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     

The Abscission of Rose Petals

Petal abscission was studied in twelve hybrid tea rose (Rosahybrida L.) cultivars. At about 20 °C the time to petalabscission in uncut stems in greenhouses was the same as incut stems placed in water in the greenhouse or in a climate-controlledroom. The time between petal unfolding and abscission dependedon the cultivar, and varied between 12 and 35 d. The time topetal abscission of the cultivars was inversely correlated withtheir flower diameter at full bloom (linear regression, r² =0·82). In the cultivars with a relatively large flowerdiameter (10-18 cm) the petals fell without visible desiccationsymptoms, whereas in the group with a small diameter the petalswere partially or fully desiccated when shed. Fertilization occurred in some flowers of a few cultivars studied.In cultivars with a relatively large flower diameter (Papa Meilland,Cocktail, Dr. Verhage, Tineke) it had no effect on the timeto abscission in Motrea, Europa, and Carolien roses, which bearsmall flowers, the petals fell after fertilization, whereasin unfertilized flowers of the latter group of cultivars anabscission zone just above the uppermost node became activeand all parts above this node (pedicel and flower) turned brownand desiccated, though remained attached for more than a month. It is concluded that in the cultivars investigated: (a) thetime to petal abscission was inversely related to their flowerdiameter, (b) abscised petals were more desiccated in cultivarsin which the time to abscission was longer, (c) fertilizationhad little effect on the time to abscission in most cultivars,whereas the absence of fertilization prevented petal abscissionin a number of the small-diameter cultivars where it was replacedby flower abscission, and (d) cutting and placement in waterat 20 °C did not affect the time to abscission.Copyright1995, 1999 Academic Press Abscission, fertilization, flowers, petals, Rosa hybrida L., rose, water stress, carbohydrate stress     

Anatomy of Ethylene-induced Petal Abscission in Pelargonium x hortorum

When viewed under the light microscope, the abscission zoneat the petal base of Pelargonium x hortorum consisted of smallcells which, when stained with Toluidine Blue, possessed denselystained cells walls. After treatment with 1 µl l^-1 ethyleneat 22°C, the force required to separate the petals fromthe receptacle declined after a lag phase of only 30 min, withseparation complete 60-90 min later depending upon the stageof development of the flower. Transmission electron micrographsof the petal abscission zones showed evidence of cell wall degradation,particularly in the middle lamella. These cells also containedextensive rough endoplasmic reticulum and numerous Golgi bodiesribosomes. When abscission was complete, cells at the fractureface showed evidence of breakdown of cellular compartmentalization,often with little sign of an intact tonoplast. Scanning electronmicrographs of recently-abscissed surfaces showed that the epidermalcells surrounding the abscisson zone were turgid and rounded,whereas those of the mesophyll cells were partially collapsed.The micrographic evidence is consistent with the hypothesisthat ethylene-induced separation is caused by rapid enzymaticof the cell walls.Copyright 1993, 1999 Academic Press Abscission, cell walls, ethylene, flower, Pelargonium x hortorum     

Categories of Petal Senescence and Abscission: A Re-evaluation

In a previous paper (Woltering and van Doorn, 1988, Journalof Experimental Botany39: 1605–1616) we identified threetypes of flower life cessation: by petal wilting or withering,which was either ethylene-sensitive or insensitive, and by abscissionof turgid petals, which was ethylene-sensitive. These categoriestended to be consistent within families. Here we re-examinethese relationships by testing a further 200 species, and anumber of other families. As previously, flowering shoots wereexposed to 3 ppm ethylene for 24 h at 20 °C, in darkness.Most monocotyledonous species tested showed ethylene-insensitivepetal wilting, although ethylene-sensitive wilting occurredin the Alismataceae and Commelinaceae. Petals of the dicotyledonousspecies tested were generally sensitive to ethylene, exceptfor a few groups showing wilting (Crassulaceae, Gentianaceaeand Fumariaceae, and one subfamily in both the Ericaceae andSaxifragaceae). Petal abscission was generally ethylene-sensitive,but ethylene insensitivity was found in some Tulipa cultivarsand three Saxifraga species. In most tulip cultivars tested,the petals wilted and then fell. It is concluded that (a) theresponse to ethylene is often consistent within either familiesor subfamilies; and (b) a fourth category, ethylene-insensitivepetal abscission, exists both in monocotyledons and dicotyledons.Copyright 2001 Annals of Botany Company Ethylene sensitivity, flower longevity, petal abscission, petal wilting, petal withering, petal senescence, taxonomic categories     

Cellulase, Fruit Softening and Abscission in Red RaspberryRubus idaeusL. cv Glen Clova

The ripening of raspberry fruit (Rubus ideausL. cv Glen Clova)is associated with a climacteric rise in ethylene production.As the fruit pigments change from green to red there is a progressivesoftening, loss of skin strength and a breakdown of cell wallsin the mesocarp. An increase in cellulase (endo-1,4-ß-D-glucanase)in both drupelets and receptacles accompanies these changes.The localization of cellulase in the regions of the fruit associatedwith abscission zones suggest the enzyme may be involved infruit separation as well as softening. Rubus idaeusL; raspberry; fruit ripening; ethylene; abscission; cell wall breakdown; cellulase; endo-1,4-ß-D-glucanase     

Petal Abscission in Rose Flowers: Effects of Water Potential, Light Intensity and Light Quality

Petal abscission was studied in roses (Rosa hybrida L.), cvs.Korflapei (trade name Frisco), Sweet Promise (Sonia) and CaraMia (trade name as officially registered cultivar name). Unlikeflowers on plants in greenhouses, cut flowers placed in waterin the greenhouse produced visible symptoms of water stress,depending on the weather during the experiment and on the cultivar.Cut Frisco roses showed no visible signs of water stress andthe time to petal abscission was as in uncut flowers. In Soniaroses the symptoms of water stress varied from mild to severe,and the number of flowers in which the petals abscised variedfrom 100% (mild stress) to 0% (severe stress). An antimicrobialcompound in the vase water of Sonia roses, or removal of theleaves, alleviated the symptoms of water stress and increasedthe number of stems in which the petals abscised. Cut Cara Miaroses showed severe symptoms of water stress in all experimentsand petal abscission was found in only a few flowers, even whenthe stems were placed at 20 °C and low photon flux (15 µmolm^-2s^-1). Abscission in Sonia and Cara Mia roses was low or absentwhen the water potential of the leaves reached values below-2.0 MPa within the first 5 d of the experiment; such low valueswere not reached in Frisco roses. Addition of sucrose to the vase solution, together with an effectiveantimicrobial compound, had no effect on the time to petal abscission,at any light intensity. Placing flowers in far-red light alsohad no effect on abscission, compared with flowers placed inred light or white light of the same photon fluence. It is concluded that petal abscission in the rose cultivarsstudied is not affected by their water status unless the plantsreach a low water potential (about -2 MPa) early on during vaselife. Petal abscission is not inhibited by low light intensitynor affected by the Pr/Pfr ratio. Abscission; light intensity; petals; phytochrome; Rosa hybrida L.; rose; sugars; water potential     

Effects of Diazocyclopentadiene (DACP) and Silver Thiosulphate (STS) on Ethylene Regulated Abscission of Sweet Pea Flowers (Lathyrus odoratus L.)

The ethylene production rate of cut sweet pea flower buds increased37-fold during the first 48 h of their vase life. This increasein ethylene production was accompanied by petal wilting at 72h and abscission of the buds 24 h later. Exposure of the cutspikes to the ethylene action inhibitor diazocyclopentadiene(DACP, 170 µI 1^-1) for 18 h under fluorescent lights delayedsubsequent wilting and abscission and promoted bud opening.Silver thiosulphate (0·2 mM) was more effective thanDACP, delaying wilting for longer and preventing abscissionentirely.Copyright 1995, 1999 Academic Press Ethylene, abscission, silver thiosulphate, diazocyclopentadiene, flower senescence, wilting, sweet pea, Lathyrus odoratus L     

Role of ethylene in the senescence of isolated hibiscus petals

Senescence of petals isolated from flowers of Hibiscus rosa-sinensis L. (cv Pink Versicolor) was associated with increased ethylene production. Exposure to ethylene (10 microliters per liter) accelerated the onset of senescence, as indicated by petal in-rolling, and stimulated ethylene production. Senescence was also hastened by basal application of 1-aminocyclopropane-1-carboxylic acid (ACC). Aminooxyacetic acid, an inhibitor of ethylene biosynthesis, effectively inhibited ethylene production by petals and delayed petal in-rolling. In marked contrast to these results with mature petals, immature petals isolated from flowers the day before flower opening did not respond to ethylene in terms of an increase in ethylene production or petal in-rolling. Furthermore, treatment with silver thiosulfate the day before flower opening effectively prevented petal senescence, while silver thiosulfate treatment on the morning of flower opening was ineffective. Application of ACC to both immature and mature petals greatly stimulated ethylene production indicating the presence of an active ethylene-forming enzyme in both tissues. Immature petals contained less free ACC than mature, presenescent petals and appeared to possess a more active system for converting ACC into its conjugated form. Thus, while the nature of the lack of responsiveness of immature petals to ethylene is unknown, ethylene production in hibiscus petals appears to be regulated by the control over ACC availability.     

Role of Aminocyclopropane-1-carboxylic Acid (ACC) in Control of Ethylene Production in Fresh and Cold-stored Rose Flowers

The relationships between ethylene production, aminocyclopropane-1-carboxylicacid (ACC) content and ethylene-forming-enzyme (EFE) activityduring ageing and cold storage of rose flower petals (Rose hybridaL. cv. Gabriella) were investigated. During flower ageing at20 °C there was a climacteric rise in petal ethylene production,a parallel increase in ACC content, but a continuous decreasein EFE activity. Applied ACC increased petal ethylene productionc. 200-fold. During cold storage of flowers at 1 °C therewere parallel increases in petal ethylene production and ACCcontent, to levels greater than those reached in fresh flowersheld at 20 °C. EFE activity decreased during storage. Immediatelyafter cold-stored flowers were transferred to 20 °C ethyleneproduction and ACC levels were c. four times greater than infreshly cut flowers. These levels increased to maximum valuesof two to four times the maximum values reached during ageingof fresh, unstored, flowers. It was concluded that in rose petalsethylene synthesis is probably regulated by ACC levels and thatcold storage stimulates ethylene synthesis because it increasesthe levels of ACC in the petals. Key words: Rose flower, senescence, ethylene     

Sites of ethylene production in the pollinated and unpollinated senescing carnation (Dianthus caryophyllus) inflorescence

Production of endogenous ethylene from the styles, ovary and petals of pollinated and unpollinated flowers of Dianthus caryophyllus L. was measured. The rate of ethylene production of cut, unpollinated flowers aged in water at 18°C was low until the onset of petal wilting, when a rapid surge of ethylene occurred in all tissues. The flower ethylene production was evolved mostly from the styles and petals. The bases of petals from unpollinated, senescing flowers evolved ethylene faster and sometimes earlier than the upper parts. Treatment of cut flowers with propylene, an ethylene analogue, accelerated wilting of flower petals and promoted endogenous ethylene production in all flower tissues. Pollination of intact flowers also promoted endogenous ethylene production and caused accelerated petal wilting within 2–3 days from pollination. Although the data are consistent with the hypothesis that ethylene forms a link between pollination of the style and petal wilting, in the unpollinated flower the style and petals can evolve a surge of ethylene independently of each other, about the time when the petals irreversibly wilt. The results are discussed in relation to the role of ethylene in flower senescence.     

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.     

Cell Separation Processes in Plants--Models, Mechanisms and Manipulation

Abscission and dehiscence are developmental processes that involvethe co-ordinated breakdown of the cell wall matrix at discretesites and at specific stages during the life cycle of a plant.In this review we examine the events that influence the differentiationof abscission and dehiscence zone cells and the changes thatare associated with wall degradation. There is convincing evidenceto believe that ethylene and auxin co-ordinate the timing ofleaf, flower and fruit abscission but the events that regulatedehiscence and seed abscission are unclear. The use of transgenicplants and model systems such as Arabidopsis is assisting ourunderstanding of the mechanisms that regulate abscission anddehiscence and the application of this information will advanceour understanding of cell separation processes in general. Armedwith this knowledge it should be possible to either delay oraccelerate abscission and dehiscence, and this could have majorbenefits for the agricultural and horticultural industries.Copyright 2000 Annals of Botany Company Abscission, dehiscence, cell separation, wall degradation, gene expression, polygalacturonase, ß-1,4-glucanase, pathogenesis-related proteins, ethylene     

Relationship between petal abscission and programmed cell death in <Emphasis Type="Italic">Prunus yedoensis</Emphasis> and <Emphasis Type="Italic">Delphinium belladonna</Emphasis>

Depending on the species, the end of flower life span is characterized by petal wilting or by abscission of petals that are still fully turgid. Wilting at the end of petal life is due to programmed cell death (PCD). It is not known whether the abscission of turgid petals is preceded by PCD. We studied some parameters that indicate PCD: chromatin condensation, a decrease in nuclear diameter, DNA fragmentation, and DNA content per nucleus, using Prunus yedoensis and Delphinium belladonna which both show abscission of turgid petals at the end of floral life. No DNA degradation, no chromatin condensation, and no change in nuclear volume was observed in P. yedoensis petals, prior to abscission. In abscising D. belladonna petals, in contrast, considerable DNA degradation was found, chromatin was condensed and the nuclear volume considerably reduced. Following abscission, the nuclear area in both species drastically increased, and the chromatin became unevenly distributed. Similar chromatin changes were observed after dehydration (24 h at 60°C) of petals severed at the time of flower opening, and in dehydrated petals of Ipomoea nil and Petunia hybrida, severed at the time of flower opening. In these flowers the petal life span is terminated by wilting rather than abscission. It is concluded that the abscission of turgid petals in D. belladonna was preceded by a number of PCD indicators, whereas no such evidence for PCD was found at the time of P. yedoensis petal abscission. Dehydration of the petal cells, after abscission, was associated with a remarkable nuclear morphology which was also found in younger petals subjected to dehydration. This nuclear morphology has apparently not been described previously, for any organism.     

Ethylene-promoted tomato flower abscission and the possible involvement of an inhibitor

The abscission zone in tomato (Lycopersicon esculentum (L.) Mill. flower pedicels is morphologically distinguishable prior to separation and is delineated by an indentation of the epidermis. Exposure of excised pedicels with the flower attached to ethylene results in abscission within 12 h and this can be accelerated by flower removal. Abscission of excised pedicels with the flower removed takes place in the absence of exogenous ethylene but this is delayed by pretreatment with aminoethoxyvinyl glycine, an inhibitor of ethylene biosynthesis. The data presented support the hypothesis that flower tissue is the source of an abscission inhibitor.Abbreviations AVG aminoethoxyvinyl glycine - IAA indole-3-acetic acid     

A Flower and Pod Staging System for Soybean

Flower and pod abscission limit soybean yield. A system forquantifying flower and pod development based on the morphologicalappearance of the flower prior to and following anthesis hasbeen developed to aid in studies of pod abscission. Changesin the appearance of the corolla, primarily the banner petal,are used to distinguish the different stages of the system.External pistil dimensions have been correlated with internalfeatures for each stage of development. From anthesis to podset, pistil length and weight increase almost two- and fivefold,respectively, and ovule development progresses from unfertilizedegg cells to embryos surrounded by cellular endosperm. Pod determinedare correlated with ovule length and width and embryo cell number.Flower and pod stages can be determined in situ, thus permittingnon-destructive observation and experimental manipulation offlowers or pods without necessarily impeding their development.Stages have been identified that indicate precisely when podset occurs and when young pods cease growing and ultimatelyabscise. This system of flower and pod staging is useful instudies designed to assess effects of abiotic or biotic stressand genetic factors on pod set and abortion. Abscission, anthesis, Glycine max (L.) Merr, embryo development, pod set     

Factors affecting the Abscission of Reproductive Organs in Yellow Lupms (Lupinus luteus L.): II. THE EFFECTS OF GROWTH SUBSTANCES, DEFOLIATION, AND REMOVAL OF LATERAL GROWTH

Abscission of flowers in Lupinus luteus L. (var. Weiko II) withoutgrowth of ovaries is followed by abscission of small pods (15–20mm. long). Normally flower abscission is much more pronouncedthan pod abscission. Abscission was delayed on plants from which laterals or theirterminal and axifliary buds were removed. Flower abscissionwas not affected, but pod abscission increased as a result ofdefoliation. When flowers at the base of the main inflorescence were replacedby auxins and anti-auxins flower abscission was induced in eitheran auxin pattern in which most of the flowers near the siteof application dropped, and pods developed on the apical whorls,or an anti-auxin pattern in which pods developed on basal whorlsnear the site of application but not higher up. The anti-auxinpattern was similar to the pattern of abscission normally inducedby developing pods on basal whorls. -Naphthylacetic acid (NAA) was much more effective in inducingabscission than ß-indolylacetic acid (IAA). 2:3:5-triiodobenzoicacid (TIBA), NAA, and IAA applied in mixtures at various concentrationsacted mainly antagonistically, i.e. the abscission-inducingeffect of NAA and LAA was depressed in basal whorls, and inapical whorls the effect of TIBA was less prevalent. Consequentlythe effect of the mixtures on the total number of pods was aboutequal to that of the most active component by itself. All growth substances seemed to move much more efficiently inacropetal direction than in basipetal direction in the flowerstalk. Transport in lateral direction was very limited. The effect of growth substances applied on laterals was enhancedby defoliating the main 8tem. The influence of assimilates on flower and pod abscission andtransport of growth substances is discussed.     

Interrelationship between the Different Flower Parts during Emasculation-Induced Senescence in Cymbidium Flowers

In Cymbidium flowers emasculation by removal of the anther capand the pollinia, led to rapid colouration of the lip and advancedwilting of the petals and sepals. The ethylene production ofwhole flowers showed an emasculation-induced early peak in ethyleneevolution followed some days later by a second increase concomitantwith the wilting of the flower. In non-emasculated flowers theethylene production increased later and simultaneously withcolouration of the lip and wilting of the petals and sepals.At all stages of senescence, the contribution of the lip, petals,and sepals to the total amount of ethylene produced was negligible. Parallel to the increase in ethylene production of whole flowers,an increase in 1-aminocyclopropane-l-carboxylic acid (ACC) andmalonyl-ACC (MACC) in the central column and, to a lesser extent,in the ovary was observed. Also an increase in internal ethyleneconcentration was demonstrated and this, in contrast, was apparentin all the different flower parts. The activity of the ethylene-formingenzyme in lips, petals, and sepals showed an increase afteremasculation and such an effect could also be induced by treatmentof isolated lips with low concentrations of ethylene. The data indicate that senescence in Cymbidium flowers is regulatedby the central column and perhaps the ovary and that both ACCand ethylene may play a signalling role in inter-organ communication. Key words: 1-aminocyclopropane-l-carboxylic acid, ethylene, Cymbidium, senescence     

A study of ethylene in apple, red raspberry, and cherry

High ethylene levels were associated with flower abscission in apple (Malus sylvestris) and cherry (Prunus avium and Prunus cerasus), “June drop” of immature cherries, and harvest drop of apple and red raspberry (Rubus idaeus). However, an increase in ethylene content was not associated with June drop of apples and harvest drop of cherries. During the period of fruit ripening on the plant, the largest increases in ethylene occurred in apple flesh and red raspberry receptacular tissue. Ethylene remained low throughout the period of sweet and tart cherry ripening. The data obtained indicated marked ethylene gradients between adjacent tissues. Increases of ethylene in some tissues may have resulted from ethylene diffusion from adjacent tissues containing high levels of ethylene.     

Physiological Responses of Cut Rose Flowers to Exposure to Low Temperature: Changes in Membrane Permeability and Ethylene Production

Senescence of cut rose flowers (Rosa hybrida L. cv. Mercedes)at 22 °C occurred earlier in flowers previously held at2 °C for 10 d or 17 d than in freshly cut flowers. Thisadvanced senescence was observed as an earlier increase in bothethylene production rate and membrane permeability. The risein ethylene production preceded the rise in the level of ionleakage from petals, and this in turn preceded visible symptomsof petal death. Applied ethylene stimulated ion leakage andinhibitors of ethylene synthesis and action (amino-oxyaceticacid and silver thiosulphate respectively) inhibited the normalincrease in ion leakage. The maximum rate of ethylene productionof 22 °C increased markedly in petals of flowers previouslyheld at 2 °C, up to nine times the level in fresh flowers.We conclude that during exposure of rose flowers to 2 °C,in addition to senescence, processes were induced which ledto stimulated ethylene production after transferral to 22 °C.Ethylene apparently caused the subsequent advance in membranepermeability and senescence. Key words: Rose flower, Low temperature, Senescence