A substantial body of literature supports the involvement of ethylene in abscission of leaves, flowers and flower parts, and fruit. Exogenous ethylene accelerates abscission in many, but not all, abscising plant systems. In some cases, endogenous ethylene production increases before abscission, while in others a change in ethylene sensitivity can be demonstrated. Ethylene interacts with auxin in regulating abscission, particularly in leaves and fruit. Ethylene has been shown to regulate expression of genes involved in cell separation, including cellulases and pectinases. Such regulation has been demonstrated at the level of mRNA abundance, but post-transcriptional regulation may also be important.

Ethylene and abscission

The reproductive biology of the lychee

The high level of physiological fruitlet abscission in litchi (Litchi chinensis Sonn.) causes severe yield loss. Cell separation occurs at the fruit abscission zone (FAZ) and can be triggered by ethylene. However, a deep knowledge of the molecular events occurring in the FAZ is still unknown. Here, genome-wide digital transcript abundance (DTA) analysis of putative fruit abscission related genes regulated by ethephon in litchi were studied. More than 81 million high quality reads from seven ethephon treated and untreated control libraries were obtained by high-throughput sequencing. Through DTA profile analysis in combination with Gene Ontology and KEGG pathway enrichment analyses, a total of 2,730 statistically significant candidate genes were involved in the ethephon-promoted litchi fruitlet abscission. Of these, there were 1,867 early-responsive genes whose expressions were up- or down-regulated from 0 to 1 d after treatment. The most affected genes included those related to ethylene biosynthesis and signaling, auxin transport and signaling, transcription factors, protein ubiquitination, ROS response, calcium signal transduction and cell wall modification. These genes could be clustered into 4 groups and 13 subgroups according to their similar expression patterns. qRT-PCR displayed the expression pattern of 41 selected candidate genes, which proved the accuracy of our DTA data. Ethephon treatment significantly increased fruit abscission and ethylene production of fruitlet. The possible molecular events to control the ethephon-promoted litchi fruitlet abscission were prompted out. The increased ethylene evolution in fruitlet would suppress the synthesis and polar transport of auxin and trigger abscission signaling. To the best of our knowledge, it is the first time to monitor the gene expression profile occurring in the FAZ-enriched pedicel during litchi fruit abscission induced by ethephon on the genome-wide level. This study will contribute to a better understan

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Genome-wide digital transcript analysis of putative fruitlet abscission related genes regulated by ethephon in litchi.

Auxin, a vital plant hormone, regulates a variety of physiological and developmental processes. It is involved in fruit abscission through transcriptional regulation of many auxin-related genes, including early auxin responsive genes (i.e., auxin/indole-3-acetic acid (AUX/IAA), Gretchen Hagen3 (GH3) and small auxin upregulated (SAUR)) and auxin response factors (ARF), which have been well characterized in many plants. In this study, totally five auxin-related genes, including one AUX/IAA (LcAUX/IAA1), one GH3 (LcGH3.1), one SAUR (LcSAUR1) and two ARFs (LcARF1 and LcARF2), were isolated and characterized from litchi fruit. LcAUX/IAA1, LcGH3.1, LcSAUR1, LcARF1 and LcARF2 contain open reading frames (ORFs) encoding polypeptides of 203, 613, 142, 792 and 832 amino acids, respectively, with their corresponding molecular weights of 22.67, 69.20, 11.40, 88.20 and 93.16 kDa. Expression of these genes was investigated under the treatment of girdling plus defoliation which aggravated litchi fruitlet abscission due to the blockage of carbohydrates transport and the reduction of endogenous IAA content. Results showed that transcript levels of LcAUX/IAA1, LcGH3.1 and LcSAUR1 mRNAs were increased after the treatment in abscission zone (AZ) and other tissues, in contrast to the decreasing accumulation of LcARF1 mRNA, suggesting that LcAUX/IAA1, LcSAUR1 and LcARF1 may play more important roles in abscission. Our results provide new insight into the process of fruitlet abscission induced by carbohydrate stress and broaden our understanding of the auxin signal transduction pathway in this process at the molecular level.

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Carbohydrate Stress Affecting Fruitlet Abscission and Expression of Genes Related to Auxin Signal Transduction Pathway in Litchi

Massive young fruit abscission usually causes low and unstable yield in litchi (Litchi chinensis Sonn.), an important fruit crop cultivated in tropical and subtropical areas. However, the molecular mechanism of fruit drop has not been fully characterized. This study aimed at identification of molecular components involved in fruitlet abscission in litchi, for which reference genome is not available at present. An improved de novo transcriptome assembly was firstly achieved by using an optimized assembly software, Trinity. Using improved transcriptome assembly as reference, digital transcript abundance (DTA) profiling was performed to screen and identify candidate genes involved in fruit abscission induced by girdling plus defoliation (GPD), a treatment significantly decreased the soluble sugar contents causing carbohydrate stress to fruit. Our results showed that the increasing fruit abscission rate after GPD treatment was associated with higher ethylene production and lower glucose levels in fruit. A total of 2,771 differentially expressed genes were identified as GPD-responsive genes, 857 of which were defined by GO and KEGG enrichment analyses as the candidate genes involved in fruit abscission process. These genes were involved in diverse metabolic processes and pathways, including carbohydrate metabolism, plant hormone synthesis and signaling, transcription factor activity and cell wall modification that were rapidly induced in the early stages (within 2 d after treatment). qRT-PCR was used to explore the expression pattern of 15 selected candidate genes in the abscission zone, pericarp and seed, which confirmed the accuracy of our DTA data. More detailed information for different functional categories was also analyzed. This study profiled the gene expression related to fruit abscission induced by carbohydrate stress at whole transcriptome level and thus provided a better understanding of the regulatory mechanism of young fruit abscission in litchi.

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An improved fruit transcriptome and the identification of the candidate genes involved in fruit abscission induced by carbohydrate stress in litchi

Fruit development and abscission in 'Mauritius' lychee (Litchi chinensis Sonn.) were studied over three consecutive seasons. Each season, two distinct abscission periods were observed. The first started at the end of full female bloom and continued for ≈ 4 weeks. Of the initial number of female flowers, 85 % to 90 % abscised during this period. The second period began after a lag period of ≈ ≈ 1 week and lasted ≈ 2 weeks. About half of the remaining fruitlets abscised during this wave. AU of these fruitlets contained an embryo. The second wave coincided with a period of rapid embryo growth and endosperm loss. Tipimon (a commercial product containing the triethanolamine salt of the synthetic auxin 2,4,5-TP) consistently and significantly increased marketable fruit yield when applied between the two abscission periods. Chemical name used: 2,4,5 -trichlorophenoxy propionic acid (2,4,5 -TP).

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R.A. Stern

Papers

`Mauritius' Lychee Fruit Development and Reduced Abscission after Treatment with the Auxin 2,4,5-TP