Frontiers of Agricultural Science and Engineering
Cover Story   2021, Volume 8 Issue 2
As indicated by the overarching theme of Beijing’s Horticultural Exhibition “Live Green, Live Better”, horticulture contributes to promote respect for nature and a better life in harmony with nature. Horticultural industry comprises a wide range of plants and crops including fruit and nut trees, vegetables, edible fungi, and ornamental plants, as w [Detail] ...
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, Volume 8 Issue 2

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EDITORIAL
HIGHLIGHTS OF THE SPECIAL ISSUE ON “HORTICULTURE RESEARCH FOR GREEN AND SUSTAINABLE DEVELOPMENT”
Xiuxin DENG, Yujin HAO, Jingquan YU, Qixiang ZHANG, Zhonghua LIU
Front. Agr. Sci. Eng. . 2021, 8 (2): 193-195.  
https://doi.org/10.15302/J-FASE-2021396

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RESEARCH ARTICLE
BULKED SEGREGANT RNA SEQUENCING (BSR-SEQ) IDENTIFIES A NOVEL ALLELE ASSOCIATED WITH WEEPING TRAITS IN PRUNUS MUME
Xiaokang ZHUO, Tangchun ZHENG, Zhiyong ZHANG, Suzhen LI, Yichi ZHANG, Lidan SUN, Weiru YANG, Jia WANG, Tangren CHENG, Qixiang ZHANG
Front. Agr. Sci. Eng. . 2021, 8 (2): 196-214.  
https://doi.org/10.15302/J-FASE-2020379

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• Five QTLs associated with weeping traits on chromosome 7 were identified by BSR-seq.

• The novel allele PmUGT72B3 has a synonymous transition of T66 (upright) to C (weeping) in the coding sequence and a 470-bp deletion in the promoter region.

PmUGT72B3 was associated with hormone and lignin regulation by WGCNA.

Weeping species are used both as ornamental plants and for breeding dwarf plant types. However, exploration of casual genes controlling weeping traits is rather limited. Here, we identified individuals with contrasting phenotypes from an F1 bi-parental mapping population of Prunus mume which was developed from a cross between the upright cultivar ‘Liuban’ and the weeping cultivar ‘Fentai Chuizhi’. Bulked segregant RNA sequencing was used and five QTLs on Chromosome 7 were identified. The Pm024074 (PmUGT72B3) allele, belonging to the UDP-glycosyltransferase superfamily containing the coniferyl-alcohol glucosyltransferase domain, was identified in a genomic region overlapping with a previously identified QTL, and had a synonymous transition of T66 (upright) to C (weeping) in the coding sequence and a 470-bp deletion in the promoter region. Pm024074 had exceptionally high expression in buds and stems of weeping P. mume. Weighted correlation network analysis indicates that genes neighboring Pm024074 were significantly associated with plant architecture. In addition, a reliable single nucleotide polymorphism marker was developed based on the variation in the Pm024074 gene, providing precise marker-assisted breeding for weeping traits. This study provides insights into the genetic mechanism governing the weeping trait in P. mume, and indicates potential applications for the manipulation of tree architecture.

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METABOLIC AND TRANSCRIPTOME ANALYSIS REVEALS METABOLITE VARIATION AND FLAVONOID REGULATORY NETWORKS IN FRESH SHOOTS OF TEA (CAMELLIA SINENSIS) OVER THREE SEASONS
Chen-Kai JIANG, De-Jiang NI, Ming-Zhe YAO, Jian-Qiang MA, Liang CHEN
Front. Agr. Sci. Eng. . 2021, 8 (2): 215-230.  
https://doi.org/10.15302/J-FASE-2021382

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• Metabolites of fresh tea shoots at harvest were profiled.

• Season-dependent metabolites were identified.

• Key genes responsible for flavonoid metabolism are proposed.

• Regulated relationships among the main compounds were investigated.

Metabolites, especially secondary metabolites, are very important in the adaption of tea plants and the quality of tea products. Here, we focus on the seasonal variation in metabolites of fresh tea shoots and their regulatory mechanism at the transcriptional level. The metabolic profiles of fresh tea shoots of 10 tea accessions collected in spring, summer, and autumn were analyzed using ultra-performance liquid chromatography coupled with quadrupole-obitrap mass spectrometry. We focused on the metabolites and key genes in the phenylpropanoid/flavonoid pathway integrated with transcriptome analysis. Multivariate statistical analysis indicates that metabolites were distinctly different with seasonal alternation. Flavonoids, amino acids, organic acids and alkaloids were the predominant metabolites. Levels of most key genes and downstream compounds in the flavonoid pathway were lowest in spring but the catechin quality index was highest in spring. The regulatory pathway was explored by constructing a metabolite correlation network and a weighted gene co-expression network.

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LETTER
THE ANTHOCYANIN BIOSYNTHETIC REGULATOR MDMYB1 POSITIVELY REGULATES ASCORBIC ACID BIOSYNTHESIS IN APPLE
Jianping AN, Xiaofei WANG, Chunxiang YOU, Yujin HAO
Front. Agr. Sci. Eng. . 2021, 8 (2): 231-235.  
https://doi.org/10.15302/J-FASE-2020367

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• The contents of anthocyanin and AsA in red-flesh apples are higher than that in non-red-flesh apples.

• The anthocyanin biosynthetic regulator MdMYB1 directly activates the expression of dehydroascorbate reductase gene MdDHAR, thus promoting the activity of the DHAR enzyme and the accumulation of AsA.

• MdMYB1-MdDHAR module may play a key role in AsA-DHA homeostasis.

Ascorbic acid (AsA, vitamin C) is involved in the regulation of many aspects of plant growth and development. It is an essential micronutrient for humans and can prevent scurvy, maintain the health of gums and blood vessels, reduce the level of plasma cholesterol and enhance the immune systen. Apple cultivars Orin and Guanghui were crossed to obtain a group of hybrid offspring with and without red flesh in the course of assessing apple germplasm resources. Unexpectedly, the red-flesh apples had higher AsA contents than other apples. Further studies showed that the anthocyanin biosynthetic regulator MdMYB1 directly activates the expression of dehydroascorbate reductase gene MdDHAR, thus promoting the activity of the DHAR enzyme and the accumulation of AsA. This finding reveals the mechanism leading to high AsA levels in red-flesh apples and suggests a new idea to cultivate red-flesh apples with high AsA contents and produce AsA efficiently and without pollution.

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RESEARCH ARTICLE
OVEREXPRESSION OF PTRLEA7, A LATE EMBRYOGENESIS ABUNDANT FAMILY GENE FROM PONCIRUS TRIFOLIATA, CONFERS ENHANCED DROUGHT TOLERANCE BY ENHANCING ANTIOXIDANT CAPACITY
Tonglu WEI, Dalong GUO, Jihong LIU
Front. Agr. Sci. Eng. . 2021, 8 (2): 236-246.  
https://doi.org/10.15302/J-FASE-2020368

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• A LEA family gene (PtrLEA7)was cloned from Poncirus trifoliata.

PtrLEA7was strongly induced by stresses and ABA.

PtrLEA7played a positive role in modulation of drought tolerance.

• Overexpression of PtrLEA7elevated antioxidant capacity.

Late embryogenesis abundant (LEA) genes encode highly hydrophilic proteins that are essential in abiotic stress responses. However, most LEA genes in higher plants have not yet been investigated. This study identified an LEA family gene (PtrLEA7) from Poncirus trifoliata and studied its function in drought tolerance. The full-length coding sequence of PtrLEA7 was 420 bp encoding a protein of 139 amino acids. Phylogenetic analysis shows that PtrLEA7 protein belongs to the LEA_4 subfamily. Expression profiling by qPCR found that PtrLEA7 was strongly induced by dehydration, cold and ABA treatments, and slightly induced by salt stress. Subcellular localization reveals that PtrLEA7 protein was located in both cytoplasm and nucleus. To investigate its function, transgenic plants of both tobacco and Poncirus trifoliata overexpressing PtrLEA7 were obtained. Stress tolerance assays show that overexpression lines had enhanced dehydration and drought tolerance compared with wild type plants, indicating that PtrLEA7 positively regulates drought tolerance. In addition, transgenic plants had much higher expression levels of three antioxidant enzyme genes (CAT, SOD and POD) and significantly increased catalase enzyme activity, accompanied by reduced reactive oxygen species accumulation in comparison with wild type plants. Collectively, this study demonstrates that PtrLEA7 can confer enhanced drought tolerance partially via enhancing antioxidant capacity.

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APPLE SUMO E3 LIGASE MDSIZ1 NEGATIVELY REGULATES DROUGHT TOLERANCE
Baohua CHU, Jia SUN, Huan DANG, Ziqing MA, Shuang ZHAO, Qingmei GUAN, Xuewei LI
Front. Agr. Sci. Eng. . 2021, 8 (2): 247-261.  
https://doi.org/10.15302/J-FASE-2021388

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MdSIZ1 RNAi transgenic apple trees are drought tolerance than wild type—GL-3.

MdSIZ1 RNAi plants get enhanced ability to keep water and scavenge ROS under drought conditions.

•MdSIZ1 may participate in apple drought tolerance by affecting ABA biosynthesis.

Drought stress typically causes heavy losses in apple production and uncovering the mechanisms by which apple tolerates drought stress is important in apple breeding. MdSIZ1 is a SUMO (small ubiquitin-like modifier) E3 ligase that promotes SUMO binding to substrate proteins. Here, we demonstrate that MdSIZ1 in apple has a negative relationship with drought tolerance. MdSIZ1 RNAi transgenic apple trees had a higher survival rate after drought stress. During drought stress they had higher leaf water potential, reduced ion leakage, lower H2O2 and malondialdehyde contents, and higher catalase activity. In addition, MdSIZ1 RNAi transgenic plants had a higher net photosynthetic rate during the latter period of drought stress. Finally, the transgenic apple trees also altered expression levels of some microRNAs in response to drought stress. Taken together, these results indicate that apple MdSIZ1 negatively regulates drought stress by enhancing leaf water-holding capacity and antioxidant enzyme activity.

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PROTECTIVE ROLES OF D1 PROTEIN TURNOVER AND THE XANTHOPHYLL CYCLE IN TOMATO (SOLANUM LYCOPERSICUM) UNDER SUB-HIGH TEMPERATURE AND HIGH LIGHT STRESS
Tao LU, Jiazhi LU, Mingfang QI, Zhouping SUN, Yufeng LIU, Tianlai LI
Front. Agr. Sci. Eng. . 2021, 8 (2): 262-279.  
https://doi.org/10.15302/J-FASE-2021383

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• D1 turnover plays a more important role than xanthophyll cycle in photoprotection under sub-high temperature and high light (HH) conditions in tomato.

• D1 protein turnover and xanthophyll cycle avoided the accumulation of excess energy and photooxidative damage of photosystem through maintaining photosynthetic pigments content and inducing changes in NPQ core components.

• D1 protein turnover and xanthophyll cycle can also avoided accumulation of ROS and membrane lipid peroxidation by maintaining the activity of the ROS scavenging system.

D1 protein turnover and the xanthophyll cycle (XC) are important photoprotective mechanisms in plants that operate under adverse conditions. Here, streptomycin sulfate (SM) and dithiothreitol (DTT) were used in tomato plants as inhibitors of D1 protein turnover and XC to elucidate their photoprotective impacts under sub-high temperature and high light conditions (HH, 35°C, 1000 µmol·m-2·s-1). SM and DTT treatments significantly reduced the net photosynthetic rate, apparent quantum efficiency, maximum photochemical efficiency, and potential activity of photosystem II, leading to photoinhibition and a decline in plant biomass under HH. The increase in reactive oxygen species levels resulted in thylakoid membrane lipid peroxidation. In addition, there were increased non-photochemical quenching and decreased chlorophyll pigments in SM and DTT application, causing an inhibition of D1 protein production at both transcriptional and translational levels. Overall, inhibition of D1 turnover caused greater photoinhibition than XC inhibition. Additionally, the recovery levels of most photosynthesis indicators in DTT-treated plants were higher than in SM-treated plants. These findings support the view that D1 turnover has a more important role than XC in photoprotection in tomato under HH conditions.

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FUNCTIONAL GAIN OF FRUIT NETTED-CRACKING IN AN INTROGRESSION LINE OF TOMATO WITH HIGHER EXPRESSION OF THE FNC GENE
Chunli ZHANG, Taotao WANG, Jing LI, Danqiu ZHANG, Qingmin XIE, Shoaib MUNIR, Jie YE, Hanxia LI, Yongen LU, Changxian YANG, Bo OUYANG, Yuyang ZHANG, Junhong ZHANG, Zhibiao YE
Front. Agr. Sci. Eng. . 2021, 8 (2): 280-291.  
https://doi.org/10.15302/J-FASE-2020374

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• A novel netted-cracking fruit phenotype was discovered in tomato introgression line IL4-4.

• A single dominant gene (FNC) determined the fruit netted-cracking phenotype.

• The high transcript level of FNC results in the functional gain of fruit netted-cracking and it was found to be a common mechanism in a diverse range of plant species.

Fruit cracking is a major disorder that affects the integrity of fruit and reduces the commercial value of tomato and other fleshy fruit. Here, we have found a novel fruit ‘netted-cracking’ (FNC) phenotype in tomato introgression line IL4-4 which is present in neither the donor parent (LA0716) nor the receptor parent (M82). An F2 population was generated by crossing IL4-4 with M82 to genetically characterize the FNC gene and this showed that a single dominant gene determined fruit netted-cracking. Further map-based cloning narrowed down the FNC locus to a 230 kb region on chromosome 4. Sequencing and annotation analysis show that FNC (Solyc04 g082540) was the most likely candidate gene. Functional characterization of FNC by overexpressing FNCAC and FNCIL4-4 resulted in the fruit netted-cracking phenotype, suggesting that the FNC transcript level results in the functional gain of fruit netted-cracking. These findings were further confirmed by FNC ortholog in netted-cracking pepper and melon, indicating a common regulatory mechanism in different plant species. Furthermore, cytoplasm and nucleus-localized FNC indicates increased expression of genes involved in suberin, lignin, lipid transport and cell wall metabolism. These findings provide novel genetic insights into fruit netted-cracking and offer a way to promote molecular improvement toward cracking resistant cultivars.

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CHARACTERISTICS OF HERBIVORY/WOUND-ELICITED ELECTRICAL SIGNAL TRANSDUCTION IN TOMATO
Chaoyi HU, Siqi DUAN, Jie ZHOU, Jingquan YU
Front. Agr. Sci. Eng. . 2021, 8 (2): 292-301.  
https://doi.org/10.15302/J-FASE-2021395

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• Herbivory and mechanical wounding elicited electrical signals.

• Petiole wounding elicited stronger electrical signals than did leaflet wounding.

• Leaflet wounding elicited electrical signals and JA signaling within a compound leaf.

• GLR3.3 and GLR3.5 mediated leaflet-to-leaflet electrical signal transduction.

• JA synthesis and Helicoverpa armigera resistance were reduced in glr3.3/3.5 plants.

Electrical signals commonly occur in plants in response to various environmental changes and have a dominant function in plant acclimation. The transduction of wound-elicited electrical signals in the model plant species Arabidopsis has been characterized but the characteristics of electrical signal transduction in response to herbivory or wounding in crop species remain unknown. Here, the features of electrical signals elicited by insect herbivory and wounding in tomato were investigated. Unlike those in Arabidopsis, wounding tomato leaves did not cause leaf-to-leaf electrical signal transduction. In contrast, electrical signals elicited in response to petiole wounding were stronger and more strongly transduced. Leaflet wounding also activated electrical signal transduction and jasmonic acid (JA) signaling within the whole compound leaf. It was also demonstrated that tomato glutamate receptor-like 3.3 (GLR3.3) and GLR3.5 mediated leaflet-to-leaflet electrical signal transduction. Herbivory-induced JA accumulation and Helicoverpa armigera resistance were reduced in glr3.3/3.5 plants. This work reveals the nature of electrical signal transduction in tomato and emphasizes the key roles of GLR3.3 and GLR3.5 in electrical signal transduction and JA signaling activation.

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UNREDUCED MEGAGAMETOPHYTE FORMATION VIA SECOND DIVISION RESTITUTION CONTRIBUTES TO TETRAPLOID PRODUCTION IN INTERPLOIDY CROSSES WITH ‘ORAH’ MANDARIN (CITRUS RETICULATA)
Qiangming XIA, Wei WANG, Kaidong XIE, Xiaomeng WU, Xiuxin DENG, Jude W. GROSSER, Wenwu GUO
Front. Agr. Sci. Eng. . 2021, 8 (2): 302-313.  
https://doi.org/10.15302/J-FASE-2021385

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• In addition to triploid progeny, tetraploid hybrids derived from the fertilization of 2n megagametophytes are frequently regenerated from 2x × 4x crosses that utilize ‘Orah’ mandarin as the female parent.

• Data here indicate that ‘Orah’ mandarin is a cultivar that readily produces 2n megagametophytes.

• Second division restitution is the mechanism underlying 2n megagametophyte formation in ‘Orah’ mandarin.

Seedless fruits are desirable in the citrus fresh fruit market. Triploid production via diploid × tetraploid interploidy crosses is thought to be the most efficient and widely-used strategy for the breeding of seedless citrus. Although ‘Orah’ mandarin has desirable organoleptic qualities, seeds in the fruits weaken its market competitiveness. To produce new seedless cultivars that are similar to ‘Orah’ mandarin, we performed three 2x × 4x crosses using ‘Orah’ mandarin as the seed parent to regenerate triploid plantlets. A total of 182 triploid and 36 tetraploid plantlets were obtained. By analyzing their genetic origins using nine novel single nucleotide polymorphism (SNP) markers, all of the triploids and tetraploids derived from these three crosses were proven to be hybrids. Also, we demonstrated that 2n megagametophyte formation in ‘Orah’ mandarin result in tetraploid production in these three interploidy crosses. These tetraploid plantlets were genotyped using eight pericentromeric SNP markers and nine centromere distal SNP markers. Based on the genotypes of the 2n megagametophytes, the parental heterozygosity rates in 16 SNP loci and all 2n megagametophytes were less than 50%, indicating that second division restitution was the mechanism underlying 2n megagametophyte formation at both the population and individual levels. These triploid hybrids enrich the germplasm available for seedless breeding. Moreover, the tetraploid hybrids are valuable as parents for ploidy breeding for the production of seedless citrus fruits.

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REVIEW
RECENT ADVANCES IN THE REGULATION OF CLIMACTERIC FRUIT RIPENING: HORMONE, TRANSCRIPTION FACTOR AND EPIGENETIC MODIFICATIONS
Yinglin JI, Mingyang XU, Aide WANG
Front. Agr. Sci. Eng. . 2021, 8 (2): 314-334.  
https://doi.org/10.15302/J-FASE-2021386

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• The dynamic interplay between phytohormones plays an important part in climacteric fruit ripening.

• Transcription factors are critical for the regulation of climacteric fruit ripening.

• Epigenetic modifications act as important regulators of fruit ripening.

Fruit ripening is a complex developmental process made up of genetically programmed physiological and biochemical activities. It culminates in desirable changes in the structural and textural properties and is governed by a complex regulatory network. Much is known about ethylene, one of the most important metabolites promoting the ripening of climacteric fruits. However, the dynamic interplay between phytohormones also plays an important part. Additional regulatory factors such as transcription factors (TFs) and epigenetic modifications also play vital role in the regulation of climacteric fruit ripening. Here, we review and evaluate the complex regulatory network comprising interactions between hormones and the action of TFs and epigenetic modifications during climacteric fruit ripening.

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COMPARISON OF POMELO (CITRUS MAXIMA) GROWN IN CHINA AND THAILAND
Warangkana MAKKUMRAI, Yue HUANG, Qiang XU
Front. Agr. Sci. Eng. . 2021, 8 (2): 335-352.  
https://doi.org/10.15302/J-FASE-2021391

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• China is the largest producer of pomelo globally.

• Chinese pomelos are adapted to subtropical climates and Thai pomelos to tropical climates.

• Guanxi pomelo is a popular cultivar in China and Thong Dee is the most popular in Thailand.

• Naringin is the most abundant flavonoid in Chinese and Thai pomelos.

• Fruity, sweet, sour, juicy and overall flavor attributes are important in consumer preference.

Pomelo is a member of the genus Citrus that is a key contributor to the breeding of modern citrus cultivars. China is the largest producer of pomelo and one of the top five pomelo exporting countries. Pomelos from Thailand are also well-known for their excellent quality and flavor and are ranked in the top ten export countries. This review introduces pomelo planting locations and conditions in China and Thailand. The characteristics and qualities of some commercial pomelo cultivars in China and Thailand are summarized to introduce them to international consumers and to document their similarities and dissimilarities. Data on bioactive compounds and antioxidant capacity are also included for most Chinese and Thai pomelos to highlight how they differ in this aspect because consumers are increasingly interested in healthier foods. In addition, the sensory perception in terms of aroma, flavor, texture and taste attributes and consumer perspective and preferences are discussed.

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PERSPECTIVE
GLOBAL GENOMIC PREDICTION IN HORTICULTURAL CROPS: PROMISES, PROGRESS, CHALLENGES AND OUTLOOK
Craig HARDNER, Satish KUMAR, Dorrie MAIN, Cameron PEACE
Front. Agr. Sci. Eng. . 2021, 8 (2): 353-355.  
https://doi.org/10.15302/J-FASE-2021387

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COMMENTS
A MYB-INDUCED MECHANISM FOR ENHANCED ASCORBATE IN RED-FLESHED APPLES
Richard V. ESPLEY, Kui LIN-WANG
Front. Agr. Sci. Eng. . 2021, 8 (2): 356-357.  
https://doi.org/10.15302/J-FASE-2021381

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The results presented in “The anthocyanin biosynthetic regulator MdMYB1 positively regulates ascorbic acid biosynthesis in apple” (An et al., this issue) provide evidence for a new mechanism for the elevation of ascorbate concentration in apple. Using a red-fleshed apple breeding population, the authors show how the anthocyanin-regulating MYB transcription factor, MdMYB1, also increases ascorbate concentrations by directly activating transcription of the dehydroascorbate reductase gene MdDHAR. This gene recycles oxidized ascorbate back to ascorbate, leading to elevated concentrations of vitamin C. These red-fleshed apples have enhanced concentrations of both anthocyanins and ascorbate, both of which are appealing traits for the development of healthier apples.

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