Fruit age and changes in abscisic Acid content, ethylene production, and abscission rate of cotton fruits

The relationships of fruit age, abscisic acid (ABA) concentration, ethylene evolution, and abscission rates were studied in an effort to determine why cotton (Gossypium hirsutum L., cv. Deltapine 16) fruits rarely abscise more than 15 days after anthesis. Because abscission of cotton fruits is increased by conditions that limit photosynthesis, greenhouse-grown plants with fruits of various ages were placed in dim light for 3 days to induce high rates of fruit abscission. Abscission rates, ABA concentrations, and ethylene evolution rates were determined for fruits of various ages. Almost all of the young fruits abscised, but abscission rate declined with age until almost no abscission was observed in fruits that were 15 or more days past anthesis.     

Abscisic Acid: correlations with abscission and with development in the cotton fruit

Abscisic acid was measured in developing cotton fruit (Gossypium hirsutum) by means of gas-liquid chromatography. High levels of abscisic acid occurred in correlation with abortion and abscission of young fruit, with low germination of immature seed, and with senescence and dehiscence of mature fruit. Declining or low levels of abscisic acid occurred in correlation with the period of most rapid fruit growth and with high germination of immature and mature seed. Young fruit of cultivar Acala 4-42 contained about twice as much abscisic acid as young fruit of cultivar Acala SJ-1, and this difference is correlated with a higher rate of young fruit abscission in Acala 4-42. Young fruit abscising late in the fruiting season contained about twice as much abscisic acid as young fruit abscising early in the fruiting season.     

Physiology and Chemistry of Substances Accelerating Abscission in Senescent Petioles and Fruit Stalks

Senescent petioles of Coleus rehneltianus Berger. Phaseolus vulgaris L. cv. Saxa. Acer pseudoplatanus L., and senescent fruit stalks of Malus domestica Borkh. cv. Golden Delicious contain at least three abscission accelerating substances, which were isolated by extraction with methanol or with water and by diffusion into agar. They were purified by thin-layer chromatography and bioassayed in a special abscission test using Coleus explants. Two of these abscission accelerators could be conclusively identified by thin-layer chromatography and by gas chromatography as abscisic acid and xanthoxin. The third substance, which has acidic properties and is less polar than abscisic acid, could not be identified. The concentration and the absolute amount of abscisic acid in Coleus petioles were found to decrease during their development, young petioles having the highest concentration. No evidence was found that the three abscission accelerators or synthetic abscisic acid and xanthoxin affect the production of ethylene in Caleus explatns. The results obtained do not support the hypothesis that senescent petioles contain a specific “senescence factor”, which stimulates abscission via ethylene production.     

The physiological significance of phenylacetic Acid in abscising cotton cotyledons

The physiological role of phenylacetic acid (PAA) as an endogenous regulator of cotyledon abscission was examined using cotton (Gossypium hirsutum L. cv LG 102) seedlings. Application of 100 micromolar or more PAA to leafless cotyledon abscission-zone explants resulted in the retardation of petiole abscission and a decrease in the rise of ethylene evolution that normally accompanies aging of these explants in vitro. The partial inhibition of ethylene evolution in these explants by PAA was indirect since application of this compound stimulated short-term (<24 hours) ethylene production. PAA treatment partially suppressed the stimulation of petiole abscission elicited by either ethylene or abscisic acid. Both free and an acid-labile, bound form of PAA were identified in extracts prepared from cotyledons. No discernible pattern of changes in free or bound PAA was found during the course of ethylene-induced cotyledon abscission. Unlike indole-3-acetic acid, transport of PAA in isolated petiole segments was limited and exhibited little polarity. On the whole, these results are not consistent with the direct participation of PAA in the endogenous regulation of cotyledon abscission.     

Cherry fruit abscission: evidence for time of initiation and the involvement of ethylene

Initiation of abscission at the pedicel-fruit zone in the sour cherry (Prunus cerasus L. cv. Montmorency) occurs near the transition of Stage II to Stage III of fruit growth. The preinitiation phase is characterized by a high fruit removal force (FRF) and explants prepared from fruits during this period do not undergo abscission as indexed by a reduction in FRF. Ethylene does not cause a significant reduction in FRF either in attached fruit or in explants prepared during this period. By contrast, after initiation (Stage III of fruit growth), there is a marked decrease in FRF with fruit development, explants prepared from fruits during this period undergo abscission, and ethylene markedly promotes the loss in break-strength. Neither the rate of evolution nor the internal concentration of ethylene in the fruit were correlated with fruit abscission. Similar abscission responses, as indexed by FRF and sensitivity to ethylene, were observed in attached fruit and in detached fruit explants.     

Role of ethylene in stimulating stylar abscission in pistil explants of lemons

Abscission in styles of excised Citrus limon (cv. Lisbon) pistils was stimulated by addition of 70 μ M 2-chloroethylphosphonic acid (ethephon) or 0.1 m M 1-aminocyclopropane-1-carboxylic acid (ACC) to the defined medium of cultures. To study the relationship between ethylene and abscission, we used gas chromatography to analyze ethylene in cultures containing a test medium plus or minus abscission-active chemicals. In the presence of ethephon or ACC, ethylene levels in sealed tubes increased rapidly, suggesting that these compounds stimulated abscission because they were converted to ethylene. In the presence of test medium or the inhibitor of abscission 2 μ M picloram, the low ethylene levels found in sealed tubes did not differ strikingly in the two treatments. Ethylene production rates measured prior to abscission with test medium or in the presence of picloram were not markedly different either, although picloram completely inhibited abscission. Stylar abscission was delayed but not prevented by 50 μ M aminoethoxyvinylglycine, an inhibitor of ethylene biosynthesis, and by hypobaric conditions (280 mm Hg) which removed ethylene from cultures. We concluded that ethylene is an important factor regulating stylar abscission in vitro and suggest that the inhibitory effect of picloram involves a process other than detectable ethylene production. Our results do not exclude the possibility that picloram affects enodgenous ethylene biosynthesis and/or metabolism and/or tissue retention.     

Involvement of ethylene in the action of the cotton defoliant thidiazuron

The effect of the defoliant thidiazuron (N-phenyl-N′-1,2,3-thiadiazol-5-ylurea) on endogenous ethylene evolution and the role of endogenous ethylene in thidiazuron-mediated leaf abscission were examined in cotton (Gossypium hirsutum L. cv Stoneville 519) seedlings. Treatment of 20- to 30-day-old seedlings with thidiazuron at concentrations equal to or greater than 10 micromolar resulted in leaf abscission. At a treatment concentration of 100 micromolar, nearly total abscission of the youngest leaves was observed. Following treatment, abscission of the younger leaves commenced within 48 hours and was complete by 120 hours. A large increase in ethylene evolution from leaf blades and abscission zone explants was readily detectable within 24 hours of treatment and persisted until leaf fall. Ethylene evolution from treated leaf blades was greatest 1 day posttreatment and reached levels in excess of 600 nanoliters per gram fresh weight per hour (26.7 nanomoles per gram fresh weight per hour). The increase in ethylene evolution occurred in the absence of increased ethane evolution, altered leaf water potential, or decreased chlorophyll levels. Treatment of seedlings with inhibitors of ethylene action (silver thiosulfate, hypobaric pressure) or ethylene synthesis (aminoethoxyvinylglycine) resulted in an inhibition of thidiazuron-induced defoliation. Application of exogenous ethylene or 1-aminocyclopropane-1-carboxylic acid largely restored the thidiazuron response. The results indicate that thidiazuron-induced leaf abscission is mediated, at least in part, by an increase in endogenous ethylene evolution. However, alterations of other phytohormone systems thought to be involved in regulating leaf abscission are not excluded by these studies.     

Endogenous ethylene and abscisic Acid relative to phytogerontology

Endogenous production of ethylene and endogenous levels of abscisic acid were measured from Hibiscus rosa-sinensis L. abscission zone explants at six stages of development: tight bud, open flower, closed flower, petal abscission, calyx abscission, and peduncle abscission.     

Role of abscisic acid in the phenomena of abscission of flower buds and bolls of cotton (Gossypium hirsutum L.) and its reversal with other plant regulators

The investigations carried out to find the role of abscisic acid in the phenomena of abscission of flower buds and bolls of cotton (Gossypium hirsutum L. cv. ‘H-14’) have shown abscisic acid content to be low in retained bolls as compared to that in the abscising ones of the same age, suggesting that relatively higher endogenous abscisic acid content to be promotive of abscission. Abscisic acid applied exogenously either to intact flower buds/bolls or boll explants promoted their abscission. Naphthalene acetic acid not only reduced abscission but also could erase completely the promotive effect of abscisic acid on abscission. Gibberellic acid promoted abscission in intact buds and boll explants but applied to intact bolls it reduced their shedding even more than naphthalene acetic acid. Gibberellic acid could also counteract the promotive effect of abscisic acid in the case of intact bolls but enhanced that of boll explants. All the cytokinin-furfurylamino-purine treatments given other than at the abscission zone promoted abscission. Furfurylaminopurine applied in combination with abscisic acid showed some antagonistic effect in the case of intact bolls and boll explants abscission zone treatments. Ascorbic acid applied at a relatively lower dose (0.025 mM) reduced shedding but applied at a higher dose it showed promotion. Ascorbic acid could erase the promotive effect of abscisic acid on abscission to a significant extent.     

Mechanical wounding and abscission in citrus

Fruit detachment force (FDF), ethylene evolution, fruit and leaf drop were determined in Citrus sinensis for periods up to 96 h after mechanical wounding. Injury by removing a thin section of mature fruit flavedo reduced FDF, increased ethylene evolution and promoted abscission. Injuring flavedo 1 cm below the calyx was more effective at reducing FDF than injuring flavedo at the equator or the blossom‐end of mature fruit. Injuring the calyx or peduncle of mature fruit, or injuring three leaves closest to the mature fruit did not reduce FDF. Immature fruitlets either did not abscise or underwent low rates of abscission in response to mechanical wounding, depending on age. Inhibiting ethylene binding in wounded mature fruit with 1‐methylcyclopropene (1‐MCP) increased ethylene evolution compared with wounded fruit alone, but the reduction in FDF was similar. When an ethylene biosynthesis inhibitor (aminoethoxyvinylglycine, AVG) was used, reduction in FDF of wounded mature fruit exposed to AVG was similar to that of wounded fruit alone but ethylene production was markedly reduced. Wounding mature leaf blades in the presence or absence of 1‐MCP resulted in elevated but equal ethylene evolution up to 48 h after wounding, however, no leaf drop occurred. Thereafter, ethylene evolution was higher in 1‐MCP‐treated wounded leaves. Removing up to 77% of the total mature leaf area did not cause leaf drop, nor did wounding tissue across the laminar or petiolar abscission zones. Leaflets of 5 mm length reached nearly 100% abscission after mechanical wounding, whereas wounding leaves 20 mm length resulted in 15% abscission. The data suggest that mechanical wounding of flavedo results in mature fruit abscission, and ethylene binding may not be mandatory to initiate abscission in citrus fruit. The differential response of fruit and leaves at different ages to wounding may be related to potential contribution to carbohydrate accumulation, and production and sensitivity of tissues to an abscission signal(s).     

The Role of Abscisic Acid in Flower Abscission of Lupinus luteus

The role of abscisic acid in the control of flower abscission in Lupinus luteus L. was examined. Using a modified extraction and purification technique, endogenous abscisic acid levels in the upper flowers of an inflorescence were found to increase markedly some days before abscission could be detected. When abscisic acid was injected into flower-bearing nodes or fed via the roots, no increase in the abscission rate was obtained at any position in the flowerhead. Application of abscisic acid to only the leaves resulted in a marked increase in flower abscission. The role of abscisic acid per se as a primary controlling factor of flower abscission in yellow lupin is questioned.     

Abscission: role of abscisic Acid

The effect of abscisic acid on cotton (Gossypium hirsutum L. cv. Acala 4-42) and bean (Phaseolus vulgaris L. cv. Red Kidney) explants was 2-fold. It increased ethylene production from the explants, which was found to account for some of its ability to accelerate abscission. Absci is acid also increased the activity of cellulase. Increased synthesis of cellulase was not du to an increase in aging of the explants but rather was an effect of abscisic acid on the processes that lead to cellulase synthesis or activity.     

Focus on Ethylene: Ethylene and Hormonal Cross Talk in Vegetative Growth and Development

Ethylene is a gaseous plant hormone that most likely became a functional hormone during the evolution of charophyte green algae, prior to land colonization. From this ancient origin, ethylene evolved into an important growth regulator that is essential for myriad plant developmental processes. In vegetative growth, ethylene appears to have a dual role, stimulating and inhibiting growth, depending on the species, tissue, and cell type, developmental stage, hormonal status, and environmental conditions. Moreover, ethylene signaling and response are part of an intricate network in cross talk with internal and external cues. Besides being a crucial factor in the growth control of roots and shoots, ethylene can promote flowering, fruit ripening and abscission, as well as leaf and petal senescence and abscission and, hence, plays a role in virtually every phase of plant life. Last but not least, together with jasmonates, salicylate, and abscisic acid, ethylene is important in steering stress responses.This Update provides recent insights into the role of ethylene on vegetative growth, both at the cellular and the whole-plant levels, with special attention to hormonal cross talk. Due to space restrictions, this Update is mainly focused on Arabidopsis (Arabidopsis thaliana).     

Arrested Leaf Abscission in the Non-Abscising Variety of Pubescent Birch: Developmental,Morphological and Hormonal Aspects

A novel type of abscission development, arrested abscission,is described for a rare variety of pubescent birch (Betula pubescensEhrh. f. hibernifolia Ulvinen), the leaves of which dehydrateand remain attached to the plant. Anatomical examination ofthe abscission zone revealed that its development is temporallyand spatially similar to that of normal leaves of Betula pubescens.Floodingand exogenous ethylene were effective in inducing shedding ofthe leaves at the beginning of the growing season, butthis effectwas lost in senescing leaves. Leaf hydration and abscissionwere retained in the presence of external abscisic acid (ABA)inthe non-abscising variety, which also had a lower level of endogenousABA. The observed responses together with the alteredhormonallevel in this variety suggest that apart from ethylene, ABAis involved in autumnal abscission indirectly by regulatingtheleaf water status. Key words: ABA deficiency, abscisic acid, abscission zone, ethylene, senescenc     

Involvement of Abscisic Acid in Ethylene-Induced Cotyledon Abscission in Cotton Seedlings

Cotton (Gossypium hirsutum L. cv LG102) seedlings raised from seeds exposed to 100 [mu]M norflurazon (NFZ) during imbibition contained reduced levels of free abscisic acid (ABA) and were visibly achlorophyllous. Exposure of untreated cotton seedlings to ethylene concentrations >1 [mu]L/L for 24 h resulted in cotyledon abscission. In contrast, exposure of NFZ-treated seedlings to concentrations of ethylene [less than or equal to]50 [mu]L/L elicited no cotyledon abscission. Application of ABA, an ABA analog, or jasmonic acid to NFZ-treated seedlings restored ethylene-induced abscission. Isolated cotyledonary node explants prepared from NFZ-treated seedlings exhibited an altered dose-response pattern of ethylene-induced petiole abscission. Endogenous levels of free IAA were unaltered in NFZ-treated seedlings. Ethylene treatment (50 [mu]L/L, 24 h) had no effect on free indoleacetic acid (IAA) levels in either control or NFZ-treated seedlings. Levels of conjugated (ester plus amide) IAA were substantially increased in NFZ-treated seedlings regardless of ethylene treatment. These results indicate that endogenous ABA plays an essential, but physiologically undefined, role in ethylene-induced cotyledon abscission in cotton.     

The Role of Ethylene in the Shedding of Red Raspberry Fruit

The production of ethylene by red raspberry (Rubus idaeus L.cv. Glen Clova) fruit increased climacterically during development.The concentration of ethylene within green fruit was low butincreased substantially as fruit abscission and ripening commenced.The receptacle contained higher concentrations than the drupeletsat all stages measured. In the mature ripening fruit the ethyleneconcentrations were found to be physiologically significant,and would accelerate the abscission of large green non-abscisingfruit if supplied as a fumigant. The addition of ethylene toripe fruit did not accelerate abscission, probably because saturatinglevels occurred naturally within these fruit. Reduction of ethylenesynthesis rates using the inhibitor of ethylene production aminoethoxyvinylglycine(AVG) reduced the rate of abscission zone weakening which occursin detached large green fruit. The rate of ethylene productionwas found to be dependent on the supply of the precursor l-aminocyclopropane-l-carboxylicacid (ACC). This only accumulated to any extent in those ripefruit with high rates of ethylene production. Rubus idaeus, raspberry, abscission, fruit ripening, ethylene, aminocyclopropane-l-carboxylic acid     

Is uronic acid oxidase involved in the hormonal regulation of abscission in explants of citrus leaves and fruits?

The role of uronic acid oxidase in abscission was studied in explants of citrus ( Citrus sinensis L. Osbeck; var. Shamouti) leaves and fruits. In leaf explants, activity of uronic acid oxidase prior to onset of abscission and the rate of abscission were markedly accelerated by ethylene and delayed by 2,4-dichlorophenoxyacetie acid. Similar results were obtained for uronic acid oxidase activity in the exocellular fraction of young fruit explants. In mature fruit explants, treated with ethylene, an immediate increase in activity was evidenty in the non-active shoot/peduncle abscission zone, whereas in the calyx abscission zone the rise in activity occurred after a prolonged exposure to ethylene, when most of the fruits had already abscised. Whenever ethylene enhanced uronic acid oxidase activity, 2,4-dichlorophenoxyacetic acid delayed it. A gradient of decreasing activity or uronic acid oxidase was recorded from both sides of the abscission zone in leaves and fruits toward the separation line, where activity was the lowest as compared with the activity found in adjacent tissues. It is suggested that uronic acid oxidase is involved in senescence and cell wall degradation. However, it is yet questionable whether this enzyme is directly related to the control mechanism of abscission.     

Abscisic acid- and ethylene-induced defoliation of Radermachera sinica L.

Radermachera sinica L. is an ornamental plant with demonstrated sensitivity to ethylene-induced leaf abscission. In this study, we examine the relationship between abscisic acid (ABA) and ethylene in initiating the abscission response. Treatment with 1 l L^\s-1 of ethylene, 1 mM 1-aminocyclopropane-1-carboxylic acid (ACC) or 1 mM ABA resulted in complete defoliation of leaf explants. Application of 0.125 mM silver thiosulfate (STS) inhibited ethylene- and ACC-induced abscission but had no effect on explants treated with ABA. The ABA-induced abscission was unaffected by treatment with aminoethoxyvinylglycine (AVG) or aminooxyacetic acid (AOA). Treatment of explants with 1 mM cobalt chloride (CoCl₂) or 2000 l L^\s-1 of norbornadiene (NBD) completely inhibited abscission in explants treated with 1 l L^\s-1 ethylene or 1 mM ACC but they were only marginally effective in blocking ABA-induced abscission despite the lower level of endogenous ethylene. ABA appeared to increase the sensitivity of explants to ethylene. However, the evidence suggests that ABA may also function independent of ethylene to induce leaf abscission in R. sinica.Abbreviations ABA abscisic acid - ACC 1-aminocyclopropane-1-carboxylic acid - AOA aminooxyacetic acid - AVG aminoethoxyvinylglycine - CoCl₂ cobalt chloride - NBD norbornadiene - STS silver thiosulfate     

Relationship between Leaf Water Status and Endogenous Ethylene in Detached Leaves

The pattern of changes in the internal concentration of ethylene in response to water stress was investigated in species with leaves that do abscise and leaves that do not abscise. When leaves which abscise were detached and exposed to dry air for up to 6 hours, a continuous increase of internal ethylene was observed. In water-stressed leaves which do not abscise only a transient rise in ethylene occurred. The peak, which was attained after 30 to 120 minutes, depending on the species studied, was followed by a sharp decline to the initial level. The principal site of ethylene production in response to a short period of water stress was in the blades rather than the petioles in both types of leaves. The internal ethylene level in leaves was reduced by pretreatment with the ethoxy analog of rhizobitoxine (an inhibitor of ethylene biosynthesis) or by maintaining the leaves under subatmospheric pressure. The results obtained by these methods showed that ethylene was not involved in the mechanism of stomatal movement in either turgid or in stressed leaves. Also, the increase in leaf abscisic acid content and the depletion of gibberellins induced by water stress were not related to the internal concentration of ethylene in the detached leaf. The different patterns of drought-induced ethylene production observed in the blades of leaves which exhibit abscission compared with those which do not exhibit abscission may indicate the involvement of ethylene in a primary event in the process of leaf abscission induced by water stress.