Delay of early fruitlet abscission by branch girdling in citrus coincides with previous increases in carbohydrate and gibberellin concentrations

Current evidence in citrus indicates that gibberellins (GAs) are main determinants of early fruit set while subsequent growth of developing fruits is mostly dependent upon carbohydrate availability. In this work, branch girdling performed at anthesis in Satsuma mandarin (Citrus unshiu (Mak.) Marc.) cv. Okitsu transitorily reduced early abscission rates (12–32 days after anthesis, DAA) delaying initially the process of natural fruitlet drop. The effects of girdling on growth, gibberellin (GA) and carbohydrate concentrations in developing ovaries and fruitlets were assessed during this initial growth stage (0–69 DAA). In girdled branches, abscission rate reduction was preceded by elevated concentrations of carbohydrate and GA in developing ovaries and fruitlets. Girdling at anthesis stimulated higher hexose (21 DAA) and starch (6–20 DAA) concentrations and also higher GA₁ (6 DAA), GA₁₉ (13–20 DAA) and GA₂₀ (6–20 DAA). The results established a relationship between the reduction of early abscission rates and higher concentrations of carbohydrates and GAs induced by girdling in developing fruitlets. These findings revealed that girdling certainly increased GA concentration and strongly suggested that its effect on early fruitlet abscission delay is likely mediated by both GA and carbohydrates.     

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.     

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.     

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     

Dry matter accumulation in citrus fruit is not limited by transport capacity of the pedicel

The vascularization of the pedicel in Marisol clementine (Citrus clementina Hort. ex Tanaka) has been characterized in relation to fruit growth. Phloem and xylem formation occurred during the first half of the period of fruit growth. Phloem cross-sectional area reached its maximum value by the end of fruitlet abscission, 78 d after anthesis (DAA), shortly after the rate of accumulation of dry matter in fruitlets reached its maximum value. Secondary xylem formation occurred until day 93, well after the end of fruitlet abscission. At fruit maturity, xylem accounted for 42-46 % of the cross-section of the pedicel. Vessels differentiated in this late-formed xylem. Formation of phloem and early xylem was directly related to fruitlet size (and growth rate). Differences in the rate of formation of conductive tissues in the pedicel of the developing fruitlets followed rather than preceded the differences in growth rate. Specific mass transfer (SMT) in the phloem was highest in the fastest growing fruitlets, and peaked during the late stages of fruitlet abscission (72-78 DAA) and during the main period of fruit growth (107-121 DAA). Application of a synthetic auxin to developing fruits, either at the end of flowering (2,4-D) or by day 64 after flowering (2,4-DP), increased the growth rate of the fruit and fruit size at maturity (8-13 % increase in fruit diameter at maturity). These auxin applications also enhanced the formation of conductive tissues in the pedicel, with a specific effect on phloem formation. Applying auxin at flowering resulted in a reduction in the phloem SMT by days 72-78, whereas auxin application on day 64 increased this parameter. Despite this difference in behaviour, which resulted from the different time-course of the growth response of the fruit to auxin applications, these applications increased fruit size to a similar extent. Severing 37 % of the phloem of the pedicel during the main period of fruit growth resulted in an increase in the specific mass transfer in the phloem but had no influence on fruit growth. These observations demonstrate that the transport capacity in the phloem of the pedicel does not limit fruit growth and, within the limits of our experiments, an increase in demand by the fruit appeared to be matched by an increase in SMT. The dependence of late xylem formation (after the period of fruitlet abscission) on fruitlet growth was demonstrated in Salustiana orange [Citrus sinensis (L.) Osbeck] by means of controlling fruit growth through the manipulation of leaf area. Fruit growth at this time was more closely related to leaf area than to carbohydrate levels, suggesting that it may be limited by current photosynthesis.     

Pollination and Resource Constraints on Fruit Set and Fruit Size ofPersoonia rigida(Proteaceae)

Flower and immature fruit abscission can be caused by proximatefactors preventing the development of all flowers into maturefruits. Two potential limiting factors, pollen supply and carbohydrateavailability, were assessed as relative constraints on accumulationof fruit dry matter inPersoonia rigida(Proteaceae). Naturalpollen transfer was highly efficient, and no evidence of self-incompatibilitywas found. Levels of fruit set (67.4% following open-pollinationand 50.4–58.8% following hand-pollinations) may be thehighest recorded for a species of Proteaceae with hermaphroditicflowers. Fruit abscission occurred principally in a single phasefrom 4–10 weeks post-anthesis, with a peak during theeighth week post-anthesis, and this early abscission was theresult of fertilization failure in some flowers. Manipulationof carbohydrate availability by girdling and/or defoliationof branches provided evidence that carbohydrate supply affectedfruit set only when supply was very low. Fruit abscission dueto very low carbohydrate availability on defoliated brancheswas evident during the early abscission phase but was more pronouncedduring the second half of the fruit development period (>17weeks post-anthesis), coinciding with the period of greatestdry matter demand by the fruits. Minimal fruit abscission occurredduring this later period on leaf-bearing branches. Increasesin carbohydrate supply had no effect on fruit set, but fruitsize was highly sensitive to carbohydrate availability. Subtendingand adjacent leaves were identified as major contributors tothe carbohydrate supply of fruits, although some carbohydratecould be obtained from beyond the fruiting branch. Despite thehigh levels of natural fruit set, both pollen supply and resourceavailability were indicated as potential constraints on accumulationof fruit dry matter inP. rigida.Copyright 1999 Annals of BotanyCompany. Persoonia rigidaR.Br., Proteaceae, pollination, carbohydrates, fruit set, fruit size, abscission, girdling.     

Partitioning of [14C]sucrose and Acid Invertase Activity in Reproductive Organs of Pepper Plants in Relation to Their Abscission Under Heat Stress

The effect of heat stress on processes related to carbohydratepartitioning was investigated in young bell pepper (Capsicumannum L. cv. Maor) plants in relation to abscission of theirreproductive organs at different stages of development. None of the reproductive organs abscised after 5 d in a normalday/night temperature regime (25/18°C). With a temperatureregime of 35°C day, 25°C night, abscission occurredin only a small portion of the flower buds and none of the flowersand fruitlets. However, when temperatures in the day and nightwere reversed (25/35°C, day/night) all the buds and someof the flowers abscised during that time period. The young fruitat the first node did not abscise under any temperature regime.The abscission rate of the flower buds was reduced under heatstress if the developing fruit at the first node had been removed. High temperature during either the light or dark periods reducedthe export of [¹⁴C]sucrose from the source leaf (fed for 48h with [¹⁴C]sucrose). Both heat stress and fruit presence reduced the relative amountof [¹⁴C]sucrose which was exported to the flower buds, flowersand roots. Likewise, these treatments reduced the concentrationof reducing sugars in the reproductive organs. Concomitantly,the heat stress and fruit presence on the first node reducedthe activity of soluble acid invertase in the flower buds andthe roots, but not in young leaves. Overall, the results show that heat stress causes alternationin sucrose distribution in the plant, but may also have specificeffects on metabolic activities related to sucrose import andutilization in flower buds and flowers which in turn may enhancetheir abscission. Bell pepper, (Capsicum annuum L. cv. Maor), abscission, acidinvertase, heat stress, reproductive organs, sink leaves. Bell pepper, (Capsicum annuum L. cv. Maor), abscission, acid invertase, heat stress, reproductive organs, sink leaves.     

Partitioning of [14C]sucrose and Acid Invertase Activity in Reproductive Organs of Pepper Plants in Relation to Their Abscission Under Heat Stress

The effect of heat stress on processes related to carbohydratepartitioning was investigated in young bell pepper (Capsicumannum L. cv. Maor) plants in relation to abscission of theirreproductive organs at different stages of development. None of the reproductive organs abscised after 5 d in a normalday/night temperature regime (25/18 °C). With a temperatureregime of 35 °C day, 25 °C night, abscission occurredin only a small portion of the flower buds and none of the flowersand fruitlets. However, when temperatures in the day and nightwere reversed (25/35 °C, day/night) all the buds and someof the flowers abscised during that time period. The young fruitat the first node did not abscise under any temperature regime.The abscission rate of the flower buds was reduced under heatstress if the developing fruit at the first node had been removed. High temperature during either the light or dark periods reducedthe export of [¹⁴C]sucrose from the source leaf (fed for 48h with [¹⁴C]sucrose). Both heat stress and fruit presence reduced the relative amountof [¹⁴C]sucrose which was exported to the flower buds, flowersand roots. Likewise, these treatments reduced the concentrationof reducing sugars in the reproductive organs. Concomitantly,the heat stress and fruit presence on the first node reducedthe activity of soluble acid invertase in the flower buds andthe roots, but not in young leaves. Overall, the results show that heat stress causes alternationin sucrose distribution in the plant, but may also have specificeffects on metabolic activities related to sucrose import andutilization in flower buds and flowers which in turn may enhancetheir abscission. Bell pepper, (Capsicum annuum L. cv. Maor), abscission, acid invertase, heat stress, reproductive organs, sink leaves     

Anatomical Investigation of Abscissed Avocado Flowers and Fruitlets

The anatomy of abscissed avocado fruitlets of the Hass varietywas compared with that of actively growing control fruitletsto investigate the reason for abscission. Control fruitletshad a small embryo (4–14 cells) and numerous endospermcells by seven days after pollination. Cotyledon developmenthad commenced by 21 days and a root/shoot axis was present at28 days. Over 90 per cent of the fruitlets which abscissed during thefirst week after the end of flowering were unfertilized and18 per cent were abnormal. By the fourth week after the endof flowering all abscissed fruitlets were fertilized and noneabnormal. Abnormalities included ovaries with more than oneembryosac or ovule, with an immature embryo-sac, with an ovulein an abnormal position or with a deficiency in ovule structure.In a few fertilized abscissed fruitlets either the embryo orendosperm had ceased development first. In the majority of casesembryo and endosperm were anatomically normal. The majorityof fertilized abscissed fruitlets had reached a stage correspondingto 14 days after pollination of the control tree. Degenerationwas often observed in both unfertilized and fertilized abscissedfruitlets. It is suggested that this degeneration occurred inthe period between the cessation of development and abscissionof the fruit. This period was approximately one week. The proportion of abnormal ovaries was too low to significantlyaffect yields, and sufficient flowers had been fertilized togive an adequate crop. No anatomical reason for the high rateof fruitlet abscission was observed. Persea americana Mill, avocado, abscission, embryo-sac, embryo, endosperm     

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.     

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.     

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.     

Changes in carbohydrates and mineral elements in Citrus leaves during flowering and fruit set

Mineral elements and metabolizable carbohydrates in Citrus leaves [ Citrus sinensis (L.) Osbeck cv. Washington navel] have been determined from bud sprouting until the end of the June drop and related to fruitlet growth and abscission. Mineral elements in old leaves decreased during the spring flush of growth and reached minimum values at flower opening, coinciding with a peak in abscission. This was followed by a rapid recovery in potassium and nitrogen to the initial values, with little change afterwards. Old leaves accumulate carbohydrates until flowering, and lose them during post-anthesis at a constant rate for more than 4 months; this rate of export is unaltered by the presence of a nearby growing fruit. Inflorescence leaves accumulate carbohydrates and mineral elements during post-anthesis; during the June drop there is an interruption in the accumulation of nitrogen and a net loss of phosphorus, potassium and carbohydrates from these leaves, coinciding with the attainment of the maximum growth rate of the fruit.
The two main periods of abscission coincide with minima in the amount of reserves in leaves, suggesting that a limitation in metabolite supply may be the primary cause of drop. There is a closer relationship of the fruit with inflorescence leaves than with old mature ones; however, the regulation of carbohydrate levels in the inflorescence leaves cannot be simply explained in terms of source-sink relationships with the nearby growing fruit, and the smaller size of inflorescence leaves vs. vegetative ones is not due to the presence of the flower during leaf development.     

Seasonal Accumulation of Starch by Components of the Kiwifruit Vine

The accumulation of starch by various components of 6-year-oldkiwifruit vines (Actinidia deliciosa var dehciosa cv Hayward)was recorded over one season Twenty vines were harvested periodicallythroughout the year and separated into perennial components(fibrous roots, structural roots, stump, stem, cordon, laterals)and current season's growth (shoots, leaves, and fruit) The concentration of starch in the fibrous roots followed asinusoidal trend Minimum concentrations occurred 98 d afterbudbreak, while the maximum concentrations occurred 182 d laterCorresponding times in the structural roots were approximately42 d earlier In the above-ground perennial components, elevatedconcentrations of starch in the cordon, fruiting wood and barkof the stem were evident at budbreak and fruit harvest (approx220 d later) In the case of the stem, concentrations were greatestat fruit harvest Because the biomass of the perennial componentswas found to be relatively constant throughout the year, starchconcentrations and contents were directly proportional in thesetissues For current season's growth, peak concentrations and contentsin leaves and shoots were observed at fruitset and fruit harvest,respectively For fruit, starch increased continuously untilharvest Approximately 30% of the total starch content accumulated inthe perennial components by leaf abscission was lost duringwinter and early summer Quantitative losses were greatest forthe roots Regeneration of the starch pools in the perennialcomponents of the vine occurred from midseason until leaf abscissionAt the same time, approximately five times more starch was accumulatedby the current season's growth, in particular the fruit, thanby the perennial components As a result of the difference inthe rate of accumulation, the starch content of the currentseason's growth increased from less than 10% midseason to nearly60% of the total starch content of the vine by fruit harvest The results were discussed in relation to the carbon economyof the kiwifruit vine, and compared with seasonal trends instarch concentrations found for other deciduous crops Actinidia deliciosa, kiwifruit, seasonal changes, starch content, whole plant     

Changes of Starch and Sorbitol in Leaves Before and After Removal of Fruits from Peach Trees

Changes in contents of nonstructural carbohydrates in leaves,as well as some characteristics of leaves before and after fruitremoval, were investigated in potted peach (Prunus persica L.)trees. Leaf area and dry mass per unit leaf area (SLW) at thefruit-maturation stage decreased with increasing numbers ofpeaches per tree, whereas the chlorophyll content per unit areain leaves of fruiting trees increased. The chlorophyll contentdecreased more rapidly upon removal of fruit than that in leavesof fruiting trees. The starch content per unit dry mass in leavesof fruiting trees at the fruit-maturation stage was lower thanthat in leaves of non-fruiting trees. Starch accumulated significantlyin leaves within 1 d of removal of fruit during the fruit-maturationstage and continued to increase thereafter. The accumulationof starch after removal of fruit occurred more rapidly thanthe decrease in chlorophyll content. Reducing and non-reducingsugars (total sugars) per unit dry mass in the leaves were higherin fruiting trees than in non-fruiting trees. After fruit removal,the total sugar content of leaves increased temporarily andthen gradually decreased. The sorbitol content per unit freshmass in leaves of fruiting trees during the fruit-maturationstage was slightly higher than that in leaves of non-fruitingtrees. One day after removal of fruit, the sorbitol contentincreased in parallel with the accumulation of starch and remainedhigh. The sucrose content of leaves did not change markedlyupon removal of fruit. Prunus persica L.; peach leaves; nonstructural carbohydrate; starch and sorbitol; fruit removal     

Nucellar Senescence and Ethylene Production as they Relate to Avocado Fruitlet Abscission

Events preliminary to avocado (Persea americana Mill) fruitletabscission include senescence of the nucellus and seed coat.The dynamics of nucellar deterioration and ethylene productionleading to seed abortion and abscission in avocado was examined.Excised branches bearing clusters of fruit from 1.0–2.5cm diameter were placed in humid chambers to reduce transpirationalwater loss. Fruitlets synchronously began nucellar and seedcoat deterioration 27–33 h after excision and rapidlyprogressed through stages of increasing degradation culminatingin abscission approximately 2 days later. The nucellus-seedcoat produced a temporary burst of ethylene at the first visiblesign of nucellar senescence followed by less ethylene productionin the mesocarp approximately 12 h later. All fruit underwentnucellar degradation prior to abscission. Exogenously appliedethylene accelerated fruitlet abscission with concentrationsas low as 1.0µ 1^–1 and with maximum response at100µl^–1 or greater. Maximal response took 2 days.Aminoethoxyvinyl-glycine (AVG) at 30 µ M inhibited ethyleneproduction and fruitlet abscission. The senescence process,however, was not af fected in any way by ethylene or AVG treatments.Observations of attached fruit suggest that nucellar-seed coatsenescence, concomitant ethylene production, and resulting abscissiontake place in a manner and within a time period similar to thatobserved on detached branches. It is concluded that nucellarand seed coat senescence is prerequisite to avocado fruitletabscission, and the time required from the first indicationof nucellar breakdown to abscission of that fruitlet appearsto be approximately 2 days. The senescence process is responsiblefor a large, transient rate increase in ethylene productionby the nucellus and perhaps seed coat. Ethylene is consideredto be the result rather than the cause of nucellar-seed coatsenescence. The ethylene thus produced induces fruit abscission.     

Seasonal Concentrations of Non-structural Carbohydrates of Five Actinidia Species in Fruit, Leaf and Fine Root Tissue

To characterize seasonal patterns of carbohydrate concentrationsin Actinidia species from different natural habitats, leaf,fruit and fine root tissue samples from five species (A. arguta,A. deliciosa, A. chinensis, A. polygama and A. eriantha) werecollected over one season, and analysed for fructose, glucose,sucrose, myo -inositol and starch concentrations. Sucrose andstarch peaked in leaf tissue around flowering time. In fruit,hexose sugars and/or myo -inositol transiently increased earlyin development. Starch accumulated in fruit of all species,beginning sooner after anthesis in A. arguta and A. polygamathan in the other species. Sucrose accumulation coincided withonset of net starch degradation in A. arguta but was delayedin the other species. At final fruit sampling, concentrationsof glucose and fructose were greater than sucrose in all speciesexceptA. arguta . myo -Inositol concentrations constituted >10%of total sugars for most of the season in leaf and fruit tissuesof all species except A. polygama. Fine roots of A. arguta andA. polygama contained significantly more starch and sucrosefor most of the year than those of the other species. Observeddifferences in seasonal carbohydrate patterns may reflect differentnatural habitats, with A. arguta and A. polygama growing naturallyin colder climates than the other species. Transient accumulationof sugars in fruit during early stages of development has beenconsidered to act as primary osmoticum for cell expansion. However,the presence of only low sugar concentrations in A. erianthaquestions this hypothesis. Copyright 2000 Annals of Botany Company Actinidia arguta, Actinidia deliciosa, Actinidia chinensis, Actinidia eriantha,Actinidia polygama , kiwifruit, carbohydrates, fruit, leaves, fine roots, seasonal