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Modelling branching patterns on 1-year-old trunks of six apple cultivars.

The structure resulting from branching on 1-year-old apple tree trunks was analysed in a set of apple cultivars with diverse branching and fruiting habits. Four different lateral types borne on successive nodes were observed when vegetative and flowering fates, as well as sylleptic and proleptic branching, were taken into account. The location and grouping of lateral types along the trunk were analysed for all cultivars, but are detailed for one cultivar only. This cultivar showed a succession of zones, each zone being characterized by its composition of lateral types. Statistical models-hidden semi-Markov chains-were built to take this structure into account and to characterize the cultivar's specific branching pattern. The models showed that most of the branching zones had a similar location in the different cultivars, even though zone composition and zone length differed among cultivars. On a more detailed scale, the nodes bearing a lateral, regardless of its type, were frequently followed by latent buds. The validity of the models and their biological interpretation are discussed with respect to parent shoot dynamics, hormonal gradients and competition between neighbouring buds.

Malus↗

Application of architectural analysis and AMAPmod methodology to study dwarfing phenomenon: the branch structure of 'Royal Gala' apple grafted on dwarfing and non-dwarfing rootstock/interstock combinations.

Architectural analysis was applied to study branch development of 'Royal Gala' apple trees grafted with dwarfing and non-dwarfing rootstock/interstock combinations, which had been chosen to produce trees with a wide range of vigour. Using AMAPmod methodology, the structure of 3-year-old branches was described at four levels of representation: branch; annual shoot; growth unit; and node. Three types of growth units were distinguished: extension growth unit (vegetative unit with internode extension); vegetative spur with minimal internode extension; and fruiting spur or bourse. The aim of the analysis was to describe exactly how the rootstock/interstock combinations affected the structure building process. The number of extension growth units, vegetative spurs and fruiting spurs per annual shoot changed over the years, but this was not affected by rootstock/interstock combination. Compared with MM.106 rootstock, M.9 rootstock reduced the number of nodes per extension growth unit. In most cases, rootstock/interstock combination had no effect on the linear relationship between extension growth unit length and node number (R(2) = 0.88). Average internode length depended on unit node number, with internodes being shorter for units with fewer nodes. Thus the difference in apple branch size induced by the rootstock/interstock combinations was mainly due to a reduction in the length and number of neoformed nodes produced on extension growth units. As percentage budbreak of axillary buds on extension growth units was not affected by rootstock/interstock combination, differences in numbers of axillary annual shoots per branch were entirely due to differences in the total numbers of nodes extended during the previous year.

Algorithms↗

Co-ordinated growth between aerial and root systems in young apple plants issued from in vitro culture.

BACKGROUND AND AIMS: In several species exhibiting a rhythmic aerial growth, the existence of an alternation between root and shoot growth has been demonstrated. The present study aims to investigate the respective involvement of the emergence of new organs and their elongation in relation to this phenomenon and its possible genotypic variation in young apple plants. METHODS: Two apple varieties, X6407 (recently named 'Ariane') and X3305 ('Chantecler' x 'Baujade'), were compared. Five plants per variety, issued from in vitro culture, were observed in minirhizotrons over 4 months. For each plant, root emergence and growth were observed twice per week. Growth rates were calculated for all roots with more than two segments and the branching density was calculated on primary roots. On the aerial part, the number of leaves, leaf area and total shoot length were observed weekly. KEY RESULTS: No significant difference was observed between varieties in any of the final characteristics of aerial growth. Increase in leaf area and shoot length exhibited a 3-week rhythm in X3305 while a weaker signal was observed in Ariane. The primary root growth rate was homogeneous between the plants and likewise between the varieties, while their branching density differed significantly. Secondary roots emerged rhythmically, with a 3-week and a 2-week rhythm, respectively, in X3305 and 'Ariane'. Despite a high intra-variety variability, significant differences were observed between varieties in the secondary root life span and mean length. A synchronism between leaf emergence and primary root growth was highlighted in both varieties, while an opposition phase was observed between leaf area increments and secondary root emergence in X3305 only. CONCLUSION: A biological model of dynamics that summarizes the interactions between processes and includes the assumption of a feedback effect of lateral root emergence on leaf emergence is proposed.

Cells, Cultured↗

A fast DNA sequence handling program for Apple II computer in BASIC and 6502 assembler.

A fast general purpose DNA handling program has been developed in BASIC and machine language. The program runs on the Apple II plus or on the Apple IIe microcomputer, without additional hardware except for disk drives and printer. The program allows file insertion and editing, translation into protein sequence, reverse translation, search for small strings and restriction enzyme sites. The homology may be shown either as a comparison of two sequences or through a matrix on screen. Two additional features are: (i) drawing restriction site maps on the printer; and (ii) simulating a gel electrophoresis of restriction fragments both on screen and on paper. All the operations are very fast. The more common tasks are carried out almost instantly; only more complex routines, like finding homology between large sequences or searching and sorting all the restriction sites in a long sequence require longer, but still quite acceptable, times (generally under 30 s).

Base Sequence↗

Cloudy apple juice decreases DNA damage, hyperproliferation and aberrant crypt foci development in the distal colon of DMH-initiated rats.

Clear (CleA) and cloudy (CloA) apple juices containing different amounts of analyzed procyanidins and pectin were investigated for preventive effects of colon cancer and underlying molecular mechanisms in F344 rats given intraperitoneal injections of 1,2-dimethylhydrazine (DMH; 20 mg/kg body wt) once a week for 4 weeks. Rats received either water (Cont), CleA or CloA (ad libitum) for 7 weeks starting 1 week before the first DMH injection. CloA inhibited DMH induced genotoxic damage in mucosa cells of the distal colon compared with Cont as investigated by single-cell microgel electrophoresis assay. The mean tail intensity in mucosa cells of DMH-treated controls (Cont/DMH: 6.1+/-0.9%) was significantly reduced by CloA (2.4+/-0.8%; P<0.01) but not by CleA intervention (4.1+/-1.2%; P>0.05). The crypt cell proliferation index induced by DMH (Cont/NaCl: 10.0+/-0.7%; Cont/DMH: 19.9+/-1.0%; P<0.001) was significantly decreased by CleA (15.7+/-0.7%; P<0.001) and CloA intervention (11.9+/-0.4%; P<0.001). CloA but not CleA significantly reduced the number of large aberrant crypt foci (ACF) consisting of more than four aberrant crypts (AC) (Cont/DMH: 37.4+/-5.4; CleA/DMH: 32.8+/-4.4, P>0.05; CloA/DMH: 18.8+/-2.5 ACF; P<0.05) and the overall mean ACF size in the distal colon (Cont/DMH: 2.31+/-0.09; CleA/DMH: 2.27+/-0.05; CloA/DMH: 2.04+/-0.03 AC/ACF; P<0.05). After treatment with DMH and/or apple juices there were no changes in transcript levels of colonic cyclooxygenase isoforms (COX-1, COX-2) or glutathione-associated enzymes (GST-M2, gamma-GCS, GST-P), the splenocyte natural killer cell activity and plasma antioxidant status. However, CloA but not CleA prevented the DMH-induced reduction of splenocyte CD4/CD8 (T-helper cells to cytotoxic lymphocytes) ratio. Since both formulations contained comparable concentrations and types of monomeric polyphenols, complex polyphenols or non-polyphenolic compounds, such as pectin might be responsible for the stronger cancer-preventive effect by CloA.

Animals↗

The neurodegeneration mutant löchrig interferes with cholesterol homeostasis and Appl processing.

The novel Drosophila mutant löchrig (loe) shows progressive neurodegeneration and neuronal cell death, in addition to a low level of cholesterol ester. loe affects a specific isoform of the gamma-subunit of AMP-activated protein kinase (AMPK), a negative regulator of hydroxymethylglutaryl (HMG)-CoA reductase and cholesterol synthesis in vertebrates. Although Drosophila cannot synthesize cholesterol de novo, the regulatory role of fly AMPK on HMG-CoA reductase is conserved. The loe phenotype is modified by the level of HMG-CoA reductase and suppressed by the inhibition of this enzyme by statin, which has been used for the treatment of Alzheimer patients. In addition, the degenerative phenotype of loe is enhanced by a mutation in amyloid precursor protein-like (APPL), the fly homolog of the human amyloid precursor protein involved in Alzheimer's disease. Western analysis revealed that the loe mutation reduces APPL processing, whereas overexpression of Loe increases it. These results describe a novel function of AMPK in neurodegeneration and APPL/APP processing which could be mediated through HMG-CoA reductase and cholesterol ester.

Animals↗

Pest phenology and evaluation of traps and pheromone lures for monitoring flight activity of obliquebanded leafroller (Lepidoptera: Tortricidae) in Minnesota apple orchards.

The efficacy of two trap designs (Pherocon 1C "wing" trap versus Pherocon VI "delta" trap) and two pheromone blends for monitoring obliquebanded leafroller, Choristoneura rosaceana (Harris), was evaluated in Minnesota apple orchards during the 2001 and 2002 field seasons. Two distinct flight periods of C. rosaceana were recorded yearly in Minnesota. Overwintered C. rosaceana larvae resumed activity in the spring at approximately 60 degree-days (DD) base 10 degrees C, whereas adult emergence began approximately 275 DD base 10 degrees C. To determine the optimal pheromone blend for monitoring C. rosaceana in Minnesota, traps were baited with either a three-component pheromone blend (Z11-14:Ac, E11-14:Ac, and Z11-14:OH) produced by females in eastern North America, or a four-component blend (Z11-14:Ac, E11-14:Ac, Z11-14:OH, and Z11-14:Al) commonly produced by females in western North America. Of the four pheromone-baited traps evaluated, delta traps baited with the four-component western pheromone lure captured the highest number of C. rosaceana males, followed by wing traps baited with western lure. Male C. rosaceana were less attracted to traps containing the three-component eastern lure, and both lure types seemed to be considerably selective against sympatric redbanded leafroller, Argyrotaenia velutinana (Walker). These results suggest that the pheromonal response of the predominant endemic population of C. rosaceana in Minnesota is similar to the response of the pest in many parts of western North America. The delta trap baited with western pheromone lure of C. rosaceana is recommended for monitoring the pest in Minnesota, and the results are discussed in relation to the development of effective management strategies against this important pest of apple.

Animals↗

Rubisco activation state decreases with increasing nitrogen content in apple leaves.

Based on the curvilinear relationship between leaf nitrogen content and the initial slope of the response of CO(2) assimilation (A:) to intercellular CO(2) concentrations (C:(i)) in apple, it is hypothesized that Rubisco activation state decreases with increasing leaf N content and this decreased activation state accounts for the curvilinear relationship between leaf N and CO(2) assimilation. A range of leaf N content (1.0-5.0 g m(-2)) was achieved by fertilizing bench-grafted Fuji/M.26 apple (Malus domestica Borkh.) trees for 45 d with different N concentrations, using a modified Hoagland's solution. Analysis of A:/C:(i) curves under saturating light indicated that CO(2) assimilation at ambient CO(2) fell within the Rubisco limitation region of the A:/C:(i) curves, regardless of leaf N status. Initial Rubisco activity showed a curvilinear response to leaf N. In contrast, total Rubisco activity increased linearly with increasing leaf N throughout the leaf N range. As a result, Rubisco activation state decreased with increasing leaf N. Both light-saturated CO(2) assimilation at ambient CO(2) and the initial slope of the A:/C:(i) curves were linearly related to initial Rubisco activity, but curvilinearly related to total Rubisco activity. The curvatures in the relationships of both light-saturated CO(2) assimilation at ambient CO(2) and the initial slope of the A:/C:(i) curves with total Rubisco activity were more pronounced than in their relationships with leaf N. This was because the ratio of total Rubisco activity to leaf N increased with increasing leaf N. As leaf N increased, photosynthetic N use efficiency declined with decreasing Rubisco activation state.

Carbon Dioxide↗

The relationship between photosystem II efficiency and quantum yield for CO(2) assimilation is not affected by nitrogen content in apple leaves.

Bench-grafted Fuji/M.26 apple (Malus domestica Borkh.) trees were fertigated with different concentrations of nitrogen by using a modified Hoagland's solution for 45 d. CO(2) assimilation and photosystem II (PSII) quantum efficiency in response to incident photon flux density (PFD) were measured simultaneously in recent fully expanded leaves under low O(2) (2%) and saturated CO(2) (1300 micromol mol(-1)) conditions. A single curvilinear relationship was found between true quantum yield for CO(2) assimilation and PSII quantum efficiency for leaves with a wide range of leaf N content. The relationship was linear up to a quantum yield of approximately 0.05 mol CO(2) mol(-1) quanta. It then became curvilinear with a further rise in quantum yield in response to decreasing PFD. This relationship was subsequently used as a calibration curve to assess the rate of non-cyclic electron transport associated with Rubisco and the partitioning of electron flow between CO(2) assimilation and photorespiration in different N leaves in response to intercellular CO(2) concentration (C(i)) under normal O(2) conditions. Both the rate of non-cyclic electron flow and the rate of electron flow to CO(2) or O(2) increased with increasing leaf N at any given C(i). The percentage of non-cyclic electron flow to CO(2) assimilation, however, remained the same regardless of leaf N content. As C(i) increased, the percentage of non-cyclic electron flow to CO(2) assimilation increased. In conclusion, the relationship between PSII quantum efficiency and quantum yield for CO(2) assimilation and the partitioning of electron flow between CO(2) assimilation and photorespiration are not affected by N content in apple leaves.

Carbon Dioxide↗

Role of neonicotinyl insecticides in Washington apple integrated pest management. Part I. Control of lepidopteran pests.

Three neonicotinyl insecticides, acetamiprid, thiacloprid and clothianidin, were evaluated for their impact on four species of lepidopteran pests of apple in Washington, the codling moth, Cydia pomonella (L.), the Pandemis leafroller, Pandemis pyrusana Kearfott, and the obliquebanded leafroller, Choristoneura rosaceana (Harris), and Lacanobia subjuncta (Grote & Robinson). None of the neonicotinyl insecticides demonstrated sufficient activity against P. pyrusana, C. rosaceana, or L. subjuncta to warrant field trials. Conversely, all had some activity against one or more stages of C. pomonella. Acetamiprid was highly toxic to larvae in laboratory bioassays, and had relatively long activity of field-aged residues (21 days). It also showed some toxicity to C. pomonella eggs (via topical exposure) and adults. Acetamiprid provided the highest level of fruit protection from C. pomonella attack in field trials conducted over five years in experimental orchards with extremely high codling moth pressure. Thiacloprid performed similarly in bioassays, but fruit protection in field trials was slightly lower than acetamiprid. Clothianidin showed moderate to high toxicity in bioassays, depending on the C. pomonella stage tested, but poor fruit protection from attack in field trials. None of the neonicotinyl insecticides were as toxic to larvae or effective in protecting fruit as the current standard organophosphate insecticide used for C. pomonella control, azinphosmethyl. However, both acetamiprid and thiacloprid should provide acceptable levels of C. pomonella control in commercial orchards where densities are much lower than in the experimental orchards used for our trials. The advantages and disadvantages of the neonicotinyl insecticides as replacements for the organophosphate insecticides and their role in a pest management system for Washington apple orchards are discussed.

Agriculture↗

Air method measurements of apple vessel length distributions with improved apparatus and theory.

Studies showing that rootstock dwarfing potential is related to plant hydraulic conductance led to the hypothesis that xylem properties are also related. Vessel length distribution and other properties of apple wood from a series of varieties were measured using the 'air method' in order to test this hypothesis. Apparatus was built to measure and monitor conductivity to air of fresh wood segments of different lengths. Theory for determining vessel length distribution was improved to give a single parameter uni-modal vessel length probability density function. The function, derived by combining the exponential extinction (with extinction coefficient k) of wood conductivity to air (C) as wood length (x) increases (i.e. C=Co exp (-kx)) with the differential double difference formula, is Px=xxk2 exp (-kx), where Px is the fraction of vessels of length x. The main parameter of the distribution, k, was found to be the inverse of the mode of the distribution, i.e. the most common vessel length, Lo. Lo for ten apple rootstock and scion varieties varied from 5.6+/-0.1 cm (+/-SE) for MM.111 to 9.0+/-1.0 for Prunifolia (P <0.05). Average maximum vessel length was approximately 50 cm, and differences were not significant. Effective vessel radii ranged from 14 for Prunifolia to 24.3+/-0.7 micro m for M.26, with standard errors less than 12% of the mean. Specific conductivity of a 15 cm wood segment ranged from 2x109-4) to 1.6+/-0.2x10(-2) dm3 s(-1) kPa(-1) m(-1) for maruba and M.26, respectively, with standard errors up to 63% of the mean. Vessel density at the air entry point ranged from 18+/-3 to 42+/-6 vessels mm-2 for M.26 and MM.106, respectively, with standard errors as high as 89% of the mean. It was concluded that there is no general relationship between the wood properties investigated and rootstock size class, and that plasticity increases from vessel lengths to radii to specific conductivity and vessel densities.

Air↗

Determining the role of N remobilization for growth of apple (Malus domestica Borkh) trees by measuring xylem-sap N flux.

The contribution of N remobilization to the seasonal growth of field-grown Malus domestica (apple) trees was measured using two different techniques. 'Fuji' trees grafted on M.9 apple rootstocks were planted in the field and fertilized and irrigated for two growing seasons. During the second year, the trees received 15N-labelled fertilizer and destructive harvests were taken during the spring and summer to determine the pattern of N remobilization and uptake. At the same time, patterns of N translocation in the xylem were measured by sampling saps at each harvest and analysing them for their constituent amino acids and amides. Total water flux through the trunk xylem was also measured throughout the sampling period using the heat balance technique. The flux of amino compounds in the xylem was then calculated to see if this approach could quantify remobilization. Most of the N for leaf growth was provided by remobilization, which lasted for some 40 d following bud-burst. The labelled N was not taken up until 14 d after remobilization had started. The predominant amino compounds recovered in the xylem were Asn, Asp, Arg, and Gln, whose concentration peaked during remobilization, except for Arg whose concentration was highest at bud-break and declined thereafter. The amount of N translocated in the xylem as Asn, Asp and Gln correlated well with the amount of N remobilized (as measured by the recovery of unlabelled N in the new above-ground growth). The data suggest that Arg is translocated predominantly as a consequence of root uptake and they are discussed in relation to measuring N remobilization in field-grown trees.

Amino Acids↗

Ethylene biosynthesis and perception in apple fruitlet abscission (Malus domestica L. Borck).

Abscission was studied in immature apple fruits (cv. Golden Delicious) during the physiological drop. Fruitlet populations, characterized by different abscission potential, were analysed. Non-abscising fruitlets (NAF) were obtained from central flowers borne in clusters where all the lateral flowers had been removed at bloom while abscising fruitlets (AF) were derived from lateral fruitlets of trees sprayed with benzylaminopurine (BAP) at 200 ppm, 17 d after petal fall (APF), when the fruit cross diameter was about 10-12 mm. Fruit shedding, monitored at the end of the June drop, was significantly different in the two populations, being less than 10%, and more than 90%, in NAF and AF, respectively. In AF, fruit drop peaked around 33 d after petal fall (APF) and was preceded by an increase in ethylene around 20 d APF. Transcript analysis was performed from 17-24 d APF, since preliminary experiments pointed out that major changes in expression of abscission related genes occurred within this period. Transcript accumulation of genes involved in ethylene biosynthesis (MdACS5B and MdACO) and action (MdERS1, MdETR1, and MdCTR1) was studied in the seed, cortex, peduncle, and abscission zone (AZ) of the two fruit populations. MdACS5B and MdACO transcripts accumulated along the experimental period in AF population, even though at a different magnitude, while ethylene evolution declined after peaking at day three. MdETR1, MdERS1, and MdCTR1 expression patterns depended on tissue and/or population. The ERS/ETR ratio was higher in AF than in NAF populations. Overall results pointed out that apple fruitlet drop is preceded by a stimulation of ethylene biosynthesis and a gain in sensitivity to the hormone.

Ethylenes↗

Polyamine biosynthesis of apple callus under salt stress: importance of the arginine decarboxylase pathway in stress response.

To clarify the involvement of the arginine decarboxylase (ADC) pathway in the salt stress response, the polyamine titre, putrescine biosynthetic gene expression, and enzyme activities were investigated in apple [Malus sylvestris (L.) Mill. var. domestica (Borkh.) Mansf.] in vitro callus under salt stress, during recovery after stress, and when ADC was inhibited by D-arginine, an inhibitor of ADC. Salt stress (200 mM NaCl) caused an increase in thiobarbituric acid-reactive substances (TBARS) and electrolyte leakage (EL) of the callus, which was accompanied by an increase in free putrescine content, during 7 d of treatment. Conjugated putrescine was also increased, but this increase was limited to the early stage of salt stress. Accumulation of putrescine was in accordance with induction of ADC activity and expression of the apple ADC gene (MdADC). When callus that had been treated with 200 mM NaCl was transferred to fresh medium with (successive stress) or without (recovery) NaCl, TBARS and EL were significantly reduced in the recovery treatment, indicating promotion of formation of new callus cells, compared with the successive stress treatment. Meanwhile, MdADC expression and ADC activity were also decreased in the callus undergoing recovery, whereas those of the callus under successive stress were increased. Ornithine decarboxylase (ODC) activity showed a pattern opposite to that of ADC in these conditions. D-Arginine treatment led to more serious growth impairment than no treatment under salt stress. In addition, accumulation of putrescine, induction of MdADC, and activation of ADC in D-arginine-treated callus were not comparable with those of the untreated callus. Exogenous addition of putrescine could alleviate salt stress in terms of fresh weight increase and EL. All of these findings indicated that the ADC pathway was tightly involved in the salt stress response. Accumulation of putrescine under salt stress, the possible physiological role of putrescine in alleviating stress damage, and involvement of MdADC and ADC in response to salt stress are discussed.

Arginine↗

Cloning of NAD-dependent sorbitol dehydrogenase from apple fruit and gene expression.

Partial amino acid sequences of NAD-dependent sorbitol dehydrogenase (NAD-SDH) were used to identify a full-length cDNA from apple fruit. This clone consisted of 1,433 bp containing an open reading frame of 1,137 bp that could code for a polypeptide with 379 amino acids. To our knowledge, this is the first report about cloning of NAD-SDH cDNA from a plant source. The deduced amino acids from cDNA revealed 43.7% identity to human NAD-SDH. The activity of this enzyme to convert sorbitol to fructose with the reduction of NAD was certified by the fusion protein of this clone expressed in Escherichia coli. Northern blot analysis showed that the mRNA was expressed in matured apple fruit.

Amino Acid Sequence↗

Sorbitol synthesis in transgenic tobacco with apple cDNA encoding NADP-dependent sorbitol-6-phosphate dehydrogenase.

The apple (Malus domestica) cDNA encoding NADP-dependent sorbitol-6-phosphate dehydrogenase (S6PDH) was stably integrated and expressed in transgenic tobacco (Nicotiana tabacum cv. SR1). Expression of the cDNA in either a sense or antisense orientation was accomplished using cauliflower mosaic virus regulatory sequences (CaMV35S). Sorbitol synthesis was confirmed by gas-chromatography-mass-spectroscopy (GC-MS). Sorbitol concentration in the leaves of the transgenic plants expressing the sense orientation varied from 186 to 446 nmol (g fr wt)-1. The concentration positively correlates with S6PDH activity in leaves. Neither sorbitol nor S6PDH activity was detected in the extracts of nontransformed tobacco or transgenic tobacco expressing the antisense orientation. These results provide key genetic evidence that S6PDH expression is sufficient for the synthesis of sorbitol in tobacco, implicating it as a key enzyme in the sorbitol biosynthetic pathway in apple and perhaps other members of the woody Rosaceae.

Base Sequence↗

Identification of sorbitol transporters expressed in the phloem of apple source leaves.

Sorbitol is a major photosynthetic product and a major phloem-translocated component in Rosaceae (e.g. apple, pear, peach, and cherry). We isolated the three cDNAs, MdSOT3, MdSOT4, and MdSOT5 from apple (Malus domestica) source leaves, which are homologous to plant polyol transporters. Yeasts transformed with the MdSOTs took up sorbitol significantly. MdSOT3- and MdSOT5-dependent sorbitol uptake was strongly inhibited by xylitol and myo-inositol, but not or only weakly by mannitol and dulcitol. Apparent K(m) values of MdSOT3 and MdSOT5 for sorbitol were estimated to be 0.71 mM and 3.2 mM, respectively. The protonophore, carbonyl cyanide m-chlorophenylhydrazone (CCCP), strongly inhibited the sorbitol transport. MdSOT3 was expressed specifically in source leaves, whereas MdSOT4 and MdSOT5 were expressed in source leaves and also in some sink organs. MdSOT4 and MdSOT5 expressions were highest in flowers. Fruits showed no or only weak MdSOT expression. Although MdSOT4 and MdSOT5 were also expressed in immature leaves, MdSOT expressions increased with leaf maturation. In addition, in situ hybridization revealed that all MdSOTs were expressed to high levels in phloem of minor veins in source leaves. These results suggest that these MdSOTs are involved in sorbitol loading in Rosaceae.

Amino Acid Sequence↗

Growth of young apple trees in relation to reserve nitrogen and carbohydrates.

Bench-grafted Fuji/M.26 apple (Malus domestica Borkh.) trees were fertilized with a nutrient solution (fertigation) containing 0, 2.5, 5, 7.5, 10, 15 or 20 mM nitrogen (N) in a modified Hoagland's solution from June 30 to September 1. In mid-October, half of the trees in each N treatment were sprayed twice with 3% urea, 1 week apart. The remaining trees served as controls. All trees were harvested after leaf fall and stored at 2 degrees C over winter. One group of trees from each treatment was destructively sampled before bud break to determine amounts of reserve N and total nonstructural carbohydrates (TNC); the remaining trees were transplanted to N-free medium in the spring. These trees were supplied with Hoagland's solution with or without 10 mM N (from 15N-depleted NH4NO3) for 60 days, starting from bud break. With increasing N supply from fertigation, tree N concentration increased, whereas TNC concentration decreased. Foliar urea applications increased tree N concentration and decreased TNC concentration in each N fertigation treatment. There was a negative linear relationship between tree N concentration and TNC concentration. Irrespective of whether N was provided the following spring, trees with high N reserves but low carbohydrate reserves produced a larger total leaf area at the end of the regrowth period than trees with low N reserves but high carbohydrate reserves. The pooled data on reserve N used for new growth showed that, regardless of the spring N supply, there was a linear relationship between total N accumulated in the tree during the previous season and the amount of reserve N remobilized for new shoot and leaf growth. About 50% of tree N content was remobilized to support new shoot and leaf growth over the range of tree N status examined. We conclude that the initial growth of young apple trees in the spring is determined mainly by reserve N, not reserve carbohydrates. The amount of reserve N remobilized for new growth in spring was proportional to tree N status and was unaffected by current N supply.

Biomass↗