Fate of 15N-labelled nitrogen fertilizer applied to kiwifruit (Actinidia deliciosa) vines

The fate of ¹⁵N-labelled ammonium fertilizer applied once to six-year-old field-grown kiwifruit (Actinidia deliciosa Hayward) vines was measured over three years. The three main treatments were nitrogen (N) applied singularly at 100 or 200 kg N ha^–1 in early spring (two weeks before bud burst) or split with 100 kg N ha^–1 (unlabelled) in early spring and 100 kg N ha^–1 (¹⁵N-labelled) ten weeks later. All N treatments were applied to vines with a history of either 50 or 200 kg N ha^–1 yr^–1. For three years after ¹⁵n application, components of the vines and soil (0–600 mm depth) were sampled at harvest in late autumn and the N and ¹⁵N contents determined.By the first harvest, all plant uptake of ¹⁵N had occurred and this represented 48–53% of the ¹⁵N applied. There was no significant effect of current N fertilizer treatment or of N history on ¹⁵N recovery by vines. Removal of ¹⁵N in harvested fruit was small at 5–6% in the first year and 8% over 3 years. After 2–3 years, most plant ¹⁵N occurred in the roots and this component declined only slowly over time. In contrast, there was a large temporal decline in ¹⁵N in above-ground plant components due to the annual removal in leaf fall and pruning. An associated experiment showed that when ¹⁵N-labelled prunings and leaves were mulched and returned to the soil, only about 9% was recovered by plants within 2 years. Almost all remaining mulched material had been immobilised into the soil organic N.In all treatments, about 20% of the added ¹⁵N remained in soil at the first harvest. This was almost entirely in organic fractions (<0.4% in inorganic N) and mostly in the surface 150-mm layer. The ¹⁵N content in soil changed little over time (from 20 to 17% between the first and third harvests respectively) and indicated that most of the N had been immobilised into stable humus forms.     

Influence of anions on the potassium status and productivity of kiwifruit (Actinidia deliciosa) vines

The effects of K fertiliser (160 kg ha^-1) applied with Cl^- or SO₄ ^2- as the accompanying anion on the K nutrition of kiwifruit (Actinidia deliciosa var. deliciosa) were assessed in a field experiment, using vines with varying degrees of K deficiency. Leaf K concentrations in spring were significantly higher for vines receiving KCl, compared to those receiving K₂SO₄. This effect did not interact significantly with the degree of K deficiency, and persisted for about 6 weeks. Subsequently there was no significant difference between the leaf K concentrations for the vines receiving KCl or K₂SO₄. Applying K as KCl increased the leaf Cl concentration, especially in spring, while applying K as K₂SO₄ had no significant effect on the leaf S concentration at that time. These results implied a greater requirement for organic acid anions for K⁺ uptake from K₂SO₄ than from KCl, and the importance of organic acid anions for K⁺ uptake from different sources of K fertiliser is discussed. This transient effect of the accompanying anion on leaf K status was associated with large effects on flowering, and fruit yields were about 28% higher for plants receiving KCl rather than K₂SO₄.The effects on growth and tissue nutrient composition of varying the concentrations of Cl^-, NO₃ ^-, SO₄ ^2- and H₂PO₄ ^- around the roots of kiwifruit vines were examined in a solution culture experiment. For H₂PO₄ ^-, plant growth was very similar over a wide range of rates of addition. For the other anions, the range between deficiency and toxicity was clearly delineated. For Cl^- and NO₃ ^-, toxicity was associated with high tissue concentrations of Cl and N, respectively, and was consistent with competition for uptake between Cl^- and NO₃ ^-. However, for SO₄ ^2-, toxicity was associated with only a small increase in the tissue S concentration relative to that associated with maximum growth, and appeared to result more from effects on uptake of other anions and cations rather than from direct effects of high tissue S concentrations.It is concluded that the sensitivity of kiwifruit to the anion accompanying K⁺ in fertiliser may be related to the unusually high requirement for Cl previously reported for this species.     

Magnesium deficiency of kiwifruit (Actinidia deliciosa)

Magnesium deficiency was associated with large yield reductions in a five-year-old commercial kiwifruit (Actinidia deliciosa) orchard. The effect on yield resulted primarily from a reduction in fruit numbers, there being no difference in mean fruit weight between fruit harvested from affected and unaffected vines. Magnesium deficiency had no deleterious effect on postharvest storage characteristics of fruit stored at 0.5–1°C for 18 weeks; fruit from deficient vines were firmer but had slightly lower soluble solids than fruit from control vines. Although deficiency symptoms were first observed on the basal leaves of the non-fruiting shoots mid season, indications of the impending deficiency could be established very early in the season using foliar analysis. Magnesium concentrations in youngest fully expanded leaves (YFEL) on the affected vines were less than 2.0 g kg⁻¹ DM four weeks after budbreak and remained below this value for the rest of the season; concentrations in YFEL on unaffected vines did not decrease below this value and gradually increased after fruitset to 4.5 g kg⁻¹ DM at harvest. To avert potential production losses, it is suggested that soluble magnesium fertilizers (containing at least 200 kg ha⁻¹ Mg) should be broadcast early in the season if foliar magnesium concentrations less than 2.0 gkg⁻¹ DM are measured four–six weeks after budbreak.     

Temporal expression of effects of varying nitrogen supply on canopy growth, photosynthesis and fruit production for Actinidia deliciosa vines in the field

The effects of varying nitrogen supply on canopy leaf area, response of leaf net photosynthesis (A_n) to quantum flux density (Q), and fruit yields of kiwifruit vines (Actinidia deliciosa var. deliciosa) were examined in a two-year field experiment. Vines were grown with 0, 250 or 750 kg N ha^?1 year^?1. The responses to nitrogen supply were compared with responses to shade, to examine the impact of reduced carbon assimilation on canopy leaf area and fruit yields. Nitrogen supply did not affect significantly any of the measured variables during the first season of the experiment. In the second season, canopy leaf area was reduced significantly where nitrogen supply was limited. The quantum efficiency of photosynthesis (φ_q) increased from 0. 03 mol CO₂ mol^?1 Q soon after leaf emergence to more than 0. 05 mol CO₂ mol^?1 Q during the middle of the growing season. The quantum saturated rate of A_n (A_sat) also increased during the season, from 7–10 μmol CO₂ m^?2 s^?1 soon after leaf emergence, to 15–20 (μmol CO₂ m^?2 s^?1 during the middle of the growing season. φ_q and A_sat increased significantly with nitrogen supply at all measurement times during the second season. For vines with high nitrogen, fruit yields in both seasons were similar, averaging 3. 05 kg m^?2. Fruit yields in the second season were reduced significantly where nitrogen supply was limited, due to reduced fruit numbers. The relative effects of reduced leaf area and reduced leaf photosynthesis for carbon assimilation by nitrogen deficient vines were examined using a mathematical model of canopy photosynthesis for kiwifruit vines. Simulations of canopy photosynthesis indicated that effects on leaf area and on leaf photosynthesis were of similar importance in the overall effects of nitrogen deficiency on carbon assimilation. The effects of nitrogen supply on fruit numbers (i. e. flower development) preceded the measured effects on carbon assimilation, indicating that the nitrogen supply affected carbon partitioning to reserves in the first season.     

Root demography in kiwifruit (Actinidia deliciosa)

A rhizotron was used to study fine-root demography in mature vines of kiwifruit (Actinidia deliciosa). The vines were grown in a deep, well drained, silt loam and received normal orchard management. Roots were measured from 10 to 160cm depth at biweekly intervals for 2 years. After an initial phase of rapid colonisation of the repacked soil behind the rhizotron windows, the total length of visible roots per vine remained quite steady. This apparent stability of the total belied fast and sustained localized turnover of the fine roots at all soil depths. Fifty-one per cent of the roots survived ≤28d, 69% died at an age ≤56d and only 8% survived >252d. For each year, the cumulative length of roots grown was equivalent to about 2·75 times the maximum net length of roots visible. These may be the largest annual rates of root turnover yet reported. This has important ramifications for the carbon balance, mineral nutrition and water relations of the plant.     

Pollen-pistil interaction in kiwifruit (Actinidia deliciosa; Actinidiaceae)

The pollen-pistil interaction has been examined in kiwifruit (Actinidia deliciosa). In this species a large number of seeds are produced in each fruit and a great many pollen grains germinate and grow to reach the ovules. This growth is assisted by an abundant secretion that is present all along the pistilar tract. At anthesis, the stigma is covered by a secretion where the pollen grains germinate and grow. The stylar transmitting tissue is initially rich in starch reserves, but the starch gradually disappears and, concomitantly, an abundant secretion that stains for carbohydrates appears in all of the intercellular spaces. Pollen tube growth relies on this secretion since it is depleted after pollen tube passage, while in unpollinated flowers it remains unaltered throughout the flower life-span. In the ovary a similar situation occurs. The placental surface, where the pollen tubes grow before reaching the ovules, is covered by a number of obturators. At anthesis, these obturators are rich in starch reserves and have an abundant secretion on their outer surface. As time passes, starch disappears while the secretion increases. It is in this secretion that the pollen tubes grow on their way toward the ovules. These observations are discussed in terms of the support given by the pistil to pollen tube growth to achieve the highly successful reproductive performance of this species.     

Seasonal changes in photosynthetic capacity of leaves of kiwifruit (Actinidia deliciosa) vines

A simplified procedure for the assay and purification of an enzyme which activates a galactosyltransferase (EC 2.4.1.96) involved in volume regulation of the unicellular alga Poterioochromonas malhamensis (Peterfi) is described. The enzyme was extracted with water from membranes, followed by chromatography on DEAE-Sephacel, phenyl-Sepharose and fetuin-agarose. Its proteinase activity was demonstrated by cleavage of oxidized insulin A- and B-Chains. The predominant cleavage site of the oxidized A-chain is the peptide bond between ¹³Leu and ¹⁴Tyr whereas ¹⁶Leu-¹⁷Glu is also hydrolyzed with minor activity. Besides this chymotrypsin-like endopeptidase activity some carboxypeptidase activity was also observed.     

Temporal variability and scaling behaviour in the fine-root system of kiwifruit (Actinidia deliciosa)

We show that temporal variability in root populations can depend upon the scale of measurement (particularly the sampled soil volume). The presence of roots in a range of volumes of soil was studied using perspex tubes installed horizontally into the soil around three mature kiwifruit vines. Roots intercepting lines scored on the tubes were counted using a periscope. For small volumes of soil (c. 2–4 cm³) the root counts varied with time in a very irregular manner, and as the interval between measurements increased the autocorrelation between the measurements decayed rapidly. At about half of the locations monitored there was no significant autocorrelation between measurements 27 d apart. Linear interpolation in these time series was unreliable, and where the correlation dimension could be resolved it was usually non-integer (suggesting chaotic behaviour). The time series measured at different locations were poorly correlated, indicating weak coordination. As the observed soil volume increased, the coordination between locations improved, the autocorrelation function increased, and linear interpolation errors decreased (although these remained substantial). Clearly there are considerable fundamental constraints on our ability to predict the root behaviour of kiwifruit vines at scales that are appropriate for mechanistic models of nutrient and water uptake. We discuss the need for a new conceptual model of the fine-root systems of kiwifruit and similar species.     

The allopolyploid origin of kiwifruit,Actinidia deliciosa (Actinidiaceae)

The genetic origin of kiwifruit (Actinidia deliciosa var.deliciosa) was studied using phylogenetic analysis of DNA sequences derived from the polygalacturonase gene. Results indicate that hexaploid kiwifruit had an allopolyploid origin with the diploidA. chinensis contributing one genome (genome A) and another (as yet unidentified) diploid species contributing a second genome (genome B). The results leave open the question of whether a third, distinct species contributed to the hexaploid kiwifruit genome. A tetraploid race ofA. chinensis is also suggested to be allopolyploid containing genomes A and B.     

Cytokinins and fruit development in the kiwifruit (Actinidia deliciosa). II. Effects of reduced pollination and CPPU application

The significance of changes in cytokinin content during early fruit growth was examined in the kiwifruit ( Actinidia deliciosa var. deliciosa cv. Hayward). Fruit growth was modified by the reduction of seed number or by the application of the synthetic phenylurea cytokinin N -(2-chloro-4-pyridyl)- N -phenylurea (CPPU). The influence of these treatments on cell division was monitored by flow cytometry and changes in the endogenous cytokinins were measured at days 10 and 20 after anthesis, using high-performance liquid chromatography and radioimmunoassay. Total cytokinin levels appeared not to be limiting growth since the highest total cytokinin concentration was detected in unpollinated fruit, which abscised by day 25 after anthesis. However, compared with control fruit which had the highest concentration of zeatin (Z) 10 days post anthesis, Z levels were low in unpollinated fruit. It is hypothesised that an increase in Z is the critical change in cytokinin metabolism required for the initiation of cell division and fruit growth. The synthetic cytokinin CPPU promoted fruit development, but there was a decrease in the endogenous cytokinin concentration. Zeatin was not detected in CPPU-treated fruit. Cell division was reduced in unpollinated fruitlets but there was no significant difference ( P > 0.05) between the other treatments. Differences in final fruit size appeared to be due to cell expansion.     

Anti-hyperglycemic activity of kiwifruit leaf (Actinidia deliciosa) in mice

The methanol extract of kiwifruit leaf suppressed the postprandial blood glucose level after an oral administration of soluble starch or sucrose in mice. The mechanism of action is proposed to be due to the alpha-amylase-inhibiting activity in the 90% aqueous methanol fraction and alpha-glucosidase-inhibiting activity in the n-buthanol fraction, based on the results of in vitro experiments.     

Purification and characterisation of a galactoglucomannan from kiwifruit (Actinidia deliciosa)

A galactoglucomannan (GGM) has been purified from the primary cell walls of ripe kiwifruit. A combination of barium hydroxide precipitation, anion exchange- and gel-permeation chromatography gave a chemically homogeneous polymer with a 1:2:2 galactose-glucose-mannose ratio and a molecular weight range of 16-42 kDa. Complete hydrolysis of the polymer with endo-1,4-beta-mannanase (EC 3.2.1.78) from Aspergillus niger gave a mixture of oligosaccharides, three of which (II, III, IV) accounted for more than 80% of the GGM. Structural characterisation of these oligosaccharides and the original polysaccharide was achieved by linkage analysis, 1D and 2D NMR spectrometry and enzymatic hydrolysis. Oligosaccharide II beta-D-Glcp-(1-->4)-beta-D-Manp-(1-->, III beta-D-Glcp-(1-->4)-[alpha-D-Galp-(1-->6)]-beta-D-Manp-(1-->, and IV beta-D-Glcp-(1-->4)-[beta-D-Galp-(1-->2)-alpha-D-Galp-(1-->6)]-beta-D-Manp-(1-->4)-beta-D-Glcp-(1-->4)-beta-D-Manp-(1-->, appeared in the molar ratio of 2:1:1. A trace amount of mannobiose (I) was detected, indicating that some of the mannosyl residues were contiguous. It is concluded that the predominant structural feature of kiwifruit GGM is a backbone of alternating beta-(1-->4)-linked D-glucopyranosyl and D-mannopyranosyl residues, with approximately one third of the latter carrying side-chains at 0-6 of single alpha-D-Galp-(1--> residues (50% of the branches) or the disaccharide beta-D-Galp-(1-->2)-alpha-D-Galp-(1--> (50% of the branches), the substituted residues being separated by three or five unsubstituted monosaccharide units.     

Solar light effects on growth,net photosynthesis,and leaf morphology of in vitro kiwifruit (Actinidia deliciosa) CV hayward

Summary Proliferating axillary shoots of kiwifruit (Actinidia deliciosa A. Chev., C. F. Liang and A. R. Ferguson), var.deliciosa, cv. ‘Hayward’ were grown under solar (SL), white (WL), and blue (BL) light regimens to determine the accumulation of fresh and dry weight, proliferation rate, shoot growth (length), and the net leaf photosynthetic capacity at the CO₂ concentration ranges of 200 to 350, 400 to 600, and 1200 to 1500 ppm. An histologic study determined the effects of light source on leaf stomatal density and tissue morphology. Dry and fresh matter accumulation was greatest, but callus development most limited under the SL regimen. Shoot proliferation was highest under WL and length under BL. Net photosynthetic capacity was highest for leaves grown under SL and lowest for those under BL; the leaves exposed to the latter regimen were also thinner and exhibited a less compact arrangement of palisade cells than those under WL and SL. Leaf stomata density was highest under the BL source.     

Stem and whole-plant hydraulics in olive (Olea europaea) and kiwifruit (Actinidia deliciosa)

A field study and an experiment under controlled conditions using pressure-flux relationships were conducted to compare the stem and whole-plant conductance in olive (Olea europaea) and kiwifruit (Actinidia deliciosa) species. Anatomical observations were also made on one-year-old stem to determine the conductive area of vessels (A _ves) and the total xylem area (A _xyl). Results show that A _ves of kiwifruit twigs was ~2.5-fold of that in olive twigs, and the hydraulically weighted mean diameter was up to threefold that of the olive ones. One-year-old olive twigs had lower hydraulic conductivity (k) than the kiwifruit, while values of leaf-specific conductivity (i.e. k normalised per unit leaf area) were higher than the kiwifruit (i.e. ~49 and 29 × 10^?6 kg m^?1 s^?1 MPa^?1, respectively). In the field experiment, the flux of sap (heat balance method) and differences in water potential through the soil–plant system (ΔP) were used for both species to calculate the whole-plant conductance that was normalised per unit leaf area (leaf-specific whole-plant conductance, K _plant,LA). Values of K _plant,LA are attributable to the combined effect of the ΔP and anatomical features of conduits. Olive species showed a larger ΔP (2.4 MPa at midday) than the kiwifruit (0.5 MPa) which contributed to lower K _plant,LA in Olea than the Actinidia plants. This information, combined with vessel density data, contributes to explain differences amidst olive and kiwifruit species, in terms of susceptibility to some drought-related hydraulic impairments induced by the Mediterranean environment.     

Cloning and characterization of five cDNAs for genes differentially expressed during fruit development of kiwifruit (Actinidia deliciosa var.deliciosa)

Five cDNAs for genes differentially expressed during fruit development of kiwifruit (Actinidia deliciosa var.deliciosa cv. Hayward) were isolated from a library made from young fruit, 8–10 days after anthesis. One gene (pKIWI503) has low levels of expression in young fruit but is induced late in fruit development and during fruit ripening, and has some homology to plant metallothionein-like proteins. The other four genes are highly expressed in young fruit with reduced expression in the later stages of fruit development. pKIWI504 has strong homology to plant metallothionein-like proteins and pKIWI505 exhibits homology to the -subunit of the mitochondrial ATP synthase gene. The two other genes (pKIWI501 and 502) encode proteins with no significant homology to other known sequences.     

Synchronising diagnosis and correction of nitrogen deficiency in barley grown in semi-arid environments

In field experiments conducted from 1986 to 1989, nitrogen (N) deficiency in barley crops was corrected with the same efficiency by applying N fertilizer at sowing or by broadcasting it at early tillering. Four plant tests for diagnosing and predicting N deficiency in barley were evaluated. Of these the basal stem nitrate-N test was best with a calibrated critical range of 12 000 to 15 000 mg kg^-1 when sampling was during early tillering.     

Cytokinins and fruit development in the kiwifruit (Actinidia deliciosa). I. Changes during fruit development

The cytokinin content in fruit tissue of the kiwifruit ( Actinidia deliciosa [A. Chev.] C. F. Liang et A. R. Ferguson var. deliciosa cv. Hayward) was monitored during fruit development to identify which cytokinins were present and if they were linked with specific stages of fruit growth. Cytokinins were isolated and purified by column chromatography and high-performance liquid chromatography and quantified by radioimmunoassay. A novel HPLC step utilising an amine column was successfully introduced as a preparative step in the separation of the O - and 9-glucosides from the free bases and ribosides. The radioimmunoassay results were validated, and the different cytokinins identified, by gas chromatography-mass spectrometry. Cytokinins detected in fruit included the cytokinin free bases, zeatin and isopentenyladenine, their ribosides, nucleotides and both O - and 9-glucosides. Both qualitative and quantitative changes of the cytokinins occurred during fruit development. A decrease in cytokinin concentration occurred after anthesis (from 342 pmol g⁻¹ fresh weight at anthesis to 41 pmol g⁻¹ fresh weight 27 days after anthesis). A large increase in cytokinin concentration and content per fruit occurred as the fruit reached commercial maturity (to 1900 pmol g⁻¹ fresh weight). Individual cytokinins showed quite different patterns. Zeatin, in particular, showed a peak in concentration (13 pmol g⁻¹ fresh weight) 11 days after anthesis that correlated with the beginning of the cell division phase of fruit growth. The accumulation of cytokinin (mostly zeatin riboside or zeatin nucleotide) in mature fruit may be of significance for the postharvest storage of kiwifruit fruit.     

Papillar integrity as an indicator of stigmatic receptivity in kiwifruit (Actinidia deliciosa)

Stigmatic receptivity is a major factor limiting fruit set inkiwifruit (Actinidia deliciosa). The aim of this work was toknow what determines the cessation of stigmatic receptivityin this species. Stigmatic receptivity has been analysed inkiwifruit through the capacity of the stigma to sustain pollengermination and has been related to stigmatic development anddegeneration.The stigma of kiwifruit flowers has a papillatesurface which from anthesis is covered with an abundant exudate.Papillae are unicellular and contain a number of phenolic deposits.During the lifetime of the flower these papillae gradually loseturgidity, while stigmatic secretion increases until 5 d afteranthesis when papillae start rupturing and the papillar contentis liberated into the germination medium. Stigmatic receptivityis high at anthesis and lasts for the next 4 d. However, itdrastically decreases 5 d after anthesis and it is nil 2 d later.The pattern of stigmatic receptivity closely fits that of papillarintegrity, indicating that stigmatic receptivity relies on thisintegrity. Since papillar integrity can easily be evaluatedin stigmatic arms, this criterion can be used as a quick methodto estimate stigmatic receptivity. Key words: Actinidia deliciosa, kiwifruit, stigma, receptivity, pollen     

Somatic embryogenesis in leaves and leaf-derived protoplasts of Actinidia deliciosa var. deliciosa cv. Hayward (kiwifruit)

Summary The obtention of embryogenic competence in Actinidia deliciosa var. deliciosa cv. Hayward is reported. Axillary buds from shoots submitted to cold (4°C) and starvation for 1.5 months, developed leaves with embryogenic competence. These leaves, cultured in darkness for 1.5 months on a medium containing zeatin as a sole growth regulator, originated compact structures from which embryos developed. The plating orientation and sectioning of leaves strongly affected the expression of the embryogenic potential. A selected fraction of the protoplasts isolated from these leaves was able to develop in an embryogenic way. The germination of the embryos is still only occasional.Abbreviations 2,4-D 2,4-dichlorophenoxyacetic acid - 2-iP 6-dimethylallyl aminopurine - IAA indole-3-acetic acid - NOA naphthoxyacetic acid - SEM Scanning Electron Microscopy