Abscission of mango fruitlets as influenced by enhanced ethylene biosynthesis

Experiments were conducted on developing fruitlet explants of two mango (Mangifera indica L.) cultivars to establish the source and dynamics of ethylene production prior to and during fruitlet abscission. Abscission of all fruits in the samples occurred at approximately 86 and 74 hours postharvest in `Keitt' and `Tommy Atkins,' respectively. Increased abscission began 26 hours from harvest and was preceded by enhanced ethylene synthesis. Enhanced ethylene production initiated approximately 48 hours prior to abscission and increased to a maximum near the time of fruitlet abscission. The seed produced the highest amount of ethylene on a per gram fresh weight basis. The pericarp, however, was the main source of ethylene on an absolute basis, since it represented more than 85% of total fruitlet weight. Pedicels containing the abscission zone produced no detectable ethylene prior to or at the moment of abscission. Fumigation of `Tommy Atkins' fruitlets with 1, 15, or 100 microliters per liter ethylene accelerated abscission by 24 to 36 hours in comparison with unfumigated controls. Diffusion of ethylene from distal fruitlet tissues to the abscission zone triggers the events leading to separation of the fruit from the tree.     

Carbohydrate and ethylene levels related to fruitlet drop through abscission zone A in citrus

Citrus fruits have two abscission zones (AZ), named A (in the pedicel) and C (in the calyx). Early fruitlet abscission takes place exclusively through AZ A, while at June drop it is progressively inactivated and AZ C begins to operate. In previous work, it has been demonstrated that carbohydrate and ethylene regulate fruit drop through abscission zone C. In this paper, we have analysed the effect of these two factors in developing fruitlets of Satsuma mandarins (Citrus unshiu [Mak.] Marc.) cv. Okitsu to elucidate their involvement on abscission through AZ A. The data indicated that ACC content and ethylene production of fruitlets paralleled abscission rates. Sucrose supplementation increased fruit set, although did not counteract the abscising effect induced by ACC. Branch girdling of terminal fruitlets carrying several leaves significatively reduced ethylene production and abscission rates, and increased sugar content. Pedicel girdling showed the opposite. Taken together, the results revealed that the carbohydrate content may be a biochemical signal involved in the mechanisms controlling abscission through AZ A. The evidence also showed parallelisms between ethylene and its activation. As the induction of higher ethylene levels after the period of AZ A activity, however, was not able to promote fruit drop, it is also concluded that solely ethylene is not sufficient to activate abscission.     

Effects of nitrogen supply on growth, contents of phenolic compounds and pathogen (scab) resistance of apple trees

The effect of long-term N-supply on growth, scab resistance and phenolic compounds in the leaves of two apple cultivars was studied. The different pools of phenylpropanoids (hydroxycinnamic acids, dihydrochalcones) and flavonoids (flavonols, catechins, procyanidins) were quanitfied by HPLC from non-infected and inoculated leaves representing different ontogenetic stages. Scab incidence was also evaluated. Strictly following the carbon-nutrient-balance hypothesis, apple trees responded to high N-supply with increased shoot growth and with a reduced accumulation of total phenolic compounds in their leaves. This was shown for the cultivar 'Golden Delicious', which is susceptible to the scab disease caused by Venturia inaequalis , and for the resistant cultivar 'Rewena'. Whereas high N-fertilization increased the susceptibility of 'Golden Delicious', it did not decrease the resistance of 'Rewena' despite of the pronounced reduction of phenolic concentrations. Thus, a simple C trade off between growth-related metabolism and secondary metabolism cannot solely explain changes in defensive potential.     

Carbohydrate Availability in Relation to Fruitlet Abscission in Citrus

Abscission of flowers and fruitlets in the Washington navelorange (Citrus sinensis [L.] Osbeck) has been characterizedin relation to carbohydrate availability. A main wave of flowerabscission occurs shortly after anthesis while the carbohydratereserves in the tree are high. Fruitlet abscission starts approx.30 d after the commencement of flowering, while carbohydrates(mainly starch) are being accumulated in the leaves. Flowerand early fruitlet abscission are not caused by carbohydrateshortage. During late fruitlet abscission sucrose concentrationin the leaves falls to a low value demonstrating a limitationin supply and competition among the developing fruitlets forcarbohydrates. Concentrations of sucrose and reducing sugarsin the peel of the fruitlets also fall to low values, and arelationship could be demonstrated between these free sugarlevels and abscission. Ringing increases carbohydrate supplyto fruit and reduces late fruitlet abscission, but only hasa marginal effect on the growth of the fruitlets, which seemsless sensitive than abscission to carbohydrate shortage. Thelimitation of carbohydrate supply to the fruitlets occurs whilestarch levels in the leaves remain high. Slow mobilization ofstarch reserves may be one factor limiting set in Citrus. Copyright2001 Annals of Botany Company Carbohydrate supply, citrus, fruit growth and abscission, ringing, navel orange, starch, sugar metabolism     

Hormonal regulation of fruitlet abscission induced by carbohydrate shortage in citrus

 The hormonal signals controlling fruitlet abscission induced by sugar shortage in citrus were identified in Satsuma mandarin, Citrus unshiu (Mak.) Marc, cv. Clausellina and cv. Okitsu. Sugar supply, hormonal responses and fruitlet abscission were manipulated through full, partial or selective leaf removals at anthesis and thereafter. In developing fruitlets, defoliations reduced soluble sugars (up to 98%), but did not induce nitrogen and water deficiencies. Defoliation-induced abscission was preceded by rises (up to 20-fold) in the levels of abscisic acid (ABA) and 1-aminocyclopropane-1-carboxylic acid (ACC) in fruitlets. Applications to defoliated plants showed that ABA increased ACC levels (2-fold) and accelerated fruitlet abscission, whereas norflurazon and 2-aminoethoxyvinyl glycine reduced ACC (up to 65%) and fruitlet abscission (up to 40%). Only the full defoliation treatment reduced endogenous gibberellin A₁ (4-fold), whereas exogenous gibberellins had no effect on abscission. The data indicate that fruitlet abscission induced by carbon shortage in citrus is regulated by ABA and ACC originating in the fruits, while gibberellins are apparently implicated in the maintenance of growth. In this system, ABA may act as a sensor of the intensity of the nutrient shortage that modulates the levels of ACC and ethylene, the activator of abscission. This proposal identifies ABA and ACC as components of the self-regulatory mechanism that adjusts fruit load to carbon supply, and offers a physiological basis for the photoassimilate competition-induced abscission occurring under natural conditions. Received: 19 February 1999 / Accepted: 14 August 1999     

Biosynthesis and Contents of Gibberellins in Seeded and Seedless Sweet Orange (<Emphasis Type="Italic">Citrus sinensis</Emphasis> L. Osbeck) Cultivars

In this work, we study the capacity to biosynthesize gibberellins (GA) of ovules (either fertilised or unfertilised), developing seeds and pericarp from fruitlets and their relation with fruit set capacity. Experiments were performed in adult, 12-year-old trees of seeded (Pineapple) and seedless parthenocarpic (Washington navel) sweet orange [Citrus sinensis L. Osbeck] cultivars. The activity of GA20-, GA3- and GA2-oxidases and gibberellin levels were measured in the ovules and pericarp of fruitlets in different development states. The results indicate that ovules are the main sites of gibberellin synthesis in fruitlets during the post-anthesis period. The most intense GA₁ synthesis—coincident with the highest expression of GA20ox2, GA3ox1 and GA2ox1—was detected in the ovules of the seeded cultivar, probably induced by fecundation and associated with low early fruitlet abscission rates. By contrast, the low activity detected in the sterile cultivar appears to be rather developmentally or constitutively regulated. As a fruitlet develops, the GA₁ concentration is augmented in the pericarp in comparison to ovules or developing seeds, and levels therein did not exhibit noticeable differences between varieties. Furthermore, developing seeds from pineapple had higher GA₁ content than the unfertilised abortive ovules from Washington navel. Taken together, data suggest a main role for this hormone in the control of fruitlet abscission, and also demonstrate a function in seed development.     

Synthetic Auxin 3,5,6-TPA Provokes <Emphasis Type="Italic">Citrus clementina</Emphasis> (Hort. <Emphasis Type="Italic">ex</Emphasis> Tan) Fruitlet Abscission by Reducing Photosynthate Availability

The aim of this study was to determine the effects of the synthetic auxin 3,5,6-trichloro-2-pirydiloxyacetic acid (3,5,6-TPA) on photosynthetic activity, photosynthate transport to the fruit, and fruitlet abscission to further explain the physiological basis of auxin-mediated citrus fruit thinning. Applying 15 mg l⁻¹ 3,5,6-TPA to trees during the fruit cell division stage significantly increased fruitlet abscission of Clementine mandarin. On treated trees, abnormal foliar development and photosynthetic damage were observed at the same time as 3,5,6-TPA reduced fruitlet growth rate. Briefly, treatment reduced chlorophyll and carotenoid concentrations and modified chlorophyll a fluorescence parameters, that is, reduced the quantum yield (ФPSII) of the noncyclic electron transport rate, diminished the capacity to reduce the quinone pool (photochemical quenching; q_p), and increased nonphotochemical quenching (q _N), thereby preventing the dissipation of excess excitation energy. In addition, the net photosynthetic flux (μmol CO₂ m⁻² s⁻¹) and leaf photosynthate content decreased in treated trees. As a result, the 3,5,6-TPA treatment significantly reduced the photosynthate accumulation in fruit from day 3 to day 8 after treatment, thus reducing fruitlet growth rate. Hence, treated fruitlets significantly increased ethylene production and abscised. Twenty days after treatment, chlorophyll a fluorescence parameters and fruitlet growth rate were reestablished. Accordingly, the thinning effect of 3,5,6-TPA may be due to a temporarily induced photosynthetic disorder that leads to reduction in photosynthate production and fruitlet uptake that temporarily slows its growth, triggering ethylene production and fruitlet abscission. Afterward, the remaining treated fruit overcame this effect, increased growth rate, and reached a larger size than control fruit.     

Effects of cropping, nutrition and water supply on accumulation and distribution of biomass and nutrients for apple trees on'M9'root systems

The effects of cropping, nutrition and water supply on accumulation and distribution of biomass and nutrients for apple trees on'M9'root systems were examined using trees growing in lysimeters. For 3-year-old 'Golden Delicious' trees, cropping significantly increased total biomass, although leaf and root biomass were reduced. For 3-year-old'Golden Delicious', 'Coxs Orange' and 'Gloster' trees, cropping only in the third year reduced root biomass by 39 to 45%. Cropping did not affect water use by trees of any cultivar early in the season, but increased water use later in the season for 'Gloster' trees. For trees with fruit, total non-structural carbohydrate (TNC) contents at the end of the season were highest in fruit (up to 1 400 g plant ⁻¹). For trees without fruit, TNC contents were generally highest in the roots (up to 110 g planr ⁻¹). Supplying trees with nutrient solution diluted to 10% of the standard composition increased the root biomass for 'Golden Delicious' trees only, but decreased biomass of above-ground tissues for all cultivars. Reducing the nutrient supply decreased water and nutrient uptake, and reduced nutrient contents within vegetative parts of the tree more than within roots. For 'Golden Delicious' trees, restricting the water supply to 50% or 25% of that consumed by control trees significantly reduced above-ground biomass, but root biomass was not significantly affected. The N, P and K. contents for the trees were also reduced by water stress, due to reduced contents within above-ground organs. Water stress reduced the TNC and starch contents of all tissues, except the roots. These results are discussed in relation to the efficiency of root systems for water and nutrient uptake.     

Regulators of Cell Division in Plant Tissues VUI. The Cytokinins of the Apple Fruit

The cytokinin activities of extracts of organs developed from the apple fruit bud were compared using the carrot phloem bioassay, and the identity of the cytokinins in the apple fruitlet was investigated. The activity of apple fruitlet extracts was slightly greater than the activity of pedicel extracts, and considerably greater than the activities of extracts of other organs. Extracts of the developing seeds of fruitlets were much more active than extracts of fruitlet flesh. Apple fruitlet extracts contained three principal cytokinins. One was identified as a 6-(substituted amino)purine and was either zeatin or some very closely related compound. The two other cytokinins exhibited the chromatographic behaviour of zeatin riboside and zeatin ribotide. A cytokinin extracted from vegetative apple shoots was chromatographically indistinguishable from zeatin.     

Influence of timing and prey availability on fruit damage to apple by Campylomma verbasci (Hemiptera: Miridae)

Campylomma verbasci Meyer is a zoophytophagous mirid that feeds on small arthropods as well as apple (Malus domestica Borkhausen) fruits, causing economic damage to some cultivars. The influence of timing and prey availability on the amount of fruit damage was studied to determine whether either factor could be used to refine a management program. C. verbasci nymphs were caged on branches of fruiting 'Golden Delicious' apple trees during the period from bloom through early fruit set. The greatest amount of fruit damage occurred during the bloom period; little or no damage occurred after fruit reached approximately equals 13 mm in diameter. The availability of prey did not reduce the incidence of fruit damage by C. verbasci, nor did it influence the survival of nymphs. Nymphal survival was higher, however, in cages where a blossom or fruitlet was present versus a vegetative spur. These data support the hypothesis that post petal fall insecticide applications (those made after the fruit is greater than approximately equals 10-13 mm in diameter) are not useful in preventing economic levels of fruit damage in Washington State, and that petal fall applications would only prevent a fraction of the total amount of damage by this pest. The data from this study do not support the hypothesis that manipulating arthropod prey species of C. verbasci will prevent fruit damage. There was evidence to support the hypothesis that nymphs can survive a relatively short period (7 d) without arthropod prey.     

Architecture of the pruned tree: impact of contrasted pruning procedures over 2 years on shoot demography and spatial distribution of leaf area in apple (Malus domestica)

BACKGROUND AND AIMS: Demography and spatial distribution of shoots are rarely studied on pruned trees. The present 2-year study deals with the effect of pruning strategies on shoot demography and development, and consequences on the spatial distribution of leaf area in three architecturally contrasted - from type II to IV - apple cultivars: 'Scarletspur Delicious', 'Golden Delicious' and 'Granny Smith'. METHODS: All trees were initially subjected during 5 years to Central Leader training with winter heading on all long shoots. For 2 years, half of the trees were further trained with Centrifugal training, where removal of flowering shoots - called extinction pruning - was carried out along the trunk and at the bottom of branches at flowering time. During these 2 years, shoot type (vegetative, inflorescence) and length, and the three-dimensional spatial distribution of all shoots were assessed with an electromagnetic digitizer. KEY RESULTS: Shoot demography, frequency of transitions toward an inflorescence from either an inflorescence (bourse-over-bourse) or a vegetative shoot (trend toward flowering), and the number of bourse-shoots per bourse were strongly affected by cultivar, with little influence of tree manipulation. In contrast, the proportion of vegetative long shoots developing from previous year latent buds was significantly lower in Centrifugal-trained trees for the three cultivars. Canopy volume showed large variations between cultivars, but only that of 'Granny Smith' was affected by tree manipulation in the 2 years. Spatial distribution of shoots varied significantly according to cultivar and manipulation. In 'Scarletspur Delicious' and, to a lesser extent 'Golden Delicious', the distribution of vegetative and flowering shoots in the outer and the inner parts, respectively, was not affected by tree manipulation. In contrast, in 'Granny Smith', vegetative shoots were stimulated in the periphery of Central Leader trees, whereas flowering shoots were stimulated in the periphery of Centrifugal-trained trees. CONCLUSIONS: In apple, the variability of responses to contrasted pruning strategies partly depends on the genetically determined growth and flowering habit of the cultivar.     

Changes in the accumulation of flavonoid and isoflavonoid conjugates associated with plant age and nodulation in alfalfa (Medicago sativa)

Carbohydrate and mineral nutrition was studied in relation to abscission in fruitlets from leafy inflorescences of the Washington navel orange ( Citrus sinensis [L.] Osbeck). Differences in the growth rate of the fruitlets permitted to predict abscission several weeks in advance. This allowed characterization of early differences in composition and behaviour of persisting and abscising fruitlets.
Inflorescences with persisting fruitlets accumulated more mineral elements than inflorescences with abscising fruitlets, and for the phloem-mobile elements the excess accumulation was allocated to the fruitlets. Starch accumulated in the inflorescence leaves during early fruitlet growth, and this accumulation was enhanced by the persisting fruitlets despite their higher growth rate and mobilizing ability. The relations between the fruitlets and the inflorescence leaves cannot be explained totally in terms of source sink relationships; a hormonal regulation of the leaves by the fruitlets is postulated.
Acid invertase activities and hexose concentration in the pericarp were higher in the abscising fruitlets. The lower early growth rate of these fruitlets is thus not caused by a limitation in carbohydrate supply. It seems more related to carbohydrate utilization, probably hormonally mediated, as demonstrated by the higher dependence on hormone supply for the growth in vitro of the endocarp explants.