Abscission Zone Cells Research Articles

Spatio-temporal localization of LlBOP following early events of floral abscission in yellow lupine

The phenomenon of excessive flower abscission in yellow lupine is a process of substantial interest to the agricultural industries, because it substantially affects the yield. The aim of this work was to provide an analysis of the changes taking place precisely in the abscission zone (AZ) during early stages of flower separation. We put particular emphasis on mRNA accumulation of BOP (BLADE ON PETIOLE) gene encoding a transcriptional factor so far considered to be essential for AZ formation. Our results show that the AZ displays a particular transcriptional network active in the specific stages of its function, as reflected by the expression profile of LlBOP. Noteworthy, spatio-temporal LlBOP transcript accumulation in the elements of pedicel vascular tissue reveals divergent regulatory mechanism of its activity. We have also found that AZ cells accumulate reactive oxidative species following abscission and what is more, become active due to the increasing amount of uridine-rich small nuclear RNA, accompanied by poly(A) mRNA intensive synthesis. Our paper is a novel report for BOP involvement in the AZ functioning in relation to the whole transcriptional activity of AZ and overall discussed regarding BOP role as a potential mobile key regulator of abscission.

Spatio-temporal IAA gradient is determined by interactions with ET and governs flower abscission

The abscission zone (AZ) is a specialized tissue that usually develops at the base of an organ and is highly sensitive to phytohormones, e.g., abscisic acid (ABA), ethylene (ET), and gibberellins (GAs). A current model of organ abscission assumes that the formation of an auxin gradient around the AZ area determines the time of shedding; however, that thesis is supported by studies that are primarily concerned with auxin transporters. To better understand the events underlying the progression of abscission, we focused for the first time on indole-3-acetic acid (IAA) distribution following AZ activation. We performed a series of immunolocalization studies in proximal and distal regions of floral AZ cells in yellow lupine, which is an agriculturally important legume. The examined phytohormone was abundant in natural active AZ cells, as well as above and below parts of this structure. A similar gradient of IAA was observed during the early steps of abscission, which was induced artificially by flower removal. Surprisingly, IAA was not detected in inactive AZ cells. This paper is also a consequence of our comprehensive studies concerning the phytohormonal regulation of flower abscission in yellow lupine. We present new data on interactions between IAA and ET, previously pointed out as a strong modulator of flower separation. The detailed analysis shows that disruption of the natural auxin gradient around the AZ area through the application of synthetic IAA had a positive effect on ET biosynthesis genes. We proved that these changes are accompanied by an accumulation of the ET precursor. On the other hand, exposure to ET significantly affected IAA localization in the whole AZ area in a time-dependent manner. Our results provide insight into the existence of a spatio-temporal sequential pattern of the IAA gradient related to the abscission process; this pattern is maintained by interactions with ET. We present new valuable evidence for the existence of conservative mechanisms that regulate generative organ separation and can help to improve the yield of agronomically significant species in the future.

Vesicular trafficking in abscission zone cells during ethylene-promoted fruit abscission in citrus

Vesicular trafficking in abscission zone cells during ethylene-promoted fruit abscission in citrus

Vesicular trafficking in abscission zone cells during ethylene-promoted fruit abscission in citrus

Abscission is a cell separation process by which plants are able to shed aerial organs. This physiological process proceeds through a breakdown in adhesion between a group of specialized cells that usually differentiate in particular positions of the plant body named abscission zones (AZs). The occurrence of fruit abscission has special relevance in fruit tree crops such as citrus since it may have negative consequences on yield. To identify AZ-specifically expressed genes involved in citrus fruit abscission, we used a combined strategy of laser microdissection and microarray analysis to compare the global expression profiles of calyx abscission zone (AZ-C) cells of citrus fruits and adjacent fruit rind cells during ethylene-promoted fruit abscission. The comparison enabled to identify a set of potentially specific genes including those associated with vesicular transport and intracellular transport vesicles. The first ultrastructural observations on AZ-C cells during fruit abscission (Iwahori and van Steveninck, 1976) showed an increase in rough endoplasmic reticulum profiles and dictyosome stacks with associated vesicles that fused with the plasma membrane to release their contents to the cell wall. In this comparative transcriptomic survey, we highlighted specific over-expression in AZ-C cells of genes belonging to the major transport steps along the secretory and the endocytic pathways of vesicle trafficking. In particular, it is likely that some of the genes identified in this study would represent major candidate genes for further biotechnological approaches aimed at trying to solve or at least mitigate agronomic issues regarding citrus fruit abscission.

Vesicular trafficking in abscission zone cells during ethylene-promoted fruit abscission in citrus

Vesicular trafficking in abscission zone cells during ethylene-promoted fruit abscission in citrus

Gibberellic acid affects the functioning of the flower abscission zone in Lupinus luteus via cooperation with the ethylene precursor independently of abscisic acid

The abscission of plant organs is a phytohormone-controlled process. Our study provides new insight into the involvement of gibberellic acid (GA3) in the functioning of the flower abscission zone (AZ) in yellow lupine (Lupinus luteus L.). Physiological studies demonstrated that GA3 stimulated flower abortion. Additionally, this phytohormone was abundantly presented in the AZ cells of naturally abscised flowers, especially in vascular bundles. Interesting interactions among GA3 and other modulators of flower separation were also investigated. GA3 accumulated after treatment with the ethylene (ET) precursor 1-aminocyclopropane-1-carboxylic acid (ACC). Abscisic acid (ABA) treatment did not cause such an effect. Furthermore, the expression of the newly identified LlGA20ox1 and LlGA2ox1 genes encoding 2-oxoglutarate-dependent dioxygenases fluctuated after ACC or ABA treatment which confirmed the existence of regulatory crosstalk. GA3 appears to cooperate with the ET precursor in the regulation of AZ function in L. luteus flowers; however, the presented mechanism is ABA-independent.

INFLORESCENCE DEFICIENT IN ABSCISSION-like is an abscission-associated and phytohormone-regulated gene in flower separation of Lupinus luteus

Abscission is a natural process that occurs to facilitate shedding of no longer needed organs, but on the other hand, can be triggered by certain environmental conditions, e.g. biotic or abiotic stresses. Regardless of the stimuli, organ shedding takes place specifically at the abscission zone (AZ). A signaling pathway that controls this process in Arabidopsis thaliana from ligand to receptors has been proposed. However, knowledge concerning the influence of plant hormones on these molecular elements still remains enigmatic. Excessive and premature flower abscission in the crop species Lupinus luteus L. is a process of substantial interest to the agricultural industry, as it can affect yield. Our strategy combined molecular studies, comprehensive ultrastructural and histological analysis, as well as exogenous hormone treatment to describe the contribution of the Lupinus IDA-like gene in flower abscission. In the AZ of the naturally abscised flowers, the differentiation of morphologically distinct cells characterized by progressive degradation processes was accompanied by LlIDL mRNA accumulation. A similar effect was observed following early steps of AZ activation and after abscisic acid or ethylene treatments. These phytohormones, previously pointed out as key stimulators of flower separation, altered the temporal expression pattern of LlIDL. Exogenous EPIP peptide synthesized on the basis of LlIDL sequence, significantly increased flower abortion rate, which indicates that this motif governs protein activity. In conclusion, our data provide new evidence for LlIDA involvement in both the early and late events of flower abscission supported by detailed spatiotemporal characterization of AZ cell structure and ultrastructure.

Visualizing Morphological Changes of Abscission Zone Cells in Arabidopsis by Scanning Electron Microscope.

Scanning electron microscope (SEM) is a type of electron microscope which produces detailed images of surface structures. It has been widely used in plants and animals to study cellular structures. Here, we describe a detailed protocol to prepare samples of floral abscission zones (AZs) for SEM, as well as further image analysis. We show that it is a powerful tool to detect morphologic changes at the cellular level during the course of abscission in wild-type plants and to establish the details of phenotypic alteration in abscission mutants.

A MADS-box transcription factor of ‘Kuerlexiangli’(Pyrus sinkiangensis Yu) PsJOINTLESS gene functions in floral organ abscission

MADS-box proteins have been implicated in many biological processes. However, plant MADS-box proteins functioning in floral organ abscission and the underlying physiological mechanisms remain poorly understood. Here, we report the identification and functional characterization of PsJOINTLESS isolated from ‘Kuerlexiangli’. PsJOINTLESS had a complete open reading frame of 672bp, encoding a 224 amino acid peptide, and shared high sequence identities with MADS-box from other plants. PsJOINTLESS was subcellularly targeted to the nucleus, supporting its role as a transcription factor. Expression levels of PsJOINTLESS in the calyx tube were strongly induced by calyx abscission treatment at 6d after full bloom. Overexpression of PsJOINTLESS in tomato enhanced the rate of pedicel abscission rate. Of special note, the transgenic plants increased the abscission zone cell layer compared with wild type. Furthermore, the tomato transgenic lines showed thinner flower pedicels, more cell number and small pedicel cell size. The cellulase activity in pedicel abscission zone of transgenic plants was higher than that of wild type. In addition, steady-state mRNA levels of five cell wall hydrolase genes coding for either functional or regulatory proteins were induced to higher levels in the transgenic lines. These results clearly demonstrate that PsJOINTLESS may affect pedicel abscission zone development.

Localization of Sphingolipid Enriched Plasma Membrane Regions and Long-Chain Base Composition during Mature-Fruit Abscission in Olive

Sphingolipids, found in membranes of eukaryotic cells, have been demonstrated to carry out functions in various processes in plant cells. However, the roles of these lipids in fruit abscission remain to be determined in plants. Biochemical and fluorescence microscopy imaging approach has been adopted to investigate the accumulation and distribution of sphingolipids during mature-fruit abscission in olive (Olea europaea L. cv. Picual). Here, a lipid-content analysis in live protoplasts of the olive abscission zone (AZ) was made with fluorescent dyes and lipid analogs, particularly plasma membrane sphingolipid-enriched domains, and their dynamics were investigated in relation to the timing of mature-fruit abscission. In olive AZ cells, the measured proportion of both polar lipids and sphingolipids increased as well as endocytosis was stimulated during mature-fruit abscission. Likewise, mature-fruit abscission resulted in quantitative and qualitative changes in sphingolipid long-chain bases (LCBs) in the olive AZ. The total LCB increase was due essentially to the increase of t18:1(8E) LCBs, suggesting that C-4 hydroxylation and Δ8 desaturation with a preference for (E)-isomer formation were quantitatively the most important sphingolipids in olive AZ during abscission. However, our results also showed a specific association between the dihydroxylated LCB sphinganine (d18:0) and the mature-fruit abscission. These results indicate a clear correlation between the sphingolipid composition and mature-fruit abscission. Moreover, measurements of endogenous sterol levels in the olive AZ revealed that it accumulated sitosterol and campesterol with a concomitant decrease in cycloartenol during abscission. In addition, underlying the distinct sterol composition of AZ during abscission, genes for key biosynthetic enzymes for sterol synthesis, for obtusifoliol 14α-demethylase (CYP51) and C-24 sterol methyltransferase2 (SMT2), were up-regulated during mature-fruit abscission, in parallel to the increase in sitosterol content. The differences found in AZ lipid content and the relationships established between LCB and sterol composition, offer new insights about sphingolipids and sterols in abscission.

Cell Wall Remodeling in Abscission Zone Cells during Ethylene-Promoted Fruit Abscission in Citrus.

Abscission is a cell separation process by which plants can shed organs such as fruits, leaves, or flowers. The process takes place in specific locations termed abscission zones. In fruit crops like citrus, fruit abscission represents a high percentage of annual yield losses. Thus, understanding the molecular regulation of abscission is of capital relevance to control production. To identify genes preferentially expressed within the citrus fruit abscission zone (AZ-C), we performed a comparative transcriptomics assay at the cell type resolution level between the AZ-C and adjacent fruit rind cells (non-abscising tissue) during ethylene-promoted abscission. Our strategy combined laser microdissection with microarray analysis. Cell wall modification-related gene families displayed prominent representation in the AZ-C. Phylogenetic analyses of such gene families revealed a link between phylogenetic proximity and expression pattern during abscission suggesting highly conserved roles for specific members of these families in abscission. Our transcriptomic data was validated with (and strongly supported by) a parallel approach consisting on anatomical, histochemical and biochemical analyses on the AZ-C during fruit abscission. Our work identifies genes potentially involved in organ abscission and provides relevant data for future biotechnology approaches aimed at controlling such crucial process for citrus yield.

The influence of abscisic acid on the ethylene biosynthesis pathway in the functioning of the flower abscission zone in Lupinus luteus

Flower abscission is a highly regulated developmental process activated in response to exogenous (e.g. changing environmental conditions) and endogenous stimuli (e.g. phytohormones). Ethylene (ET) and abscisic acid (ABA) are very effective stimulators of flower abortion in Lupinus luteus, which is a widely cultivated species in Poland, Australia and Mediterranean countries. In this paper, we show that artificial activation of abscission by flower removal caused an accumulation of ABA in the abscission zone (AZ). Moreover, the blocking of that phytohormone’s biosynthesis by NDGA (nordihydroguaiaretic acid) decreased the number of abscised flowers. However, the application of NBD – an inhibitor of ET action – reversed the stimulatory effect of ABA on flower abscission, indicating that ABA itself is not sufficient to turn on the organ separation. Our analysis revealed that exogenous ABA significantly accelerated the transcriptional activity of the ET biosynthesis genes ACC synthase (LlACS) and oxidase (LlACO), and moreover, strongly increased the level of 1-aminocyclopropane-1-carboxylic acid (ACC) – ET precursor, which was specifically localized within AZ cells. We cannot exclude the possibility that ABA mediates flower abscission processes by enhancing the ET biosynthesis rate. The findings of our study will contribute to the overall basic knowledge on the phytohormone-regulated generative organs abscission in L. luteus.

Immunolocalization of pectic polysaccharides during abscission in pea seeds (Pisum sativum L.) and in abscission less def pea mutant seeds.

BackgroundIn pea seeds (Pisum sativum L.), the presence of the Def locus determines abscission event between its funicle and the seed coat. Cell wall remodeling is a necessary condition for abscission of pea seed. The changes in cell wall components in wild type (WT) pea seed with Def loci showing seed abscission and in abscission less def mutant peas were studied to identify the factors determining abscission and non-abscission event.MethodsChanges in pectic polysaccharides components were investigated in WT and def mutant pea seeds using immunolabeling techniques. Pectic monoclonal antibodies (1 → 4)-β-d-galactan (LM5), (1 → 5)-α-l-arabinan(LM6), partially de-methyl esterified homogalacturonan (HG) (JIM5) and methyl esterified HG (JIM7) were used for this study.ResultsPrior to abscission zone (AZ) development, galactan and arabinan reduced in the predestined AZ of the pea seed and disappeared during the abscission process. The AZ cells had partially de-methyl esterified HG while other areas had highly methyl esterified HG. A strong JIM5 labeling in the def mutant may be related to cell wall rigidity in the mature def mutants. In addition, the appearance of pectic epitopes in two F3 populations resulting from cross between WT and def mutant parents was studied. As a result, we identified that homozygous dominant lines (Def/Def) showing abscission and homozygous recessive lines (def/def) showing non-abscission had similar immunolabeling pattern to their parents. However, the heterogeneous lines (Def/def) showed various immunolabeling pattern and the segregation pattern of the Def locus.ConclusionsThrough the study of the complexity and variability of pectins in plant cell walls as well as understanding the segregation patterns of the Def locus using immunolabeling techniques, we conclude that cell wall remodeling occurs in the abscission process and de-methyl esterification may play a role in the non-abscission event in def mutant. Overall, this study contributes new insights into understanding the structural and architectural organization of the cell walls during abscission.

Cell wall related gene expression during secondary physiological fruit drop in ponkan (Citrus reticulataBlanco) and hyuganatsu (C. tamuranahort. ex Tanaka)

Physiological fruit drop in citrus is the deeper problem for fruit production and many researches have suggested that decreasing of carbohydrate translocation to fruit involved in fruit abscission. The fruit abscission process requires 3 stages for successful abscission, which include “induction”, “decision”, and “execution”. However, there is almost no information in the molecular mechanism of secondary physiological fruit drop in citrus. Here, anatomical observation and microarray analysis demonstrated the candidate genes involved in the fruit abscission process after induction of fruit abscission by incubating the detached ponkan (Citrus reticulata Blanco) and hyuganatsu (C. tamurana hort. ex Tanaka) fruit on agar medium. In our study, the cumulative ratio of fruit abscission was very high in ponkan, but very low in hyuganatsu. Morphology of abscission zone (AZ) cells was drastically changed in ponkan, but almost not changed in hyuganatsu. From the 60 K microarray analysis at 0, 12, 24, or 36 h after the induction, the specific up-regulated genes in ponkan AZ tissue were identified. The expressions of many cell wall degrading related genes, such as beta-galactosidase, polygalacturonase and beta-glucosidase etc. increased rapidly at 24 h. These results indicated that “execution” stage of fruit abscission process would be initiated until 24 h after the induction of fruit abscission in citrus.

Cellular and Pectin Dynamics during Abscission Zone Development and Ripe Fruit Abscission of the Monocot Oil Palm.

The oil palm (Elaeis guineensis Jacq.) fruit primary abscission zone (AZ) is a multi-cell layered boundary region between the pedicel (P) and mesocarp (M) tissues. To examine the cellular processes that occur during the development and function of the AZ cell layers, we employed multiple histological and immunohistochemical methods combined with confocal, electron and Fourier-transform infrared (FT-IR) microspectroscopy approaches. During early fruit development and differentiation of the AZ, the orientation of cell divisions in the AZ was periclinal compared with anticlinal divisions in the P and M. AZ cell wall width increased earlier during development suggesting cell wall assembly occurred more rapidly in the AZ than the adjacent P and M tissues. The developing fruit AZ contain numerous intra-AZ cell layer plasmodesmata (PD), but very few inter-AZ cell layer PD. In the AZ of ripening fruit, PD were less frequent, wider, and mainly intra-AZ cell layer localized. Furthermore, DAPI staining revealed nuclei are located adjacent to PD and are remarkably aligned within AZ layer cells, and remain aligned and intact after cell separation. The polarized accumulation of ribosomes, rough endoplasmic reticulum, mitochondria, and vesicles suggested active secretion at the tip of AZ cells occurred during development which may contribute to the striated cell wall patterns in the AZ cell layers. AZ cells accumulated intracellular pectin during development, which appear to be released and/or degraded during cell separation. The signal for the JIM5 epitope, that recognizes low methylesterified and un-methylesterified homogalacturonan (HG), increased in the AZ layer cell walls prior to separation and dramatically increased on the separated AZ cell surfaces. Finally, FT-IR microspectroscopy analysis indicated a decrease in methylesterified HG occurred in AZ cell walls during separation, which may partially explain an increase in the JIM5 epitope signal. The results obtained through a multi-imaging approach allow an integrated view of the dynamic developmental processes that occur in a multi-layered boundary AZ and provide evidence for distinct regulatory mechanisms that underlie oil palm fruit AZ development and function.

Reactive oxygen species regulate leaf pulvinus abscission zone cell separation in response to water-deficit stress in cassava.

Cassava (Manihot esculenta Crantz) plant resists water-deficit stress by shedding leaves leading to adaptive water-deficit condition. Transcriptomic, physiological, cellular, molecular, metabolic, and transgenic methods were used to study the mechanism of cassava abscission zone (AZ) cell separation under water-deficit stress. Microscopic observation indicated that AZ cell separation initiated at the later stages during water-deficit stress. Transcriptome profiling of AZ suggested that differential expression genes of AZ under stress mainly participate in reactive oxygen species (ROS) pathway. The key genes involved in hydrogen peroxide biosynthesis and metabolism showed significantly higher expression levels in AZ than non-separating tissues adjacent to the AZ under stress. Significantly higher levels of hydrogen peroxide correlated with hydrogen peroxide biosynthesis related genes and AZ cell separation was detected by microscopic observation, colorimetric detection and GC-MS analyses under stress. Co-overexpression of the ROS-scavenging proteins SOD and CAT1 in cassava decreased the levels of hydrogen peroxide in AZ under water-deficit stress. The cell separation of the pulvinus AZ also delayed in co-overexpression of the ROS-scavenging proteins SOD and CAT1 plants both in vitro and at the plant level. Together, the results indicated that ROS play an important regulatory role in the process of cassava leaf abscission under water-deficit stress.

The IDA Peptide Controls Abscission in Arabidopsis and Citrus.

Organ abscission is an important process in plant development and reproduction. During abscission, changes in cellular adhesion of specialized abscission zone cells ensure the detachment of infected organs or those no longer serving a function to the plant. In addition, abscission also plays an important role in the release of ripe fruits. Different plant species display distinct patterns and timing of organ shedding, most likely adapted during evolution to their diverse life styles. However, it appears that key regulators of cell separation may have conserved function in different plant species. Here, we investigate the functional conservation of the citrus ortholog of the Arabidopsis peptide ligand INFLORESCENCE DEFICIENT IN ABSCISSION (AtIDA), controlling floral organ abscission. We discuss the possible implications of modifying the citrus IDA ortholog for citrus fruit production.

Profiling the BLADE-ON-PETIOLE gene expression in the abscission zone of generative organs in Lupinus luteus

The great agronomic potential of Lupinus luteus, species widely cultivated in many European countries as well as Australia, is strongly affected by premature and excessive generative organ abscission, mainly flowers. The unwanted process takes place in a specialized group of cells, called abscission zone (AZ). During their development they become competent to respond to external and internal factors, including phytohormones. Recently it was shown that the formation of AZ cells in Arabidopsis thaliana is coordinated by transcription factors, BLADE-ON-PETIOLE (BOPs). There is no such data, excluding tobacco plants, about BOP-dependent regulation of organ abscission in crop plants. In this work, we examined LlBOP mRNA content during generative organs AZ development and functioning. The high accumulation of LlBOP transcript was accompanied by the differentiation of morphologically distinct cells at the base of the mature flower pedicel. Moreover, enhanced LlBOP expression was observed in the active AZ, and was regulated by factors, which can strongly affect generative organ abscission. All these data indicate that LlBOP is involved in the abscission zone formation and functioning in L. luteus.

Beyond the meristems: similarities in the CLAVATA3 and INFLORESCENCE DEFICIENT IN ABSCISSION peptide mediated signalling pathways.

Plants form new organs throughout their lives; this requires a balance between cell proliferation and differentiation, and between the generation and loss of organs. To do this, plants must maintain a population of stem cells within the meristems, and at the same time, closely control the identity and position of cells at the meristem boundaries as they differentiate to new leaf or flower primordia. Once developed, organs may need to be shed, either as a controlled developmental decision-such as floral abscission after pollination, or as a response to disease, environmental stress, and predators. Cell wall degradation at specialized abscission zone (AZ) cells needs to occur for this to take place, but since there is little cell rearrangement in plants, cell separation events are also important for plant architecture. In this Opinion paper we discuss the role of two peptide ligand signalling systems that control stem cell homeostasis and cell separation, respectively. We draw parallels between the signalling pathways and explore on the commonalities of the downstream components activated and controlled by the signalling peptides. We provide evidence for AZ cells having a meristem identity and discuss the role of identical KNOTTED-LIKE HOMEOBOX (KNOX) transcription factors in meristem maintenance and abscission. Lastly we explore the evolutionary relationship between the pathways.

Burst of reactive oxygen species in pedicel-mediated fruit abscission after carbohydrate supply was cut off in longan (Dimocarpus longan)

Cutting off carbohydrate supply to longan (Dimocarpus longan Lour.) fruit by girdling and defoliation or by detachment induced 100% abscission within a few days. We used these treatments to study the involvement of reactive oxygen species (ROS) in fruit abscission. Girdling plus defoliation decreased sugar concentrations in the fruit and pedicel and depleted starch grains in the chloroplasts in the cells of abscission zone. Prior to the occurrence of intensive fruit abscission, there was a burst in ROS in the pedicel, which peaked at 1 day after treatment (DAT), when H2O2 in the abscission zone was found to be chiefly located along the plasma membrane (PM). H2O2 was found exclusively in the cell walls 2 DAT, almost disappeared 3 DAT, and reappeared in the mitochondria and cell walls 4 DAT. Signs of cell death such as cytoplasm breakdown were apparent from 3 DAT. The burst of ROS coincided with a sharp increase in the activity of PM-bound NADPH oxidase in the pedicel. At the same time, activities of antioxidant enzymes including superoxide dismutase (SOD), catalase, and peroxidase (POD) were all increased by the treatment and maintained higher than those in the control. Accompanying the reduction in H2O2 abundance, there was a sharp decrease in PM-bound NADPH oxidase activity after 1 DAT in the treated fruit. H2O2 scavenger dimethylthiourea (DMTU, 1 g L–1) significantly inhibited fruit abscission in detached fruit clusters and suppressed the increase in cellulase activity in the abscission zone. These results suggest that fruit abscission induced by carbohydrate stress is mediated by ROS. Roles of ROS in regulating fruit abscission were discussed in relation to its subcellular distribution.