫下,两品种叶片和根系中SOD活性显著下降,POD和CAT活性、MDA含量不同程度增加,但小金海棠SOD活性下降幅度较平邑甜茶小,POD和CAT活性增加比例大,且MDA积累量低。缺锌使两种砧木幼苗不同器官中IAA、GA3含量显著降低,ZR、ABA含量增加;小金海棠缺锌植株IAA、GA3、ZR含量较平邑甜茶高,ABA含量低。平邑甜茶缺锌植株膜质过氧化水平高,对缺锌胁迫较敏感,小金海棠在缺锌胁迫下抗氧化能力强,促进生长的激素含量高,对缺锌胁迫有较强的抵御和耐受能力;平邑甜茶缺锌植株膜质过氧化水平高,对缺锌胁迫较敏感。     

渭北黄土高原苹果不同砧穗组合幼树根系发育和分布的特征

砧穗组合选配是苹果矮砧集约栽培模式的关键环节之一,幼树的早果性和易成形性是评判砧穗组合优劣的重要指标,这与幼树根系分布密切相关。以中国苹果生产上8种常见砧穗组合的3年生幼树为试材,采用壕沟法对幼树根系分布特征进行了调查分析。结果表明:8种砧穗组合幼树根构型可以分为M系自根砧、SH系自根砧、乔化实生砧木、M系中间砧和SH系中间砧等5大类。3年生幼树根系在水平和垂直方向分布的主要区域,均为0~60 cm范围,且随着距树干距离的增大而减少。不同砧穗组合,根系长度以直径2 mm级别的根系最长,其次是直径2~5 mm的,10 mm以上的最短。乔化实生砧木较无性系矮化砧木粗根多,而须根少;矮化砧木中,M系砧木根系构成以须根为主,而SH系和青砧系砧木须根均较少。砧穗组合幼树的地上分枝数、短枝比例和花芽数等早果性指标与细根(直径2 mm)根系数量、根长密度和根表面积密度呈显著正相关关系。通过对各砧穗组合易成花和易成形的综合评价,认为在渭北黄土高原有灌溉条件的地区,M系自根砧和中间砧组合早果性好和易成形较好,而SH系组合较易成形,但早果性较差,乔化组合早果性最差,这与它们的根系分布特征密切相关。     

Sorten-Unterlagen-Kombinationen f��r den Mostapfelanbau

In a field trial 5 rootstocks (seedling, M 25, A 2, MM 106, MM 111) and 13 varieties were investigated under Northeast German climate conditions to obtain information on the optimal rootstock-variety combinations for intensive juice apple production. Further the sugar and acid content of the fruits were determined. Due to the fact that exclusively scab resistant varieties were examined, the occurrence and stability of the resistance were of special interest. The most favourable rootstocks were M 25 (highest absolute yield) and MM 106 (highest specific yield). The varieties ??Judeline??, ??Rebella??, ??Resi??, ??Angold??, ??Schwechow 3?? and ??Priam?? had the highest absolute yield (mean of all rootstocks). The suitability of these six varieties for juice apple production was described. Although fungicides were used for controlling apple scab, particularly varieties with the Vf resistance factor were infected. A strong scab resistance had the variety ??Reka?? (Vr).     

Einfluss schwach wachsender Apfelunterlagen auf Wachstum,Ertrag und Fruchtgröße verschiedener Sorten – Teil IV: Abschließende Bewertung

From 2001 to 2005 the influence of the dwarfed apple rootstocks M?9 (clone: Burgmer 984), Fleuren 56, Supporter 1, P?16, P?22, J-TE-F, J-TE-E and M?27 on growth, yield and fruit size of the apple varieties ‘Elstar, Elshof’, ‘Boskoop, Herr’ and ‘Jonagored’ was investigated on the locations Auweiler, Dresden-Pillnitz, Osnabrück and Wurzen in Germany.All rootstocks induced a less vegetative growth as the rootstock M?9. In comparison with M?9 the crown volume of the trees ranged from 92% (Supporter 1) to 52% (M?27). The highest yield potential showed the rootstock P?16. Her specific yield (kg/m³) was 17% higher than the specific yield of M?9. All rootstocks produced a lower individual fruit size as M?9. J-TE-E and J-TE-F produced the lowest specific yield.     

Performance of ‘Discovery’ apple on M.9, M.26 and MM.106 rootstocks at two densities

Summary

The apple rootstocks M.9, M.26, and MM.106 were evaluated for their efficiency in bringing ‘Discovery’ apple trees into production. The experiment, carried out over a ten- year period compared two planting densities at 1666 and 3333 trees per ha. Tree vigour differences between rootstocks were measured in term of trunk growth, tree volume, weight of branches pruned off and final weight of the above-ground parts of the trees. Fruit production is presented both as total yield and as the weight of first class fruit. Fruit colour development is also shown. Cropping efficiency is calculated and presented in relation to the different vigour measurements. The results confirm that ‘Discovery’ is slow to come into production. M.9 was the most productive rootstock, but due to vigour differences MM.106 gave the same yields per tree, although the latter had the lower yield efficiency. M.26 performed poorly; its vigour was similar to M.9 but it produced the lowest yields.     

Light penetration,growth, and fruit productivity in ‘Fuji’ apple trees trained to four growing systems

The study was conducted to document the range in soil nutrients, trunk and shoot growth, leaf characteristics and nutrient content, flower bud characteristics, yield, fruit quality, disease occurrence, light penetration, and shoot bark color in a 15-year-old ‘Fuji’ apple (Malus × domestica Borkh.) orchard on M.26 rootstocks trained to four growing systems. SS trees naturally growing had greater width and volume than the LT lightly heading, MLT heavily heading, and S trees thinning and bending. LT trees had more upward terminal shoots at old branches, large numbers of lateral shoots, and longer total shoot length. Leaf [N], [P], and [K] were the lowest for the MLT trees. SS and S trees had greater flower bud density and fruit yield, but LT trees had poor fruit color and less soluble solids and firmness. Percent light penetration into the canopy was the highest with the S system but not different from MLT or SS. The lowest was for the LT system, however, it was not statistically different from MLT or SS.     

Influence of endogenous IAA levels and exogenous IBA on rooting and quality of leafy cuttings of Prunus ‘GiSelA 5’

Summary

The influence of exogenously applied indole-3-butyric acid (IBA) on root and shoot development of leafy cuttings was analysed in Prunus cerasus P. canescens ‘GiSelA 5’, a dwarfing cherry rootstock, in two successive years. Compared to control cuttings, IBA application (4 g l^–1 in 2003; 2.5 g l^–1 in 2004) caused higher indole-3-acetic acid (IAA) accumulation in the cutting bases, but that did not influence the percentage of rooted cuttings, nor their survival in either year. However, IBA inhibited callus formation and, consequently, influenced the quality of the developed cuttings. Callus formation impeded root development, reducing the number of main roots, and inhibited the growth of the cuttings, reducing the average total length of shoots formed by individual cuttings. Callus formation was most reduced in the cuttings in the second experimental year, with high initial IAA concentrations.     

Field resistance of apple rootstocks to Phytophthora cactorum infection

Summary

Six different two year old apple rootstocks with ‘Spur Mcintosh’ scions were inoculated with Phytophthora cactorum and evaluated for susceptibility to this pathogen at the Summerland Research Station from 1982–92. In addition the effect of nursery source on the susceptibility of three rootstocks was compared. The rootstock MM.Ill remained healthy for ten years while MM.106 was very susceptible to P. cactorum crown rot. No significant difference in resistance to P. cactorum was observed between MM.lll and M.4, and M.7a. Fruit yield was significantly lower for MM.106 than with MM.lll, M.4, M.7a, and M.7 in 1990 and 1992. The susceptibility of M.26 from both nurseries to P. cactorum was similar to that of MM.106 from Traas Nursery. A significant difference in susceptibility to P. cactorum was observed between the two sources of MM.106 rootstock from 1984–1990; the MM.106 rootstock from Budwood Orchard was more susceptible to P. cactorum than that from Traas Nursery. However, in 1991 and 1992 (final observation) no significant difference was observed between them for susceptibility to P. cactorum infection. This study indicates that the observed variability in field resistance to P. cactorum crown rot infection should be explored for breeding for P. cactorum resistance.     

不同砧木苹果树细根周转动态的研究

为了分析不同砧木苹果树细根的发生和周转动态,连续4年通过微根管技术研究不同砧木的5年生‘富士’苹果砧穗组合细根中的活根根长密度与死根根长密度的动态变化,以及细根年周转率和季节周转率。结果表明,乔砧树富士/八棱海棠的活根根长密度最大,矮化中间砧富士/M9/八棱海棠和富士/SH40/八棱海棠次之,矮化自根砧富士/M9、富士/SH40和富士/小金海棠最小,所有砧穗组合的活根根长密度随着树龄的增加而逐年减小。所有砧穗组合在夏秋季出现细根发生和死亡高峰期,乔砧树富士/八棱海棠的细根发生高峰期的活根根长密度和死根根长密度均最大。细根年周转率和季节周转率年度间差异大,矮化自根砧树细根的年周转率高于乔砧树。矮化自根砧和矮化中间砧树体ARLD低于乔砧树体可能与其致矮性相关。