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Sirtuins (SRTs) are a group of nicotinamide adenine dinucleotide (NAD+)‐dependent deacetylase that target both histone and nonhistone proteins. The biological function of SRT in horticultural plants has been rarely studied. In this study, FaSRT1‐2 was identified as a key member of the 8 FaSRTs encoded in cultivated strawberry genome. Transient overexpression of FaSRT1‐2 in strawberry fruit accelerated ripening, increased the content of anthocyanins and sugars, enhanced ripening‐related gene expression. Moreover, stable transformation of FaSRT1‐2 in strawberry plants resulted in enhanced vegetative growth, increased sensitivity to heat stress and increased susceptibility to Botrytis cinerea infection. Interestingly, knocking out the homologous gene in woodland strawberry had the opposite effects. Additionally, we found the content of stress‐related hormone abscisic acid (ABA) was decreased, while the growth‐related gibberellin (GA) concentration was increased in FaSRT1‐2 overexpression lines. Gene expression analysis revealed induction of heat shock proteins, transcription factors, stress‐related and antioxidant genes in the FaSRT1‐2‐overexpressed plants while knocked‐out of the gene had the opposite impact. In conclusion, our findings demonstrated that FaSRT1‐2 could positively promote strawberry plant vegetative growth and fruit ripening by affecting ABA and GA pathways. However, it negatively regulates the resistance to heat stress and B. cinerea infection by influencing the related gene expression. Summary statement: FaSRT1‐2 is likely to exert a positive regulatory effect on both strawberry fruit ripening and vegetative growth by stimulating sugar, anthocyanin, and the gibberellin metabolism pathway. Moreover, FaSRT1‐2 appears to enhance the susceptibility of strawberry plants to heat stress while also promoting defence against Botrytis cinerea through alterations in abscisic acid metabolism and stress‐related genes expression. [ABSTRACT FROM AUTHOR]

GR24, a synthetic strigolactone analogue, plays a crucial role in a wide range of life processes. Studying the effects of the pre-harvest application of GR24 on strawberry ripening and fruit quality provides a scientific basis for the application of GR24 in horticultural crop production. GR24 solutions at concentrations of 0, 0.5, 1 and 2 µmol/L were applied to strawberries at the de-greening stage. The results showed that pre-harvest treatment with 1 µmol/L GR24 increased fruit weight and size; promoted fruit softening; increased the content of sugars, total flavonoids, total phenolics, and anthocyanins, and increased antioxidant activity. These results proved that GR24 effectively accelerated fruit ripening and improved fruit quality. In addition, to gain more insight into the biological mechanism of GR24 in fruit ripening, we conducted a combined transcriptomic and metabolomic analysis. It was found that the differences were related to sugar and flavonoid biosynthesis, particularly in the later stage of fruit ripening. qRT-PCR analysis revealed that GR24 increased gene expression activity and metabolite accumulation in the anthocyanin and sugar biosynthetic pathway, thereby promoting strawberry color and flavor. [ABSTRACT FROM AUTHOR]

Fruit softening is a crucial factor that controls shelf life and commercial value. Pectate lyase (PL) has a major role in strawberry fruit softening. However, the PL gene family in strawberry has not been comprehensively analyzed. In this study, 65 FaPL genes were identified in the octoploid strawberry genome. Subcellular localization prediction indicated that FaPLs are mostly localized to the extracellular and cytoplasmic spaces. Duplication event analysis suggested that FaPL gene family expansion is mainly driven by whole genome or segmental duplication. The FaPL family members were classified into six groups according to the phylogenetic analysis. Among them, FaPL1, 3, 5, 20, 25, 42, and 57 had gradually increased expressions during strawberry fruit development and ripening and higher expression levels in the fruits with less firmness than that in firmer fruit. This result suggested that these members are involved in strawberry softening. Furthermore, overexpression of FaPL1 significantly reduced the fruit firmness, ascorbic acid (AsA), and malondialdehyde (MDA) content but obviously increased the anthocyanins, soluble proteins, and titratable acidity (TA), while it had no apparent effects on flavonoids, phenolics, and soluble sugar content. These findings provide basic information on the FaPL gene family for further functional research and indicate that FaPL1 plays a vital role in strawberry fruit softening. [ABSTRACT FROM AUTHOR]

Ascorbic acid (AsA) is a crucial water-soluble antioxidant in strawberry fruit, but limited research is currently available on the identification and functional validation of key genes involved in AsA metabolism in strawberries. This study analyzed the FaMDHAR gene family identification, which includes 168 genes. Most of the products of these genes are predicted to exist in the chloroplast and cytoplasm. The promoter region is rich in cis-acting elements related to plant growth and development, stress and light response. Meanwhile, the key gene FaMDHAR50 that positively regulates AsA regeneration was identified through comparative transcriptome analysis of 'Benihoppe' strawberry (WT) and its natural mutant (MT) with high AsA content (83 mg/100 g FW). The transient overexpression experiment further showed that overexpression of FaMDHAR50 significantly enhanced the AsA content by 38% in strawberry fruit, with the upregulated expression of structural genes involved in AsA biosynthesis (FaGalUR and FaGalLDH) and recycling and degradation (FaAPX, FaAO and FaDHAR) compared with that of the control. Moreover, increased sugar (sucrose, glucose and fructose) contents and decreased firmness and citric acid contents were observed in the overexpressed fruit, which were accompanied by the upregulation of FaSNS, FaSPS, FaCEL1 and FaACL, as well as the downregulation of FaCS. Additionally, the content of pelargonidin 3-glucoside markedly decreased, while cyanidin chloride increased significantly. In summary, FaMDHAR50 is a key positive regulatory gene involved in AsA regeneration in strawberry fruit, which also plays an important role in the formation of fruit flavor, apperance and texture during strawberry fruit ripening. [ABSTRACT FROM AUTHOR]

The regulation of detached ripening is significant for prolonging fruit shelf life. Although light quality and sucrose affecting strawberry fruit ripening have been widely reported, little information is available about how they co-regulate the strawberry detached ripening process. In this study, different light qualities (red light—RL, blue light—BL, and white light—WL) and 100 mM sucrose were applied to regulate the ripening of initial red fruits detached from the plant. The results showed RL-treated samples (RL + H2O, RL + 100 mM sucrose) had brighter and purer skin color with a higher L*, b*, and C* value, and promoted the ascorbic acid. Almost all light treatments significantly decreased TSS/TA (total soluble solid/titratable acid) and soluble sugar/TA ratio, which is exacerbated by the addition of sucrose. Blue or red light in combination with sucrose notably increased total phenolic content and decreased malondialdehyde (MDA) accumulation. In addition, blue or red light combined with sucrose increased abscisic acid (ABA) content and promoted ABA signaling by inducing ABA-INSENSITIVE 4 (ABI4) expression and inhibiting SUCROSE NONFERMENTING1-RELATED PROTEIN KINASE 2.6 (SnRK2.6) expression. The strawberries exposed to blue and red light significantly improved auxin (IAA) content compared to the control (0 d), whereas the addition of sucrose inhibited IAA accumulation. Moreover, sucrose treatment suppressed the AUXIN/INDOLE-3-ACETIC ACID 11 (AUX/IAA11) and AUXIN RESPONSE FACTOR 6 (ARF6) expression under different light qualities. Overall, these results indicated that RL/BL + 100 mM sucrose might promote the detached ripening of strawberries by regulating abscisic acid and auxin signaling. [ABSTRACT FROM AUTHOR]

Proanthocyanidins (PAs), also known as condensed tannins, are widespread throughout the plant kingdom, presenting diverse biological and biochemical activities. Being one of the most abundant groups of natural polyphenolic antioxidant, PAs are applied to improve plant tolerance to (a)biotic stresses and delay the senescence of fruit by scavenging the reactive oxygen species (ROS) and enhancing antioxidant responses. The effects of PAs on coloring and softening of strawberries (Fragaria × ananassa Duch.), a worldwide demanded edible fruit and typical material for studying non-climacteric fruit ripening, were firstly assessed in this work. The results showed that exogenous PAs delayed the decrease in fruit firmness and anthocyanins accumulation but improved the fruit skin brightness. Strawberries treated with PAs had similar total soluble solids, total phenolics, and total flavonoids, but lower titratable acidity content. Moreover, the contents of endogenous PAs, abscisic acid and sucrose, were somehow increased by PA treatment, while no obvious change was found in fructose and glucose content. In addition, the anthocyanin- and firmness-related genes were significantly repressed, while the PA biosynthetic gene (anthocyanin reductase, ANR) was highly up-regulated by PA treatment at the key point for fruit softening and coloring. In summary, the results presented in this study suggest that PAs slow down strawberry coloration and softening by inhibiting the expression of related genes, which could be helpful for a better understanding of the biological role of PAs and provide a new strategy to regulate strawberry ripening. [ABSTRACT FROM AUTHOR]

WRKY transcription factors play a nonnegligible role in plant growth and development, but little is known about the involvement of WRKY transcription factors in the regulation of fruit ripening. In this study, FaWRKY71 was identified to be closely related to fruit maturation in octoploid strawberry. FaWRKY71 protein localized in the nucleus and responded to cold, salt, low phosphate, ABA, and light quality in strawberry seedlings. The temporal and spatial pattern expression analysis indicated that FaWRKY71 was expressed in all the detected tissues, especially in the full red fruits. In addition, FaWRKY71 gave rise to the accumulation of anthocyanin content by promoting the expression of structural genes FaF3'H, FaLAR, FaANR, and transport factors FaTT19 and FaTT12 in the flavonoid pathway, and softening the texture of strawberry via up-regulating the abundance of FaPG19 and FaPG21. Furthermore, FaWRKY71 was a positive regulator that mediated resistance against reactive oxygen species by enhancing the enzyme activities of SOD, POD, and CAT, reducing the amount of MDA. Altogether, this study provides new and comprehensive insight into the regulatory mechanisms facilitating fruit ripening in strawberry. [ABSTRACT FROM AUTHOR]

Ethylene response factors (ERFs) belonging to the APETALA2/ERF superfamily acted at the end of the ethylene signaling pathway, and they were found to play important roles in plant growth and development. However, the information of ERF genes in strawberry and their involvement in fruit ripening have been limited. Here, a total of 235 ERF members were identified from 426 AP2/ERF genes at octoploid strawberry genome level and classified into six subgroups according to their sequence characteristics and phylogenetic relationship. Conserved motif and gene structure analysis supported the evolutionary conservation of FaERFs. Syntenic analysis showed that four types of duplication events occurred during the expansion of FaERF gene family. Of these, WGD/segmental duplication played a major role. Transcriptomic data of FaERF genes during fruit ripening and in response to abscisic acid screened one activator (FaERF316) and one repressor (FaERF118) that were involved in fruit ripening. Transcriptional regulation analysis showed some transcription factors related to ripening such as ABI4, TCP15, and GLK1 could bind to FaERF316 or FaERF118 promoters, while protein–protein interaction analysis displayed some proteins associated with plant growth and development could interact with FaERF118 or FaERF316. These results suggested that FaERF118 and FaERF316 were potential genes to regulate strawberry ripening. In summary, the present study provides the comprehensive and systematic information on FaERF family evolution and gains insights into FaERF's potential regulatory mechanism in strawberry ripening. [ABSTRACT FROM AUTHOR]

Studies on many plants have shown that mitogen-activated protein kinases (MAPKs) are key proteins involved in regulating plant responses to biotic and abiotic stresses. However, their involvement in cultivated strawberry development and ripening remains unclear. In this study, 43 FaMAPK gene family members were identified in the genome of cultivated strawberry (Fragaria × ananassa), phylogenetic analysis indicated that FaMAPKs could be classified into four groups. Systematic analysis of the conserved motif, exon–intron structure showed that there were significant varieties between different groups in structure, but in the same group they were similar. Multiple cis-regulatory elements associated with phytohormone response, and abiotic and biotic stresses were predicted in the promoter regions of FaMAPK genes. Transcriptional analysis showed that all FaMAPK genes were expressed at all developmental stages. Meanwhile, the effect of exogenous ABA and sucrose on the expression profile of FaMAPKs was investigated. Exogenous ABA, sucrose, and ABA plus sucrose treatments upregulated the expression of FaMAPK genes and increased the content of endogenous ABA, sucrose, and anthocyanin in strawberry fruits, suggesting that ABA and sucrose might be involved in the FaMAPK-mediated regulation of strawberry fruit ripening. Based on the obtained results, MAPK genes closely related to the ripening of strawberries were screened to provide a theoretical basis and support for future research on strawberries. [ABSTRACT FROM AUTHOR]

Abscisic acid (ABA) and sucrose play an important role in strawberry fruit ripening, but how ABA and sucrose co-regulate this ripening progress remains unclear. The intention of this study was to examine the effect of ABA and sucrose on strawberry fruit ripening and to evaluate the ABA/sucrose interaction mechanism on the strawberry fruit ripening process. Here, we report that there is an acute synergistic effect between ABA and sucrose in accelerating strawberry fruit ripening. The time frame of fruit development and ripening was shortened after the application of ABA, sucrose, and ABA + sucrose, but most of the major quality parameters in treated-ripe fruit, including fruit weight, total soluble solids, anthocyanin, ascorbic acid, the total phenolic content, lightness (L*), chroma (C*), and hue angle (h°) values were not affected. Meanwhile, the endogenous ABA and sucrose levels, and the expression of ABA and sucrose signaling genes and ripening-related genes, such as NCED1, NCED2, SnRK2.2, SuSy, MYB5, CEL1, and CEL2, was all significantly enhanced by ABA or sucrose treatment alone, but in particular, by the ABA + sucrose treatment. Therefore, improving the ripening regulation efficiency is one synergetic action of ABA/sucrose. Another synergetic action of ABA/sucrose shows that a short inhibition of glycolysis occurs during accelerated strawberry ripening. ABA and sucrose can induce higher accumulation of H2O2, leading to a transient decrease in glycolysis. Conversely, lower endogenous H2O2 levels caused by reduced glutathione (GSH) treatment resulted in a transient increase in glycolysis while delaying strawberry fruit ripening. Collectively, this study demonstrates that the ABA/sucrose interaction affects the ripening regulation efficiency and shows inhibition of glycolysis. [ABSTRACT FROM AUTHOR]

Anthocyanins are responsible for the red color of strawberry, they are a subclass of flavonoids synthesized in cytosol and transferred to vacuole to form the visible color. Previous studies in model and ornamental plants indicated members of the glutathione S-transferase (GST) gene family were involved in vacuolar accumulation of anthocyanins. In the present study, a total of 130 FaGST genes were identified in the genome of cultivated strawberry (Fragaria × ananassa), which were unevenly distributed across the 28 chromosomes from the four subgenomes. Evolutionary analysis revealed the expansion of FaGST family was under stable selection and mainly drove by WGD/segmental duplication event. Classification and phylogenetic analysis indicated that all the FaGST genes were clarified into seven subclasses, among which FaGST1, FaGST37, and FaGST97 belonging to Phi class were closely related to FvRAP, an anthocyanin-related GST of wildwood strawberry, and this clade was clustered with other known anthocyanin-related GSTs. RNAseq-based expression analysis at different developmental stages of strawberry revealed that the expression of FaGST1, FaGST37, FaGST39, FaGST73, and FaGST97 was gradually increased during the fruit ripening, consistent with the anthocyanins accumulation. These expression patterns of those five FaGST genes were also significantly correlated with those of other anthocyanin biosynthetic genes such as FaCHI, FaCHS, and FaANS, as well as anthocyanin regulatory gene FaMYB10. These results indicated FaGST1, FaGST37, FaGST39, FaGST73, and FaGST97 may function in vacuolar anthocyanin accumulation in cultivated strawberry. [ABSTRACT FROM AUTHOR]

Cytosolic glyceraldehyde-3-phosphate dehydrogenase (GAPC) and plastid glyceraldehyde-3-phosphate dehydrogenase (GAPCp) are key enzymes in glycolysis. Besides their catalytic function, GAPC/GAPCp participates in the regulation of plant stress response and growth and development. However, the involvement of GAPC/GAPCp in the regulation of fruit ripening is unclear. In this study, FaGAPC2 and FaGAPCp1 in strawberries were isolated and analyzed. FaGAPC2 and FaGAPCp1 transcripts showed high transcript levels in the fruit. Transient overexpression of FaGAPC2 and FaGAPCp1 delayed fruit ripening, whereas RNA interference promoted fruit ripening and affected fruit anthocyanins and sucrose levels. Change in the expression patterns of FaGAPC2 and FaGAPCp1 also influenced the expression of several glycolysis-related and ripening-related genes such as CEL1, CEL2, SS, ANS, MYB5, NCED1, ABI1, ALDO, PK, and G6PDH, and H2O2 level and reduced glutathione (GSH)/glutathione disulfide (GSSG) redox potential. Meanwhile, metabolomics experiments showed that transient overexpression of FaGAPCp1 resulted in a decrease in anthocyanins, flavonoids, organic acid, amino acids, and their derivatives. In addition, abscisic acid (ABA) and sucrose treatment induced the production of large amounts of H2O2 and inhibited the expression of FaGAPC2/FaGAPCp1 in strawberry fruit. These results revealed that FaGAPC2/FaGAPCp1 is a negative regulator of ABA and sucrose mediated fruit ripening which can be regulated by oxidative stress. [ABSTRACT FROM AUTHOR]

Anthocyanins are one of the essential compounds affecting the quality of strawberry fruit. Several R2R3-MYB transcription factors have been well characterized as the anthocyanin activators in strawberry. However, we still lack valuable information on how R2R3-MYB repressors regulate strawberry anthocyanin accumulation. Here, we cloned a novel R2R3-MYB gene, FaMYB6-like , and characterized its repressive role during anthocyanin accumulation. FaMYB6-like encodes a nuclear localization protein, which was clustered into the FaMYB1-like type repressor clade in phylogenetic analysis. FaMYB6-like showed high expressions in anthocyanin-accumulating fruit and low transcript abundance in flowers, vegetative tissues, and early development stages of fruit. Transient overexpression of FaMYB6L-like repressed the expression of anthocyanin biosynthetic and transport genes, and reduced anthocyanin content in postharvest strawberry fruit. Transgenic tobacco lines overexpressing FaMYB6-like displayed a reduction in petal pigmentation. Overexpression of FaMYB6L-like in white-fleshed strawberry inhibited anthocyanin accumulation induced by FaMYB10. Protein interaction and promoter activation assays revealed that FaMYB6-like repressed the activation of target gene promoters by FaMYB10-FabHLH3 activation complex via its interaction with FabHLH3. Additionally, FaMYB6-like promoter activity can be activated by FaMYB10-FabHLH3 complex. Taken together, our study revealed that FaMYB6-like acted as a transcriptional repressor involved in an 'activator-and-repressor' feedback regulation mechanism for maintaining the balance of anthocyanin concentrations. • A new anthocyanin repressor, FaMYB6-like , was isolated from cultivated strawberry. • FaMYB6-like was highly expressed in anthocyanin-pigmented fruits. • Transgenic tobacco lines overexpressing FaMYB6-like displayed a reduction in petal pigmentation. • FaMYB6-like acts as the anthocyanin repressor by inhibiting MBW activity. [ABSTRACT FROM AUTHOR]

Fatty acid desaturases (FADs) play a crucial role in plant growth as well as in the regulation of biotic and abiotic stresses. However, their function has not been well understood in the ripening of fruit. In this study, we identified 73 FADs in the octoploid strawberry genome and analyzed their physicochemical characteristics, chromosomal localization, phylogeny, gene structures, covariance, cis -acting elements, and transcript levels during fruit ripening stages. According to the predicted subcellular localization, FADs are mostly located in the plasma membrane, with a lower abundance in the cell wall, cytoplasm, and nucleus. Numerous cis -elements in the FADs promoter are responsive to light, hormones, and stress. Here, combining comparative transcriptomics, we identified a Δ9 stearoyl-acyl carrier protein (ACP) desaturase gene (FxaC_8 g24100, namely FaSAD3) that is negatively regulated by both key glycolysis genes, cytoplasmic glyceraldehyde-3-phosphate dehydrogenase (FaGAPC2) and pyruvate kinase (FaPKc2.2). The spatiotemporal expression analysis revealed that FaSAD3 is expressed in both vegetative and reproductive organs. Subcellular localization analysis confirmed that FaSAD3 is expressed in both the cytoplasm and nucleus. Meanwhile, the transient overexpression of FaSAD3 reduced anthocyanin, sugar, organic acid, total phenol, and flavonoid contents, while increased fruit firmness. Furthermore, the yeast one hybridization and dual-luciferase assay revealed that the ripening regulator FaGAMYB negatively regulates the transcriptional activity of FaSAD3 promoter, promoting strawberry fruit ripening. These findings indicated that FaSAD3 acts as a negative regulator in strawberry fruits ripening, providing a novel model for comprehending the mechanisms involved in strawberry fruit ripening. [ABSTRACT FROM AUTHOR]

• A total of 53 PKs were identified in the genome of octoploid strawberry, seven of which specifically express in the fruits. • FaPKc2.2 was significantly inhibited by both ABA and sucrose, either alone or synergistically. • Transient overexpression of FaPKc2.2 significantly inhibited the ripening of strawberry fruits. • Upregulation of FaPKc2.2 systematically reprogramed metabolisms, mainly by channeling the main flux from carbon to amino acid related nitrogen biomolecules. Although the main contribution of pyruvate kinases in the energy providing and regulation of plant growth or seed development have been discovered, the potential roles of these enzymes in the fruit ripening process has not been well elucidated so far. In this study, a total of 53 pyruvate kinases (PKs) were identified in the genome of octoploid strawberry. Expression specificity in the fruit led to the identification of a cytosolic PK (FxaC_15g00080, namely FaPKc2.2), which was inhibited by both ABA and sucrose, either alone or synergistically. It was expressed in both vegetative and reproductive organs. Subcellular localization analysis confirmed that the FaPKc2.2 was a cytosolic PK. Transient overexpression of the FaPKc2.2 obviously inhibited the ripening of strawberry fruits. Biochemical and transcriptomic data further revealed that the upregulation of the FaPKc2.2 systematically reprogrammed metabolisms, mainly by channeling the main flux from carbon to amino-acid related nitrogen biomolecules. These results demonstrated the regulatory role of the FaPKc2.2 in the fruit ripening and highlighted the probability of utilizing the gene in strawberry fruit quality improvements. [ABSTRACT FROM AUTHOR]

• A total of four FvbZIP-S1 genes were identified in strawberry genome database. • In FvbZIP-S1 subfamily, FvbZIP11 most highly expressed in different tissues, and especially it had a very remarkable transcript accumulation during the fruit development. • Constitutive expression of FvbZIP11 in tomato caused a plant growth impairment, but significantly increased the fruit TSS and SS content and SS/TA ratio. Consumers and breeders value strawberry fruit with high sugar content and sweetness. However, many factors negatively affect this trait. bZIP-S1 subfamily was closely correlated with the carbohydrate metabolism. Moreover, it can regulate plant development and growth, and respond to various stress. In the present study, a total of four bZIP-S1 members were identified in strawberry. Quantitative RT-PCR analysis showed a member of this subfamily, designated as bZIP11 , most highly expressed in different tissues of strawberry. Especially, it had a significant transcript accumulation during the fruit development and ripening. Constitutive expression of FvbZIP11 in tomato caused severe plant growth retardation which was accompanied by wrinkled and curly leaves. The leaf yellowish and chlorotic patches were reflected in decrease of chlorophyll. Meanwhile, epidermal cells were smaller in leaves of transgenic plant. In addition, the total soluble solid and soluble sugar content of the fruit was significantly higher than WT during the fruit ripening. Taken together, the heterologous overexpression of strawberry bZIP11 induced sugar accumulation and hindered plant growth in tomato, which provided more information for improving strawberry fruit quality. [ABSTRACT FROM AUTHOR]