記述言語：英語   掲載種別：研究論文（学術雑誌）  

Deep neural network (DNN) techniques, as an advanced machine learning framework, have allowed various image diagnoses in plants, which often achieve better prediction performance than human experts in each specific field. Notwithstanding, in plant biology, the application of deep neural networks is still mostly limited to rapid and effective phenotyping. Recent development of explainable CNN frameworks has allowed visualization of the features in the prediction by convolutional neural network (CNN), which potentially contributes to the understanding of physiological mechanisms in objective phenotypes. In this study, we propose an integration of explainable CNN and transcriptomic approach to make a physiological interpretation of a fruit internal disorder in persimmon, rapid over-softening. We constructed CNN models to accurately predict the fate to be rapid softening in persimmon cv. Soshu, only with photo images. The explainable CNNs, such as Grad-CAM and Guided Grad-CAM, visualized specific featured regions relevant to the prediction of rapid-softening, which would correspond to the premonitory symptoms in a fruit. Transcriptomic analyses to compare the featured regions of predicted rapid-softening and control fruits suggested that rapid softening is triggered by precocious ethylene signal-dependent cell wall modification, despite exhibiting no direct phenotypic changes. Further transcriptomic comparison between the featured and non-featured regions in predicted rapid-softening fruit suggested that premonitory symptoms reflected hypoxia and the related stress signals finally to induce ethylene signals. These results would provide a good example for the collaboration of image analysis and omics approaches in plant physiology, which uncovered a novel aspect of fruit premonitory reactions in the rapid softening fate.

DOI： 10.1093/pcp/pcad050

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記述言語：英語   掲載種別：研究論文（学術雑誌）  

Sexuality is the main strategy for maintaining genetic diversity within a species. In flowering plants (angiosperms), sexuality is derived from ancestral hermaphroditism and multiple sexualities can be expressed in an individual. The mechanisms conferring chromosomal sex determination in plants (or dioecy) have been studied for over a century by both biologists and agricultural scientists, given the importance of this field for crop cultivation and breeding. Despite extensive research, the sex determining gene(s) in plants had not been identified until recently. In this review, we dissect plant sex evolution and determining systems, with a focus on crop species. We introduced classic studies with theoretical, genetic, and cytogenic approaches, as well as more recent research using advanced molecular and genomic techniques. Plants have undergone very frequent transitions into, and out of, dioecy. Although only a few sex determinants have been identified in plants, an integrative viewpoint on their evolutionary trends suggests that recurrent neofunctionalization events are potentially common, in a "scrap and (re)build" cycle. We also discuss the potential association between crop domestication and transitions in sexual systems. We focus on the contribution of duplication events, which are particularly frequent in plant taxa, as a trigger for the creation of new sexual systems.

DOI： 10.1270/jsbbs.22082

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掲載種別：研究論文（学術雑誌）   出版者・発行元：Springer Science and Business Media LLC  

DOI： 10.1038/s41477-023-01361-9

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その他リンク： https://www.nature.com/articles/s41477-023-01361-9

DOI： 10.2503/hortj.QH-012

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掲載種別：研究論文（学術雑誌）   出版者・発行元：Japanese Society for Horticultural Science  

DOI： 10.2503/hrj.22.303

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掲載種別：論文集(書籍)内論文   出版者・発行元：CABI  

DOI： 10.1079/9781789247534.0017

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記述言語：英語   掲載種別：研究論文（学術雑誌）  

Recent advances in the genomics of polyploid species answer some of the long-standing questions about the role of polyploidy in crop species. Here, we summarize the current literature to reexamine scenarios in which polyploidy played a role both before and after domestication. The prevalence of polyploidy can help to explain environmental robustness in agroecosystems. This review also clarifies the molecular basis of some agriculturally advantageous traits of polyploid crops, including yield increments in polyploid cotton via subfunctionalization, modification of a separated sexuality to selfing in polyploid persimmon via neofunctionalization, and transition to a selfing system via nonfunctionalization combined with epistatic interaction between duplicated S-loci. The rapid progress in genomics and genetics is discussed along with how this will facilitate functional studies of understudied polyploid crop species.

DOI： 10.1016/j.pbi.2022.102255

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掲載種別：論文集(書籍)内論文   出版者・発行元：Springer International Publishing  

DOI： 10.1007/978-3-031-05584-3_8

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掲載種別：研究論文（学術雑誌）   出版者・発行元：International Society for Horticultural Science (ISHS)  

DOI： 10.17660/actahortic.2022.1338.49

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記述言語：英語   掲載種別：研究論文（学術雑誌）  

In the evolutionary history of plants, variation in cis-regulatory elements (CREs) resulting in diversification of gene expression has played a central role in driving the evolution of lineage-specific traits. However, it is difficult to predict expression behaviors from CRE patterns to properly harness them, mainly because the biological processes are complex. In this study, we used cistrome datasets and explainable convolutional neural network (CNN) frameworks to predict genome-wide expression patterns in tomato (Solanum lycopersicum) fruit from the DNA sequences in gene regulatory regions. By fixing the effects of trans-acting factors using single cell-type spatiotemporal transcriptome data for the response variables, we developed a prediction model for crucial expression patterns in the initiation of tomato fruit ripening. Feature visualization of the CNNs identified nucleotide residues critical to the objective expression pattern in each gene, and their effects were validated experimentally in ripening tomato fruit. This cis-decoding framework will not only contribute to the understanding of the regulatory networks derived from CREs and transcription factor interactions, but also provides a flexible means of designing alleles for optimized expression.

DOI： 10.1093/plcell/koac079

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記述言語：英語   掲載種別：研究論文（学術雑誌）  

In flowering plants, different lineages have independently transitioned from the ancestral hermaphroditic state into and out of various sexual systems1. Polyploidizations are often associated with this plasticity in sexual systems2,3. Persimmons (the genus Diospyros) have evolved dioecy via lineage-specific palaeoploidizations. More recently, hexaploid D. kaki has established monoecy and also exhibits reversions from male to hermaphrodite flowers in response to natural environmental signals (natural hermaphroditism, NH), or to artificial cytokinin treatment (artificial hermaphroditism, AH). We sought to identify the molecular pathways underlying these polyploid-specific reversions to hermaphroditism. Co-expression network analyses identified regulatory pathways specific to NH or AH transitions. Surprisingly, the two pathways appeared to be antagonistic, with abscisic acid and cytokinin signalling for NH and AH, respectively. Among the genes common to both pathways leading to hermaphroditic flowers, we identified a small-Myb RADIALIS-like gene, named DkRAD, which is specifically activated in hexaploid D. kaki. Consistently, ectopic overexpression of DkRAD in two model plants resulted in hypergrowth of the gynoecium. These results suggest that production of hermaphrodite flowers via polyploidization depends on DkRAD activation, which is not associated with a loss-of-function within the existing sex determination pathway, but rather represents a new path to (or reinvention of) hermaphroditism.

DOI： 10.1038/s41477-022-01107-z

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掲載種別：研究論文（学術雑誌）   出版者・発行元：Elsevier {BV}  

DOI： 10.1016/j.postharvbio.2021.111787

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記述言語：英語   掲載種別：研究論文（学術雑誌）  

Proper anther and pollen development are important for plant reproduction. The plant hormone gibberellin is important for anther development in rice, but its gametophytic functions remain largely unknown. Here, we report the functional and evolutionary analyses of rice gibberellin 3-oxidase 1 (OsGA3ox1), a gibberellin synthetic enzyme specifically expressed in the late developmental stages of anthers. Enzymatic and X-ray crystallography analyses reveal that OsGA3ox1 has a higher GA7 synthesis ratio than OsGA3ox2. In addition, we generate an osga3ox1 knockout mutant by genome editing and demonstrate the bioactive gibberellic acid synthesis by the OsGA3ox1 action during starch accumulation in pollen via invertase regulation. Furthermore, we analyze the evolution of Oryza GA3ox1s and reveal that their enzyme activity and gene expression have evolved in a way that is characteristic of the Oryza genus and contribute to their male reproduction ability.

DOI： 10.1038/s42003-022-03008-5

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記述言語：英語   掲載種別：研究論文（学術雑誌）  

Peel degreening is the most conspicuous aspect of fruit ripening in many citrus fruits because of its importance for marketability. In this study, peel degreening in response to propylene (an ethylene analog) and at varying storage temperatures was characterized in Satsuma mandarin (Citrus unshiu Marc.) fruit. Propylene treatment triggered rapid peel degreening (within 4-6 days), indicated by an increase in the citrus color index (CCI) and chlorophyll loss. Peel degreening was also observed in fruit at 10°C and 15°C after 28-42 days, with gradual CCI increase and chlorophyll reduction. However, fruit at 5°C, 20°C, and 25°C remained green, and no substantial changes in peel CCI and chlorophyll content were recorded during the 42-day storage duration. The transcriptomes of peels of fruit treated with propylene for 4 days and those stored at varying temperatures for 28 days were then analyzed by RNA-Seq. We identified three categories of differentially expressed genes that were regulated by (i) propylene (and by analogy, ethylene) alone, (ii) low temperature (5°C, 10°C, or 15°C vs. 25°C) alone, and (iii) either propylene or low temperature. Gene-encoding proteins associated with chlorophyll degradation (such as CuSGR1, CuNOL, CuACD2, CuCAB2, and CuLHCB2) and a transcription factor (CuERF114) were differentially expressed by propylene or low temperature. To further examine temperature-induced pathways, we also monitored gene expression during on-tree fruit maturation vs. postharvest. The onset of on-tree peel degreening coincided with autumnal drops in field temperatures, and it was accompanied by differential expression of low temperature-regulated genes. On the contrary, genes that were exclusively regulated by propylene (such as CuCOPT1 and CuPOX-A2) displayed insignificant expression changes during on-tree peel degreening. These findings indicate that low temperatures could be involved in the fruit ripening-related peel degreening independently of ethylene.

DOI： 10.3389/fpls.2022.918226

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掲載種別：研究論文（学術雑誌）  

In contrast to the progress in the research on physiological disorders relating to shelf life in fruit crops, it has been difficult to non-destructively predict their occurrence. Recent high-tech instruments have gradually enabled non-destructive predictions for various disorders in some crops, while there are still issues in terms of efficiency and costs. Here, we propose application of a deep neural network (or simply deep learning) to simple RGB images to predict a severe fruit disorder in persimmon, rapid over-softening. With 1,080 RGB images of ‘Soshu’ persimmon fruits, three convolutional neural networks (CNN) were examined to predict rapid over-softened fruits with a binary classification and the date to fruit softening. All of the examined CNN models worked successfully for binary classification of the rapid over-softened fruits and the controls with > 80% accuracy using multiple criteria. Furthermore, the prediction values (or confidence) in the binary classification were correlated to the date to fruit softening. Although the features for classification by deep learning have been thought to be in a black box by conventional standards, recent feature visualization methods (or “explainable” deep learning) has allowed identification of the relevant regions in the original images. We applied Grad-CAM, Guided backpropagation, and layer-wise relevance propagation (LRP), to find early symptoms for CNNs classification of rapid over-softened fruits. The focus on the relevant regions tended to be on color unevenness on the surface of the fruit, especially in the peripheral regions. These results suggest that deep learning frameworks could potentially provide new insights into early physiological symptoms of which researchers are unaware.

DOI： 10.2503/hortj.UTD-323

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記述言語：英語   掲載種別：研究論文（学術雑誌）  

Kiwifruit (Actinidia chinensis) is a dioecious, long-living woody perennial vine. Reduced generation time and induction of hermaphroditism can accelerate crop improvement and facilitate alternative farming for better food security in the face of climate change. Previous studies identified that CENTRORADIALIS genes CEN and CEN4 act to repress flowering, whilst the male-specific Shy Girl (SyGl) gene with homology to type-C cytokinin response regulators could repress gynoecium development in model plants. Here we use CRISPR/Cas9 to mutagenize CEN, CEN4 and SyGl in the male kiwifruit A. chinensis 'Bruce'. Biallelic mutations of CEN and CEN4 generated rapid-flowering male plants, and simultaneous targeting of CEN4 and SyGl gave rise to rapid-flowering hermaphrodites with restored gynoecial function and viable pollen, providing functional evidence for the role of SyGl in suppression of feminization. Analysis of ovary tissues identified genes that contribute to carpel development and revealed that SyGl affected both cytokinin profiles and the expression of genes involved in cytokinin metabolism and signalling. The plant lines generated by CEN4/SyGl knockout could self-pollinate and produce fast-flowering offspring. These results establish that SyGI acts as the suppressor of feminization in kiwifruit and demonstrate the potential for accelerated breeding in an outcrossing horticultural woody perennial.

DOI： 10.1111/nph.17234

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掲載種別：研究論文（学術雑誌）  

Noninvasive diagnosis of internal traits in fruit crops is a high unmet need; however it generally requires time, costs, and special methods or facilities. Recent progress in deep neural network (or deep learning) techniques would allow easy, but highly accurate diagnosis with single RGB images, and the latest applications enable visualization of “the reasons for each diagnosis” by backpropagation of neural networks. Here, we propose an application of deep learning for image diagnosis on the classification of internal fruit traits, in this case seedlessness, in persimmon fruit (Diospyros kaki). We examined the classification of seedlessness in persimmon fruit by using four convolutional neural networks (CNN) models with various layer structures. With only 599 pictures of ‘Fuyu’ persimmon fruit from the fruit apex side, the neural networks successfully made a binary classification of seedless and seeded fruits with up to 85% accuracy. Among the four CNN models, the VGG16 model with the simplest layer structure showed the highest classification accuracy of 89%. Prediction values for the binary classification of seeded fruits were significantly increased in proportion to seed numbers in all four CNN models. Furthermore, explainable AI methods, such as Gradient-weighted Class Activation Mapping (Grad-CAM) and Guided Grad-CAM, allowed visualization of the parts and patterns contributing to the diagnosis. The results indicated that finer positions surrounding the apex, which correspond to hypothetical bulges derived from seeds, are an index for seeded fruits. These results suggest the novel potential of deep learning for noninvasive diagnosis of fruit internal traits using simple RGB images and also provide novel insights into previously unrecognized features of seeded/seedless fruits.

DOI： 10.2503/hortj.UTD-248

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記述言語：英語  

DOI： 10.3389/fpls.2021.719588

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：FRONTIERS MEDIA SA  

Sex expression in plants is often flexible and contributes to the maintenance of genetic diversity within a species. In diploid persimmons (the genus Diospyros), the sexuality is controlled by the Y chromosome-encoded small-RNA gene, OGI, and its autosomal counterpart, MeGI. Hexaploid Oriental persimmon (Diospyros kaki) evolved more flexible sex expression, where genetically male individuals carrying OGI can produce both male and female flowers (monoecy). This is due to (semi-)inactivation of OGI by the Kali-SINE retrotransposon insertion on the promoter region and the resultant DNA methylations. Instead, flower sex determination in Oriental persimmon is also dependent on DNA methylation states of MeGI. Here, we focused on a cultivar, Kumemaru, which shows stable male flower production. Our results demonstrated that cv. Kumemaru carries OGI with Kali-SINE, which was highly methylated as well as in other monoecious cultivars; nevertheless, OGI gene could have a basal expression level. Transcriptomic analysis between cv. Kumemaru and 14 cultivars that predominantly produce female flowers showed differentially expressed genes (DEGs) specific to cv. Kumemaru, which is mainly involved in stress responses. Co-expression gene networks focusing on the DEGs also suggested the involvement of stress signals, mainly via gibberellin (GA), salicylic acid (SA), and especially jasmonic acid (JA) signal pathways. We also identified potential regulators of this co-expression module, represented by the TCP4 transcription factor. Furthermore, we attempted to identify cv. Kumemaru-specific transcript polymorphisms potentially contributing to derepressed OGI expression by cataloging subsequences (k-mers) in the transcriptomic reads from cv. Kumemaru and the other 14 female cultivars. Overall, although the direct genetic factor to activate OGI remains to be solved, our results implied the involvement of stress signals in the release of silenced OGI and the resultant continuous male production.

DOI： 10.3389/fpls.2020.567249

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：Wiley  

Domestication and population differentiation in crops involve considerable phenotypic changes. The logs of these evolutionary paths, including natural/artificial selection, can be found in the genomes of the current populations. However, these profiles have been little studied in tree crops, which have specific characters, such as long generation time and clonal propagation, maintaining high levels of heterozygosity. We conducted exon-targeted resequencing of 129 genomes in the genus Prunus, mainly Japanese apricot (Prunus mume), and apricot (Prunus armeniaca), plum (Prunus salicina), and peach (Prunus persica). Based on their genome-wide single-nucleotide polymorphisms merged with published resequencing data of 79 Chinese P. mume cultivars, we inferred complete and ongoing population differentiation in P. mume. Sliding window characterization of the indexes for genetic differentiation identified interspecific fragment introgressions between P. mume and related species (plum and apricot). These regions often exhibited strong selective sweeps formed in the paths of establishment or formation of substructures of P. mume, suggesting that P. mume has frequently imported advantageous genes from other species in the subgenus Prunus as adaptive evolution. These findings shed light on the complicated nature of adaptive evolution in a tree crop that has undergone interspecific exchange of genome fragments with natural/artificial selections.

DOI： 10.1111/tpj.15020

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：Oxford University Press (OUP)  

<title>Abstract</title>
Recent rapid progress in deep neural network techniques have allowed various object recognitions and classifications, which often exceed the performance with human eyes. In plant biology and crop sciences, some deep neural network frameworks have been applied mainly to effective and rapid phenotyping. In this study, beyond simple optimizations of phenotyping, we propose an application of deep neural networks to make image-based internal disorder diagnosis that is hard even for experts, and to visualize the reasons of each diagnosis to provide biological interpretations. Here we exemplified classification of calyx-end cracking in persimmon fruit by using five convolutional neural network models with various layer structures, and examined potential analytic options involved in the diagnostic qualities. With 3,173 visible RGB images from the fruit apex side, the neural networks successfully made the binary classification of each degree of disorders, with up to 90% accuracy. Furthermore, feature visualizations, such as Grad-CAM and LRP, visualize the image regions that contribute to the diagnosis. They suggest that specific patterns of color unevenness, such as in fruit peripheral area, can be indexes of calyx-end cracking. These results not only provided novel insights into indexes of fruit internal disorders but proposed potential applicability of deep neural networks in plant biology.

DOI： 10.1093/pcp/pcaa111

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掲載種別：研究論文（学術雑誌）  

© 2020 Oxford University Press. All rights reserved. Peel degreening is an important aspect of fruit ripening in many citrus fruit, and previous studies have shown that it can be advanced by ethylene treatment or by low-temperature storage. However, the important regulators and pathways involved in natural peel degreening remain largely unknown. To determine how natural peel degreening is regulated in lemon fruit (Citrus limon), we studied transcriptome and physiochemical changes in the flavedo in response to ethylene treatment and low temperatures. Treatment with ethylene induced rapid peel degreening, which was strongly inhibited by the ethylene antagonist, 1-methylcyclopropene (1-MCP). Compared with 25 ºC, moderately low storage temperatures of 5-20 °C also triggered peel degreening. Surprisingly, repeated 1-MCP treatments failed to inhibit the peel degreening induced by low temperature. Transcriptome analysis revealed that low temperature and ethylene independently regulated genes associated with chlorophyll degradation, carotenoid metabolism, photosystem proteins, phytohormone biosynthesis and signalling, and transcription factors. Peel degreening of fruit on trees occurred in association with drops in ambient temperature, and it coincided with the differential expression of low temperature-regulated genes. In contrast, genes that were uniquely regulated by ethylene showed no significant expression changes during on-tree peel degreening. Based on these findings, we hypothesize that low temperature plays a prominent role in regulating natural peel degreening independently of ethylene in citrus fruit.

DOI： 10.1093/jxb/eraa206

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：Springer Science and Business Media LLC  

The size of plants is largely determined by growth of the stem. Stem elongation is stimulated by gibberellic acid(1-3). Here we show that internode stem elongation in rice is regulated antagonistically by an 'accelerator' and a 'decelerator' in concert with gibberellic acid. Expression of a gene we nameACCELERATOR OF INTERNODE ELONGATION 1(ACE1), which encodes a protein of unknown function, confers cells of the intercalary meristematic region with the competence for cell division, leading to internode elongation in the presence of gibberellic acid. By contrast, upregulation ofDECELERATOR OF INTERNODE ELONGATION 1(DEC1), which encodes a zinc-finger transcription factor, suppresses internode elongation, whereas downregulation ofDEC1allows internode elongation. We also show that the mechanism of internode elongation that is mediated byACE1andDEC1is conserved in the Gramineae family. Furthermore, an analysis of genetic diversity suggests that mutations inACE1andDEC1have historically contributed to the selection of shorter plants in domesticated populations of rice to increase their resistance to lodging, and of taller plants in wild species of rice for adaptation to growth in deep water. Our identification of these antagonistic regulatory factors enhances our understanding of the gibberellic acid response as an additional mechanism that regulates internode elongation and environmental fitness, beyond biosynthesis and gibberellic acid signal transduction.

DOI： 10.1038/s41586-020-2501-8

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その他リンク： http://www.nature.com/articles/s41586-020-2501-8

記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：Oxford University Press (OUP)  

<title>Abstract</title>
Sexuality is one of the fundamental mechanisms that work towards maintaining genetic diversity within a species. In diploid persimmons (Diospyros spp.), separated sexuality, the presence of separate male and female individuals (dioecy), is controlled by the Y chromosome-encoded small-RNA gene, OGI. On the other hand, sexuality in hexaploid Oriental persimmon (Diospyros kaki) is more plastic, with OGI-bearing genetically male individuals, able to produce both male and female flowers (monoecy). This is thought to be linked to the partial inactivation of OGI by a retrotransposon insertion, resulting in DNA methylation of the OGI promoter region. To identify the genetic factors regulating branch sexual conversion, genome-wide correlation/association analyses were conducted using ddRAD-Seq data from an F1 segregating population, and using both quantitative and diploidized genotypes, respectively. We found that allelic ratio at the Y-chromosomal region, including OGI, was correlated with male conversion based on quantitative genotypes, suggesting that OGI can be activated in cis in a dosage-dependent manner. Genome-wide association analysis based on diploidized genotypes, normalized for the effect of OGI allele dosage, detected three fundamental loci associated with male conversion. These loci underlie candidate genes, which could potentially act epigenetically for the activation of OGI expression.

DOI： 10.1093/dnares/dsaa012

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その他リンク： http://academic.oup.com/dnaresearch/article-pdf/27/3/dsaa012/33571384/dsaa012.pdf

記述言語：英語  

[This corrects the article DOI: 10.1371/journal.pgen.1008566.].

DOI： 10.1371/journal.pgen.1008845

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：Oxford University Press (OUP)  

<title>Abstract</title>
Epigenetic regulation adds a flexible layer to genetic variations, potentially enabling long-term, but reversible, changes to a trait, while maintaining genetic information. In the hexaploid Oriental persimmon (Diospyros kaki), genetically monoecious cultivars bearing male flowers require the Y-encoded small RNA (smRNA) gene, OGI. This gene represses the expression of its autosomal counterpart gene, MeGI, as part of the canonical male production system. However, a D. kaki cultivar, Saijo, which lacks the OGI gene and originally bears only female flowers, occasionally produces somaclonal mutant male and revertant female (RF) branches. In this study, we investigated the mechanisms underlying these somaclonal sex conversions in persimmon. Specifically, we aimed to unravel how a genetically female tree without the OGI gene can produce male flowers and RF flowers. Applying multi-omics approaches, we revealed that this noncanonical male production system is basically consistent with the canonical system, in which the accumulation of smRNA targeting MeGI and the considerable DNA methylation of MeGI are involved. The epigenetic status of MeGI on CGN and CHG was synchronized to the genome-wide methylation patterns, both in transition to and from the male production system. These results suggest that the somaclonal sex conversions in persimmon are driven by the genome-wide epigenetic regulatory activities. Moreover, flexibility in the epigenetic layers of long-lived plant species (e.g. trees) is important for overcoming genetic robustness.

DOI： 10.1093/pcp/pcz207

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その他リンク： http://academic.oup.com/pcp/article-pdf/61/2/393/32741507/pcz207.pdf

記述言語：英語   掲載種別：研究論文（学術雑誌）  

Most angiosperms bear hermaphroditic flowers, but a few species have evolved outcrossing strategies, such as dioecy, the presence of separate male and female individuals. We previously investigated the mechanisms underlying dioecy in diploid persimmon (D. lotus) and found that male flowers are specified by repression of the autosomal gene MeGI by its paralog, the Y-encoded pseudo-gene OGI. This mechanism is thought to be lineage-specific, but its evolutionary path remains unknown. Here, we developed a full draft of the diploid persimmon genome (D. lotus), which revealed a lineage-specific whole-genome duplication event and provided information on the architecture of the Y chromosome. We also identified three paralogs, MeGI, OGI and newly identified Sister of MeGI (SiMeGI). Evolutionary analysis suggested that MeGI underwent adaptive evolution after the whole-genome duplication event. Transformation of tobacco plants with MeGI and SiMeGI revealed that MeGI specifically acquired a new function as a repressor of male organ development, while SiMeGI presumably maintained the original function. Later, a segmental duplication event spawned MeGI's regulator OGI on the Y-chromosome, completing the path leading to dioecy, and probably initiating the formation of the Y-chromosome. These findings exemplify how duplication events can provide flexible genetic material available to help respond to varying environments and provide interesting parallels for our understanding of the mechanisms underlying the transition into dieocy in plants.

DOI： 10.1371/journal.pgen.1008566

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記述言語：英語   掲載種別：研究論文（学術雑誌）  

Sex expression in plants is often flexible and contributes to the maintenance of genetic diversity within a species. In diploid persimmons (the genus Diospyros), the sexuality is controlled by the Y chromosome-encoded small-RNA gene, OGI, and its autosomal counterpart, MeGI. Hexaploid Oriental persimmon (Diospyros kaki) evolved more flexible sex expression, where genetically male individuals carrying OGI can produce both male and female flowers (monoecy). This is due to (semi-)inactivation of OGI by the Kali-SINE retrotransposon insertion on the promoter region and the resultant DNA methylations. Instead, flower sex determination in Oriental persimmon is also dependent on DNA methylation states of MeGI. Here, we focused on a cultivar, Kumemaru, which shows stable male flower production. Our results demonstrated that cv. Kumemaru carries OGI with Kali-SINE, which was highly methylated as well as in other monoecious cultivars; nevertheless, OGI gene could have a basal expression level. Transcriptomic analysis between cv. Kumemaru and 14 cultivars that predominantly produce female flowers showed differentially expressed genes (DEGs) specific to cv. Kumemaru, which is mainly involved in stress responses. Co-expression gene networks focusing on the DEGs also suggested the involvement of stress signals, mainly via gibberellin (GA), salicylic acid (SA), and especially jasmonic acid (JA) signal pathways. We also identified potential regulators of this co-expression module, represented by the TCP4 transcription factor. Furthermore, we attempted to identify cv. Kumemaru-specific transcript polymorphisms potentially contributing to derepressed OGI expression by cataloging subsequences (k-mers) in the transcriptomic reads from cv. Kumemaru and the other 14 female cultivars. Overall, although the direct genetic factor to activate OGI remains to be solved, our results implied the involvement of stress signals in the release of silenced OGI and the resultant continuous male production.

DOI： 10.3389/fpls.2020.567249

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記述言語：英語   掲載種別：研究論文（学術雑誌）  

DOI： 10.1016/j.postharvbio.2019.110949

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：Oxford University Press (OUP)  

<title>Abstract</title>
The shapes of plant organs reflect the evolution of each lineage and have been diversified according to lineage-specific adaptations to environment. Research on the molecular pathways responsible for organ shapes has traditionally been focused mainly on leaves or flowers. Thus, little is known about the pathways controlling fruit shapes, despite their diversity in some plant species. In this study, we analyzed oriental persimmon (Diospyros kaki), which exhibits considerable diversity in fruit shapes among cultivars, to elucidate the underlying molecular mechanism using transcriptomic data and quantitative evaluation. First, to filter the candidate genes associated with persimmon fruit shapes, the whole gene expression patterns obtained using mRNA-Seq analysis from 100 individuals, including a segregated population and various cultivars, were assessed to detect correlations with principal component scores for fruit shapes characterized with elliptic Fourier descriptors. Next, a gene co-expression network analysis with weighted gene co-expression network analysis (WGCNA) package revealed that class 1 KNOX family genes and SEEDSTICK function as integrators along with some phytohormone-related genes, to regulate the fruit shape diversity. On the other hand, the OVATE family genes also contribute to fruit shape diversity, of which pathway would be potentially shared with other plant species. Evolutionary aspects suggest that acquisition of a high lineage-specific and variable expression of class 1 KNOX gene, knotted-like homeobox of Arabidopsis thaliana 1 (KNAT1), in young fruit is important for establishing the persimmon-specific mechanism that determines fruit shape diversity.

DOI： 10.1093/pcp/pcz139

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その他リンク： http://academic.oup.com/pcp/article-pdf/60/11/2464/30490055/pcz139.pdf

掲載種別：研究論文（学術雑誌）   出版者・発行元：The Royal Society  

One reason for studying sex chromosomes of flowering plants is that they have often evolved separate sexes recently, and the genomes of dioecious species may not yet have evolved adaptations to their changes from the ancestral state. An unstudied question concerns the relative importance of such adaptation, versus the effects of the mutations that led to separate sexes in the first place. Theoretical models for such an evolutionary change make the prediction that the mutations that created males must have sexually antagonistic effects, not only abolishing female functions, but also increasing male functions relative to the ancestral functional hermaphrodites. It is important to test this critical assumption. Moreover, the involvement of sexual antagonism also implies that plant sex-determining genes may directly cause some of the sexual dimorphisms observed in dioecious plants. Sex-determining genes are starting to be uncovered in plants, including species in the genera
<italic>Diospyros</italic>
and
<italic>Actinidia</italic>
(families Ebenaceae and Actinidiaceae, respectively). Here, we describe transgenic experiments in which the effects of the very different male-determining genes of these two dioecious species were studied in a non-dioecious plant,
<italic>Nicotiana tabacum</italic>
. The results indeed support the critical assumption outlined above.

DOI： 10.1098/rspb.2019.1805

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その他リンク： https://royalsocietypublishing.org/doi/full-xml/10.1098/rspb.2019.1805

掲載種別：研究論文（学術雑誌）   出版者・発行元：Springer Science and Business Media LLC  

DOI： 10.1007/s00122-019-03376-3

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その他リンク： http://link.springer.com/article/10.1007/s00122-019-03376-3/fulltext.html

記述言語：英語   掲載種別：研究論文（学術雑誌）  

DOI： 10.1038/s41477-019-0489-6

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記述言語：英語   掲載種別：研究論文（学術雑誌）  

DOI： 10.1111/tpj.14202

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掲載種別：研究論文（学術雑誌）   出版者・発行元：Elsevier BV  

DOI： 10.1016/j.scienta.2018.11.045

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記述言語：英語   掲載種別：研究論文（学術雑誌）  

The plant gibberellin (GA) receptor GID1 shows sequence similarity to carboxylesterase (CXE). Here, we report the molecular evolution of GID1 from establishment to functionally diverse forms in eudicots. By introducing 18 mutagenized rice GID1s into a rice gid1 null mutant, we identified the amino acids crucial for GID1 activity in planta. We focused on two amino acids facing the C2/C3 positions of ent-gibberellane, not shared by lycophytes and euphyllophytes, and found that adjustment of these residues resulted in increased GID1 affinity toward GA4, new acceptance of GA1 and GA3 carrying C13-OH as bioactive ligands, and elimination of inactive GAs. These residues rendered the GA perception system more sophisticated. We conducted phylogenetic analysis of 169 GID1s from 66 plant species and found that, unlike other taxa, nearly all eudicots contain two types of GID1, named A- and B-type. Certain B-type GID1s showed a unique evolutionary characteristic of significantly higher nonsynonymous-to-synonymous divergence in the region determining GA4 affinity. Furthermore, these B-type GID1s were preferentially expressed in the roots of Arabidopsis, soybean, and lettuce and might be involved in root elongation without shoot elongation for adaptive growth under low-temperature stress. Based on these observations, we discuss the establishment and adaption of GID1s during plant evolution.

DOI： 10.1073/pnas.1806040115

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：Oxford University Press  

The S-RNase-based gametophytic self-incompatibility (GSI) reproduction barrier is important for maintaining genetic diversity in species of the families Solanaceae, Plantaginaceae and Rosaceae. Among the plant taxa with S-RNase-based GSI, Prunus species in the family Rosaceae exhibit Prunus-specific self-incompatibility (SI). Although pistil S and pollen S determinants have been identified, the mechanism underlying SI remains uncharacterized in Prunus species. A putative pollen-part modifier was identified in this study. Disruption of this modifier supposedly confers self-compatibility (SC) to sweet cherry (Prunus avium) 'Cristobalina'. To identify the modifier, genome re-sequencing experiments were completed involving sweet cherry individuals from 18 cultivars and 43 individuals in two segregating populations. Cataloging of subsequences (35 bp kmers) from the obtained genomic reads, while referring to the mRNA sequencing data, enabled the identification of a candidate gene [M locus-encoded GST (MGST)]. Additionally, the insertion of a transposon-like sequence in the putative MGST promoter region in 'Cristobalina' down-regulated MGST expression levels, probably leading to the SC of this cultivar. Phylogenetic, evolutionary and gene expression analyses revealed that MGST may have undergone lineage-specific evolution, and the encoded protein may function differently from the corresponding proteins encoded by GST orthologs in other species, including members of the subfamily Maloideae (Rosaceae). Thus, MGST may be important for Prunus-specific SI. The identification of this novel modifier will expand our understanding of the Prunus-specific GSI system. We herein discuss the possible functions of MGST in the Prunus-specific GSI system.

DOI： 10.1093/pcp/pcy068

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記述言語：英語   出版者・発行元：Annual Reviews Inc.  

Dioecy, the presence of male and female flowers on separate individuals, is both widespread and uncommon within flowering plants, with only a few percent of dioecious species spread across most major phylogenetic taxa. It is therefore safe to assume that dioecy evolved independently in these different groups, which allows us to ask questions regarding the molecular and developmental mechanisms underlying these independent transitions to dioecy. We start this review by examining the problem from the standpoint of a genetic engineer trying to develop dioecy, discuss various potential solutions, and compare them to models proposed in the past and based on genetic and evolutionary considerations. Next, we present recent information regarding candidate sex determinants in three species, acquired using newly established genomic approaches. Although such specific information is still scarce, it is slowly becoming apparent that various genes or pathways can be altered to evolve dioecy.

DOI： 10.1146/annurev-arplant-042817-040615

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：American Society of Plant Biologists  

Dioecy, the presence of male and female flowers on distinct individuals, has evolved independently in multiple plant lineages, and the genes involved in this differential development are just starting to be uncovered in a few species. Here, we used genomic approaches to investigate this pathway in kiwifruits (genus Actinidia). Genome-wide cataloging of male-specific subsequences, combined with transcriptome analysis, led to the identification of a type-C cytokinin response regulator as a potential sex determinant gene in this genus. Functional transgenic analyses in two model systems, Arabidopsis thaliana and Nicotiana tabacum, indicated that this gene acts as a dominant suppressor of carpel development, prompting us to name it Shy Girl (SyGI). Evolutionary analyses in a panel of Actinidia species revealed that SyGI is located in the Y-specific region of the genome and probably arose from a lineage-specific gene duplication. Comparisons with the duplicated autosomal counterpart, and with orthologs from other angiosperms, suggest that the SyGI-specific duplication and subsequent evolution of cis-elements may have played a key role in the acquisition of separate sexes in this species.

DOI： 10.1105/tpc.17.00787

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：International Society for Horticultural Science  

Cultivated Diospyros kaki, otherwise known as persimmon or kaki, is a polygamous species, whereas most other Diospyros species are dioecious. Most commercial cultivars of D. kaki bear only female flowers
however, some cultivars bear male flowers as well. Elucidation of the genetic and molecular basis of sex expression in D. kaki may lead to the development of artificial control of sex expression in this important fruit species. To this end, Diospyros lotus was used as a model to study the sexuality of D. kaki. Using a population of 63 F1 offspring of D. lotus, a bulked segregant analysis with amplified fragment length polymorphisms (AFLP) showed that the sexuality of this species is controlled by a single locus or haploblock in a heterogametic male (XY) system. Male-specific subsequences from Illumina HiSeq genomic reads data obtained from male and female pools were cataloged and assembled. This enabled comprehensive isolation of the male-determining regions on the Y-chromosome of D. lotus. Integration of transcriptome and evolutionary analyses led to the identification of one Y-specific sex-determinant candidate gene, OGI. This gene encodes a small RNA that triggers a transitive RNAi on the feminizing gene, MeGI, which is located on the autosome or the pseudoautosomal region of the sex chromosome. Molecular markers developed from the OGI sequence and its flanking regions indicated that the genetic control of maleness expression in D. kaki is based essentially on the OGI/MeGI system.

DOI： 10.17660/ActaHortic.2018.1195.12

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：ELSEVIER SCIENCE BV  

There is considerable diversity in fruit shapes among persimmon (Diospyros kaki) cultivars. Although fruit shape is one of the most important traits affecting the commercial quality of fruits, quantitatively characterizing fruit shape features remains difficult. The objective of this study was to develop a reliable method for quantitatively analyzing persimmon fruit shapes, and to clarify the fruit development patterns as well as their relationships to the shapes of other organs across cultivars. Using 153 persimmon cultivars and two wild Diospyros relatives, we evaluated two-dimensional pictures of longitudinal and transverse fruit sections with elliptical Fourier descriptors (EFDs) and the SHAPE program. Principal component analysis (PCA) using each EFD generated representative vectors, which elucidated the fruit shape diversity among the cultivars. Additionally, a PCA of the EFDs quantified two major components of the variability in the shapes of longitudinal fruit sections, namely the ratio of length to diameter and the fruit apex shape. Seasonal characterizations of the first principal component revealed the fruit shape development patterns. The fundamental shapes in the longitudinal direction were coordinately determined during the early developmental stages across cultivars. Furthermore, a quantitative characterization of persimmon fruit, seed, and leaf shapes unveiled correlations between these traits. The data presented herein indicate that combining EFDs and PCA enables the quantitative analysis of persimmon organ shapes, which may be related to various quantitative indices.

DOI： 10.1016/j.scienta.2017.10.006

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記述言語：英語   出版者・発行元：GENETICS SOC JAPAN  

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：AMER SOC PLANT BIOLOGISTS  

Epigenetic regulation can add a flexible layer to genetic variation, potentially enabling long-term but reversible cis-regulatory changes to an allele while maintaining its DNA sequence. Here, we present a case in which alternative epigenetic states lead to reversible sex determination in the hexaploid persimmon Diospyros kaki. Previously, we elucidated the molecular mechanismof sex determination in diploid persimmon and demonstrated the action of a Y-encoded sex determinant pseudogene called OGI, which produces small RNAs targeting the autosomal gene MeGI, resulting in separate male and female individuals (dioecy). We contrast these findings with the discovery, in hexaploid persimmon, of an additional layer of regulation in the form of DNA methylation of the MeGI promoter associated with the production of both male and female flowers in genetically male trees. Consistent with this model, developing male buds exhibited higher methylation levels across the MeGI promoter than developing female flowers from either monoecious or female trees. Additionally, a DNA methylation inhibitor induced developing male buds to form feminized flowers. Concurrently, in Y-chromosome-carrying trees, the expression of OGI is silenced by the presence of a SINE (short interspersed nuclear element)-like insertion in the OGI promoter. Our findings provide an example of an adaptive scenario involving epigenetic plasticity.

DOI： 10.1105/tpc.16.00532

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：ELSEVIER SCIENCE BV  

The sexuality of crops is an important trait affecting production and breeding efficiencies. Recently, a Y-chromosome-encoded gene, designated Oppressor of MeGI (OGI), has been shown to play a central role in dioecious sex determination in diploid persimmons (Diospyros spp.). However, little is known about the relationship between OGI and the irregular sexuality in polyploid persimmon (D. kaki). D. kaki, which is mostly hexaploid with some nonaploid individuals, is mainly constituted of monoecious and female individuals. In this study, we attempted to investigate the association of sexuality with OGI allele diversities/dosages in a wide range of D. kaki cultivars. The results indicated that OGI is indispensable for the monoecious phenotype producing male flowers, which suggested a conserved role for OGI as a key to expressing maleness in D. kaki, as in dioecious diploid Diospyros. However, the existence of OGI alleles did not always lead to production of male flowers in D. kaki. The dosage or composition of OGI alleles had no substantial effect on the sexuality of the cultivars tested. We discuss the potential use of the obtained information on OGI allele diversities dosages for breeding of D. kaki cultivars. (C) 2016 Elsevier B.V. All rights reserved.

DOI： 10.1016/j.scienta.2016.10.046

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：OXFORD UNIV PRESS  

Self-incompatibility (SI) is an important plant reproduction mechanism that facilitates the maintenance of genetic diversity within species. Three plant families, the Solanaceae, Rosaceae and Plantaginaceae, share an S-RNase-based gametophytic SI (GSI) system that involves a single S-RNase as the pistil S determinant and several F-box genes as pollen S determinants that act via non-self-recognition. Previous evidence has suggested a specific self-recognition mechanism in Prunus (Rosaceae), raising questions about the generality of the S-RNase-based GSI system. We investigated the evolution of the pollen S determinant by comparing the sequences of the Prunus S haplotype-specific F-box gene (SFB) with those of its orthologs in other angiosperm genomes. Our results indicate that the Prunus SFB does not cluster with the pollen S of other plants and diverged early after the establishment of the Eudicots. Our results further indicate multiple F-box gene duplication events, specifically in the Rosaceae family, and suggest that the Prunus SFB gene originated in a recent Prunus-specific gene duplication event. Transcriptomic and evolutionary analyses of the Prunus S paralogs are consistent with the establishment of a Prunus-specific SI system, and the possibility of subfunctionalization differentiating the newly generated SFB from the original pollen S determinant.

DOI： 10.1093/pcp/pcw077

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：OXFORD UNIV PRESS  

Domestication and cultivar differentiation are requisite processes for establishing cultivated crops. These processes inherently involve substantial changes in population structure, including those from artificial selection of key genes. In this study, accessions of peach (Prunus persica) and its wild relatives were analysed genome-wide to identify changes in genetic structures and gene selections associated with their differentiation. Analysis of genome-wide informative single-nucleotide polymorphism loci revealed distinct changes in genetic structures and delineations among domesticated peach and its wild relatives and among peach landraces and modern fruit (F) and modern ornamental (O-A) cultivars. Indications of distinct changes in linkage disequilibrium extension/decay and of strong population bottlenecks or inbreeding were identified. Site frequency spectrum- and extended haplotype homozygosity-based evaluation of genome-wide genetic diversities supported selective sweeps distinguishing the domesticated peach from its wild relatives and each F/O-A cluster from the landrace clusters. The regions with strong selective sweeps harboured promising candidates for genes subjected to selection. Further sequence-based evaluation further defined the candidates and revealed their characteristics. All results suggest opportunities for identifying critical genes associated with each differentiation by analysing genome-wide genetic diversity in currently established populations. This approach obviates the special development of genetic populations, which is particularly difficult for long-lived tree crops.

DOI： 10.1093/dnares/dsw014

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記述言語：日本語   掲載種別：研究論文（学術雑誌）   出版者・発行元：園芸学会  

カキの性表現様式は雌雄混株であり，多くの栽培品種は雄花非着生形質を示し交雑育種の制限要素となっている．これまでに，カキの雌雄性の遺伝的機構は解明されていないが，最近，カキの近縁二倍体野生種であるマメガキにおいて，雄性形質連鎖領域DlSx-AF4Sが同定された．そこで本研究では，多くのカキ品種を供試して，DlSx-AF4Sとカキの雄花着生特性との関連性を検証した．3品種を除くすべての雄花着生品種でDlSx-AF4Sが検出されたことから，マメガキとカキの性決定に関与する遺伝制御因子が共通であることが示唆された．一方，雄花非着生品種の17％でマーカーが検出された．カキの性表現はマメガキと比較して環境要因や栄養状態，樹齢の影響を受けやすいことから，カキでは雄性決定因子をもっていても雄性形質を発現できない可能性が考えられた．また，マメガキで示されたように，カキ属の性決定はXY型であることが予想されるが，高次倍数性のカキでは，MSY上の雄性決定遺伝子の遺伝子量に多様性が生じる可能性があり，そのことが複雑な性表現様式に影響している可能性もある．今後は，DlSx-AF4Sに連鎖する雄性決定因子をカキより同定し解析することで，カキの複雑な性表現様式が解明されることが期待される．

DOI： 10.2503/hrj.14.121

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その他リンク： http://agriknowledge.affrc.go.jp/RN/2010892096

記述言語：英語   掲載種別：研究論文（学術雑誌）  

DOI： 10.1126/science.1257225

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：JAPAN SOC HORTICULTURAL SCI  

Sexuality of crops affects both cultivation and breeding systems. Cultivated persimmon (Diospyros kaki Thunb) has a morphologically well-characterized polygamous or gyonodioecious sexual system. However, the genetic basis of sexuality in D. kaki has yet to be characterized. Here, we used dioecious D. lotus L., a diploid wild relative species to hexaploid or nonaploid D. kaki, as a model to clarify the genetic basis of sexuality in Diospyros and to develop molecular markers associated with the sexuality of individuals. Using 62 F-1 offspring segregated into distinct male/female phenotypes, we found two amplified fragment-length polymorphism markers, DlSx-AF4 and DlSx-AF7, which cosegregated with maleness. This could suggest that the sexuality of D. lotus is controlled by a single gene/haploblock, and the male is dominant over the female. Thus, D. lotus's sexuality can be described as the heterogametic male type, the ICY-type, as reported for most other dioecious plant species. For unknown reasons, segregation of the phenotype of a sequence-characterized amplified region marker developed from DlSx-AF4 (DlSx-AF4S) and/or the male/female phenotype in two different crosses in D. lotus showed an apparent bias towards femaleness and better fitted 1:2 than 1:1, which is the theoretical segregation for a single genetic locus or haploblock in diploid D. lotus. DlSx-AF4S could distinguish D. kaki cultivars with female and male flowers from cultivars with only female flowers, strongly indicating that the same genetic system controls D. kaki's sexuality and that DlSx-AF4S could be used as a genetic marker for sexuality in D. kaki breeding programs.

DOI： 10.2503/jjshs1.CH-109

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記述言語：英語   掲載種別：研究論文（国際会議プロシーディングス）   出版者・発行元：INT SOC HORTICULTURAL SCIENCE  

For genetic analysis of persimmon (Diospyros kaki) and its relatives, we investigated the development of SSR markers with expression sequence tags (ESTs). Of 9475 EST data from the NCBI database which had isolated from D. kaki, 746 sequences contained SSRs which had more than four (tetranucleotide) or five (dinucleotide and trinucleotide) repeats. Tested on two species, D. kaki and D. lotus (a diploid relative of D. kaki), of primer pairs based on 65 SSR-containing EST sequences, 52 primer pairs resulted in proper PCR products. Some of these candidate EST-SSR markers showed polymorphism among cultivars and seedlings of D. kaki and D. lotus. Among tested seven clones of D. lotus, 32 of 52 candidate EST-SSR markers showed polymorphism. For molecular phylogenetic analysis, linkage mapping and whole genome mapping of Diospyros species, EST-SSR markers appear to be as useful as genomic SSR markers. In addition, development of EST-SSR markers seems to be easy and cost-effective when compared with genomic SSR markers.

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：SPRINGER HEIDELBERG  

Persimmon (Diospyros kaki Thunb.) is one of the major tree crops in East Asia and is generally hexaploid. A single ASTRINGENCY (AST) locus controls the astringency/non-astringency (A/NA) trait of persimmon fruit, one of the most important traits for consumption, on each of the six corresponding chromosomes. Although several molecular approaches are in progress to elucidate the molecular mechanisms of astringency trait in persimmon, the distinct polysomic behavior of the AST locus remains to be solved. The aim of this study was to perform fine genotyping of a highly polymorphic marker locus linked to the AST locus, detect the allele pairing in ten segregated F(1) lines derived from hybridization of A-type x NA-type cultivars, and identify the basis of hexaploid inheritance at the AST locus in persimmon. The results showed that persimmon cultivars frequently produce aneuploid offspring bearing an extra chromosome with the AST locus, with the incidence of aneuploidy varying among the cultivars. On the examination of hexasomic behavior in persimmon cultivars, the ratios of individuals bearing each allele pair segregated from A-type parents showed a good fit to the expected ratios in an autohexaploid inheritance model, except for cvs. Luo-tian-tian-shi and Sa-gok-shi which fitted to an autoallohexaploid inheritance model. These results suggest variable hexasomic behavior among persimmon cultivars.

DOI： 10.1007/s11295-011-0432-0

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：AMER SOC PLANT BIOLOGISTS  

Proanthocyanidins (PAs) are secondary metabolites that contribute to plant protection and crop quality. Persimmon (Diospyros kaki) has a unique characteristic of accumulating large amounts of PAs, particularly in its fruit. Normal astringent-type and mutant nonastringent-type fruits show different PA accumulation patterns depending on the seasonal expression patterns of DkMyb4, which is a Myb transcription factor (TF) regulating many PA pathway genes in persimmon. In this study, attempts were made to identify the factors involved in DkMyb4 expression and the resultant PA accumulation in persimmon fruit. Treatment with abscisic acid (ABA) and an ABA biosynthesis inhibitor resulted in differential changes in the expression patterns of DkMyb4 and PA biosynthesis in astringent-type and nonastringent-type fruits depending on the development stage. To obtain an ABA-signaling TF, we isolated a full-length basic leucine zipper (bZIP) TF, DkbZIP5, which is highly expressed in persimmon fruit. We also showed that ectopic DkbZIP5 overexpression in persimmon calluses induced the up-regulation of DkMyb4 and the resultant PA biosynthesis. In addition, a detailed molecular characterization using the electrophoretic mobility shift assay and transient reporter assay indicated that DkbZIP5 recognized ABA-responsive elements in the promoter region of DkMyb4 and acted as a direct regulator of DkMyb4 in an ABA-dependent manner. These results suggest that ABA signals may be involved in PA biosynthesis in persimmon fruit via DkMyb4 activation by DkbZIP5.

DOI： 10.1104/pp.111.191205

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：AMER SOC PLANT BIOLOGISTS  

Bud endodormancy in woody plants plays an important role in their perennial growth cycles. We previously identified a MADS box gene, DORMANCY-ASSOCIATED MADS box6 (PmDAM6), expressed in the endodormant lateral buds of Japanese apricot (Prunus mume), as a candidate for the dormancy-controlling gene. In this study, we demonstrate the growth inhibitory functions of PmDAM6 by overexpressing it in transgenic poplar (Populus tremula x Populus tremuloides). Transgenic poplar plants constitutively expressing PmDAM6 showed growth cessation and terminal bud set under environmental conditions in which control transformants continued shoot tip growth, suggesting the growth inhibitory functions of PmDAM6. In the Japanese apricot genome, we identified six tandemly arrayed PmDAM genes (PmDAM1-PmDAM6) that conserve an amphiphilic repression motif, known to act as a repression domain, at the carboxyl-terminal end, suggesting that they all may act as transcriptional repressors. Seasonal expression analysis and cold treatment in autumn indicated that all PmDAMs were repressed during prolonged cold exposure and maintained at low levels until endodormancy release. Furthermore, PmDAM4 to PmDAM6 responses to a short period of cold exposure appeared to vary between low-and high-chill genotypes. In the high-chill genotype, a short period of cold exposure slightly increased PmDAM4 to PmDAM6 expression, while in the low-chill genotype, the same treatment repressed PmDAM4 to PmDAM6 expression. Furthermore, PmDAM4 to PmDAM6 expression was negatively correlated with endodormancy release. We here discuss the genotype-dependent seasonal expression patterns of PmDAMs in relation to their involvement in endodormancy and variation in chilling requirements.

DOI： 10.1104/pp.111.181982

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記述言語：英語   出版者・発行元：ELSEVIER SCIENCE BV  

Persimmon accumulates a large amount of condensed tannins (proanthocyanidins, PAs) in the vacuoles of specific cells called &apos;tannin cells&apos; during fruit development. PAs cause astringency, which is a dry or puckering sensation due to the coagulation of oral proteins. The composition of PAs in persimmon fruit can be analysed mainly by two methods, thiolysis degradation and acid catalysis in the presence of phloroglucinol. These methods indicate that epigallocatechin (EGC) and epigallocatechin-3-O-gallate (EGCG) constitute the main subunit components of PAs in astringent-type fruit. PAs are synthesised via the phenylpropanoid and flavonoid biosynthetic pathways, which originate from substrates derived from the shikimate pathway. Subunits EGC and EGCG possibly derive from active transcription offlavonoid 3&apos;5&apos; hydroxylase and anthocyanidin reductase, and from gallic acid synthesised via the shikimate pathway for galloylation. The reason for the formation of gallates and transport systems into cell vacuoles specifically accumulating these compounds remains unknown, as well as the processes for polymerisation of fiavan 3-ol units. In addition, two Myb transcription factors, DkMyb2 and DkMyb4, which have the ability to activate the transcription of enzymatic genes, are involved in PA regulation in persimmon. In particular, DkMyb4 has been suggested to facilitate substantial PA biosynthesis in fruit. A better understanding of the mechanisms involved in PA biosynthesis in persimmon fruit will provide the basis for the breeding of non-astringent-type cultivars for fresh consumption, or for the genetic engineering of PAs for human benefit. (C) 2011 Elsevier B.V. All rights reserved.

DOI： 10.1016/j.scienta.2011.07.021

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：SPRINGER  

Persimmon fruits accumulate a large amount of proanthocyanidin (PA). Fruits of the mutant non-astringent (NA) type lose their ability to accumulate PA at an early stage of fruit development, whereas fruits of the normal astringent (A) type sustain PA accumulation until ripening. This allelotype is determined by the genotype of a single ASTRINGENCY (AST) locus. It is possible that the reduction in PA accumulation in NA-type fruits is due to phenological down-regulation of DkMyb4 (a PA regulator) and the resultant down-regulation of structural genes in the PA pathway. In this study, attempts were made to identify the regulatory mechanisms of phenological PA accumulation in A- and NA-type fruits, focusing particularly on the effects of ambient temperature. Continuous cool temperature conditions caused sustained expression of DkMyb4 in NA-type fruits, as well as in A-type fruits, resulting in increased expression of PA pathway genes and PA accumulation. However, the expression of some A/NA phenotypic marker genes was not significantly affected by the cool temperature conditions. In addition, PA composition in NA-type fruits exposed to cool temperatures differed from that in A-type fruits. These results indicate that a cool ambient temperature may have induced DkMyb4 expression and resultant PA accumulation, but did not directly affect the expression of the AST gene.

DOI： 10.1007/s00425-010-1346-z

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：JAPAN SOC HORTICULTURAL SCI  

Persimmon cultivars are classified into four types depending on the nature of astringency loss of the fruit, namely pollination constant non-astringent (PCNA), pollination variant non-astringent (PVNA), pollination variant astringent (PVA), and pollination constant astringent (PCA). Among these four types, PCNA is the most important cultivar for persimmon breeding due to the stable loss of natural astringency on the tree. The trait of natural astringency loss is recessive in Japanese PCNA cultivars, while that in the Chinese PCNA cultivar, 'Luo tian tian shi', is dominant and the latter locus, termed CPCNA, is different from that in Japanese cultivars. In order to develop a molecular marker for the selection of the CPCNA-type cultivar, we performed amplified fragment length polymorphism (AFLP) in combination with bulked segregant analysis for F-1 offspring derived from 'Luo tian tian shi', which was used as the maternal parent. A total of 384 primer combinations were tested, and three AFLP markers, namely EACT-MCCC-222 (RO1), EGGC-MCTC-309 (RO2), and EGCC-MCGA-105 (RO3), linked to the CPCNA dominant allele were obtained. Among these markers, EGGC/MCTC-309 (RO2) was converted into a sequence-characterized amplified region (SCAR) marker. PCR analysis using F-1 offspring (n = 264) revealed that the relevance ratio of the SCAR marker was 94%. The polymorphism of the RO2 marker, which was strongly linked to the CPCNA dominant allele, was detected in only two Chinese PCNA cultivars, namely 'Luo tian tian shi' and 'Tian bao gai', among the Chinese, Korean, and Japanese cultivars tested. These results indicate that the RO2 marker contributes to marker-assisted selection for breeding programs of new PCNA cultivars having the CPCNA trait.

DOI： 10.2503/jjshs1.80.150

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：SPRINGER  

Proanthocyanidins (PAs) are secondary metabolites that contribute to the protection of a plant against biotic and abiotic stresses. Persimmon (Diospyros kaki) accumulates abundant PAs in each plant organ, and some potential Myb-like transcription factors (Myb-TFs) involved in the production of PAs have been isolated. In this study, we aimed to molecularly characterize one of them, DkMyb2, which was placed in a subclade including a PA regulator of Arabidopsis (Arabidopsis thaliana), TRANSPARENT TESTA2 (TT2), and was co-induced with PA pathway genes after wound stress. Ectopic DkMyb2 overexpression caused significant up-regulation of PA pathway genes in transgenic persimmon calluses and significant accumulation of PA, and increased mean degree of polymerization of PAs in transgenic kiwifruit calluses. Analysis of the DNA-binding ability of DkMyb2 by electrophoretic mobility shift assays showed that DkMyb2 directly binds to the AC-rich cis-motifs known as AC elements in the promoters of the two PA pathway genes in persimmon, DkANR, and DkLAR. Furthermore, a transient reporter assay using a dual-luciferase system demonstrated direct transcriptional activation of DkANR and DkLAR by DkMyb2. We also discuss subfunctionalization of two PA regulators in persimmon, DkMyb2 and DkMyb4, as well as PA regulators in other plant species from the viewpoint of their ability to bind to cis-motifs and their functions in transcriptional activation. Our results provide insight into the multiple regulatory mechanisms that control PA metabolism by Myb-TFs in persimmon.

DOI： 10.1007/s00425-010-1241-7

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：JAPAN SOC HORTICULTURAL SCI  

Persimmon (Diospyros kaki Thunb.) accumulates soluble condensed tannin (CT) in fruit which is responsible for its astringency trait. In this study, we analyzed the CT composition in persimmon fruit with phloroglucinol, and identified and characterized novel epigallocatechin-3-O-gallate-phloroglucinol (EGCG-P) adducts as one of the main CT components of persimmon fruit. Analysis with phloroglucinol in persimmon cultivars revealed the different tendencies of the CT composition and component ratio among the four astringency types (PCNA, PVNA, PVA, and PCA), which are categorized by their patterns of astringency loss. The concentration of the main CT component in persimmon fruit, epigallocatechin-3-O-gallate (EGCG), was particularly different among astringency types. Further analysis of the fruit at various maturation stages will help in understanding the different mechanisms of CT accumulation among astringent types. Our results demonstrated that the phloroglucinol methodology is useful for CT composition analysis in persimmon fruit and will contribute to future studies on the astringency trait in this fruit.

DOI： 10.2503/jjshs1.79.275

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：JAPAN SOC HORTICULTURAL SCI  

In previous studies, we have developed molecular markers linked to the AST locus that controls fruit astringency type in persimmon; however, these markers are not feasible for practical application to breeding programs since they are not fully effective for discriminating the pollination constant and non-astringent (PCNA) genotype from the non-PCNA genotype in a progeny derived from 'Kurokuma'. Here we developed new sequence characterized amplified region (SCAR) markers that enable easy and reliable selection of the PCNA genotype from breeding populations. Genomic regions adjacent to probe 5R, which showed polymorphic fragments between PCNA and non-PCNA genotypes in restriction fragment length polymorphism (RFLP) analysis, were isolated from the genomic libraries of 'Nishimura-wase', 'Jiro', and 'Kurokuma'-derived offspring. The isolated genomic regions were characterized and 3 insertion/deletion mutations were observed between ast- and AST-linked regions. Several primers were designed in the flanking region of Indel-3 and, in multiplex PCR, it was shown that using 2 forward primers, AST-F and PCNA-F and a reverse primer, 5R3R, is the most useful and reliable primer set. The AST-linked 220-bp fragment proved to be a common marker of 'Kurokuma'-, 'Nisimura-wase'- and 'Aizumishirazu'- derived progenies. This multiplex PCR is considered the most practical tool for marker-assisted selection (MAS) and can enhance and accelerate progress in persimmon breeding.

DOI： 10.2503/jjshs1.79.150

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：AMER SOC HORTICULTURAL SCIENCE  

Persimmon (Diospyros kaki Thunb.) is generally hexaploid, and a single AST locus controls the pollination-constant non-astringency trait on each of six corresponding chromosomes. The pollination-constant non-astringent (PCNA) genotype is nulliplex and requires homozygous recessive alleles (ast) at the AST locus. There are several non-PCNA cultivars/selections that could be cross parents; however, the probability of yielding nulliplex offspring depends on the number of recessive alleles (ast). In genotyping for the AST locus in hexaploid persimmon, in contrast to the situation in diploid plants, we need to detect the AST/ast allele dosage; this cannot be detected by common codominant markers. In this study, we detected the allele dosage of M(ast), which is a marker allele strongly linked to the ast allele among cultivars, by quantitative real-time polymerase chain reaction (qPCR) using three reference sites, actin (DkAct), anthocyanin reductase (DkANR), and L5R, whose sequences are conserved in the genome of persimmon cultivars. Based on the allele dosage of the M(ast), AST/ast genotypes were estimated for 63 non-astringent cultivars/selections, of which only five cultivars/selections were estimated to be simplex or duplex. The quantitative genotyping method using qPCR may be generally effective for polyploid plants.

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：AMER SOC PLANT BIOLOGISTS  

Proanthocyanidins (PAs) are secondary metabolites that contribute to the protection of the plant and also to the taste of the fruit, mainly through astringency. Persimmon (Diospyros kaki) is unique in being able to accumulate abundant PAs in the fruit flesh. Fruits of the nonastringent (NA)-type mutants lose their ability to produce PA at an early stage of fruit development, while those of the normal astringent (A) type remain rich in PA until fully ripened. The expression of many PA pathway genes was coincidentally terminated in the NA type at an early stage of fruit development. The five genes encoding the Myb transcription factor were isolated from an A-type cultivar (Kuramitsu). One of them, DkMyb4, showed an expression pattern synchronous to that of the PA pathway genes in A-and NA-type fruit flesh. The ectopic expression of DkMyb4 in kiwifruit (Actinidia deliciosa) induced PA biosynthesis but not anthocyanin biosynthesis. The suppression of DkMyb4 in persimmon calluses caused a substantial down-regulation of the PA pathway genes and PA biosynthesis. Furthermore, analysis of the DNA-binding ability of DkMyb4 showed that it directly binds to the MYBCORE cis-motif in the promoters of the some PA pathway genes. All our results indicate that DkMyb4 acts as a regulator of PA biosynthesis in persimmon and, therefore, suggest that the reduction in the DkMyb4 expression causes the NA-type-specific down-regulation of PA biosynthesis and resultant NA trait.

DOI： 10.1104/pp.109.146985

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：SPRINGER  

Persimmon fruits accumulate a large amount of proanthocyanidin (PA) during development. Fruits of pollination-constant and non-astringent (PCNA) type mutants lose their ability to produce PA at an early stage of fruit development, while fruits of the normal (non-PCNA) type remain rich in PA until fully ripened. To understand the molecular mechanism for this difference, we isolated the genes involved in PA accumulation that are differentially expressed between PCNA and non-PCNA, and confirmed their correlation with PA content and composition. The expression of structural genes of the shikimate and flavonoid biosynthetic pathways and genes encoding transferases homologous to those involved in the accumulation of phenolic compounds were downregulated coincidentally only in the PCNA type. Analysis of PA composition using the phloroglucinol method suggested that the amounts of epigallocatechin and its 3-O-gallate form were remarkably low in the PCNA type. In the PCNA type, the genes encoding flavonoid 3&apos;5&apos; hydroxylase (F3&apos;5&apos;H) and anthocyanidin reductase (ANR) for epigallocatechin biosynthesis showed remarkable downregulation, despite the continuous expression level of their competitive genes, flavonoid 3&apos; hydroxylation (F3&apos;H) and leucoanthocyanidin reductase (LAR). We also confirmed that the relative expression levels of F3&apos;5&apos;H to F3&apos;H, and ANR to LAR, were considerably higher, and the PA composition corresponded to the seasonal expression balances in both types. These results suggest that expressions of F3&apos;5&apos;H and ANR are important for PA accumulation in persimmon fruit. Lastly, we tested enzymatic activity of recombinant DkANR in vitro, which is thought to be an important enzyme for PA accumulation in persimmon fruits.

DOI： 10.1007/s00425-009-0991-6

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：SPRINGER  

Fruits of persimmon (Diospyros kaki Thunb.) accumulate large amounts of proanthocyanidins (PAs) in the early stages of development. Astringent (A)-type fruits remain rich in soluble PAs even after they reach full-mature stage, whereas non-astringent (NA)-type fruits lose these compounds before full maturation. As a first step to elucidate the mechanism of PA accumulation in this non-model species, we used suppression subtractive hybridization to identify transcripts accumulating differently in young fruits of A- and NA-type. Interestingly, only a few clones involved in PA biosynthesis were identified in A-NA libraries. Represented by multiple clones were those encoding a novel 1-Cys peroxiredoxin and a new member of family 1 glycosyltransferases. Quantitative RT-PCR analyses confirmed correlation of the amount of PAs and accumulation of transcripts encoding these proteins in young persimmon fruits. Furthermore, the new family 1 glycosyltransferase was produced in Escherichia coli and shown to efficiently catalyze galactosylation at 3-hydroxyl groups of several anthocyanidins and flavonols. These findings suggest a complex mechanism of PA accumulation in persimmon fruits.

DOI： 10.1007/s00425-009-0989-0

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記述言語：英語   掲載種別：研究論文（学術雑誌）   出版者・発行元：SPRINGER HEIDELBERG  

Persimmon (Diospyros kaki Thunb.) is a polyploidy fruit tree species of economic importance to East Asia. Natural astringency loss is an important trait in persimmon breeding programs. Quantitative real-time PCR was used to determine the number of AST/ast alleles for fruit astringency in persimmon (D. kaki Thunb.). To this end, the cultivar Jiro was transformed with one or two copies of a gene encoding NADP-dependent sorbitol-6-phosphate dehydrogenase (S6PDH), which was used as a standard for measuring the allele number of a sequenced marker tightly linked to the recessive ast locus for nonastringency. Primers for markers linked to the AST or ast allele were then used to measure the AST to ast ratio directly in the progeny of a full-sib cross. From determination of the AST to ast ratio and the results of the S6PDH copy number, the number of AST and ast alleles at the AST/ast locus was estimated. This research supported the hypothesis that D. kaki is a hexaploid with six AST and/or ast alleles. In addition to the determination of the allelic status of the AST locus, the application of real-time PCR for confirmation of the ploidy level and allelic composition of target genes in autopolyploids or allopolyploids was demonstrated.

DOI： 10.1007/s11295-009-0202-4

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記述言語：英語   掲載種別：研究論文（国際会議プロシーディングス）   出版者・発行元：INT SOC HORTICULTURAL SCIENCE  

Pollination constant and non-astringent (PCNA) type persimmons, which lose their astringency naturally on the tree, are the most desirable for fresh fruit consumption. The trait of natural astringency-loss in PCNA type is recessive and controlled by a single locus, known as AST locus. Because PCNA type cultivars are of recent origin and exist only in a small number of cultivars, inbreeding depression resulting from repeated crosses among very few cultivars and/or selections is a major challenge facing the current breeding of new PCNA type cultivars. One solution to overcome this inbreeding depression is to include non-PCNA cultivars in the breeding project to extend the genetic pool by utilizing the wide diversity found in non-PCNA cultivars. However, to date, F-1 populations resulting from crosses between PCNA and non-PCNA type cultivars/selections have yielded only non-PCNA offspring. To obtain PCNA offspring, F-1 progeny needs to be backcrossed to PCNA type cultivars/selections. As persimmon is hexaploid, the obtainable rate of PCNA offspring in a backcross population is only 10-15%. Although we identified AFLP and RFLP markers tightly linked to the trait of natural astringency-loss in persimmon, these markers could not distinguish some non-PCNA progenies in other breeding populations. Previously, we demonstrated that RFLP markers derived from fosmid clones of D. lotus, diploid relatives of D. kaki, could be used to distinguish between PCNA and non-PCNA types of D. kaki, and established the possibility of extending this approach to PCR based markers. In this study, we describe the development of PCR-based markers derived from fosmid clones of D. kaki and D. lotus, and confirm their effectiveness across diverse progenies to select PCNA type.

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記述言語：英語   掲載種別：研究論文（国際会議プロシーディングス）   出版者・発行元：INT SOC HORTICULTURAL SCIENCE  

Persimmon fruit accumulates a quantity of proanthocyanidins (PAs) into "tannin cells" during development. In order to isolate genes involved in PA accumulation, suppression subtractive hybridization (SSH) libraries were constructed from astringent and non-astringent fruit as tester and driver populations for reciprocal SSH procedures. Three populations; (i) the fruit of F-1 individuals derived from the cross between Chinese and Japanese PCNA, (ii) BC1 individuals derived from the cross between F1 (Japanese PCNA x astringent cultivar) and Japanese PCNA, and (iii) astringency-removed fruit on the tree by ethanol treatment and untreated astringent fruit, were used for the SSH analysis. We identified several cDNA fragments which have similarity to flavonoid biosynthetic genes, such as PAL CHS, DFR and ANR. Additionally, we isolated gene fragments including F3GalTase, DHQ/SDH, SCPL, 1-CysPer and GST, which have not been identified for their role in PA biosynthesis. RNA gel blot analysis revealed that these genes were coincidentally expressed with PA accumulation during fruit development both in astringent and PCNA types. These genes, identified as being differentially expressed in astringent fruit by SSH, may help further study on the molecular mechanism of PA biosynthesis and its accumulation in tannin cells.

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記述言語：日本語   掲載種別：研究発表ペーパー・要旨（全国大会，その他学術会議）  

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記述言語：日本語   出版者・発行元：日本農芸化学会 ; 1962-  

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記述言語：日本語   出版者・発行元：植物化学調節学会  

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記述言語：日本語   出版者・発行元：日本園芸農業協同組合連合会  

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記述言語：英語   出版者・発行元：The Japanese Society for Horticultural Science  

© 2015 The Japanese Society for Horticultural Science (JSHS), All rights reserved.Flowering plants have developed a genetically determined self-incompatibility (SI) system to maintain genetic diversity within a species. The Solanaceae, the Rosaceae, and the Plantaginaceae have the S-RNase-based gametophytic SI (GSI) system, which uses S-RNase and F-box proteins as the pistil S and pollen S determinants, respectively. SI is associated with culture and breeding difficulties in rosaceous fruit trees, such as apple, pear, and stone fruit species; therefore, researchers in the pomology field have long studied the mechanism and genetics of SI in order to obtain clues to overcome these difficulties. Here, we investigated the evolutionary paths of the S-RNase genes by tracking their duplication patterns. Phylogenetic analysis and estimation of proxy ages for the establishment of S-RNase and its homologs in several rosaceous species showed that the divergence of S-RNase in the subtribe Malinae and the genus Prunus predated the gene in most recent common ancestors of Rosaceae species. Furthermore, the duplicated S-RNase-like genes were accompanied by duplicated pollen S-like F-box genes, suggesting segmental duplications of the S locus. Analysis of the expression patterns and evolutionary speeds of duplicated S-RNase-like genes in Prunus suggested that these genes have lost the SI recognition function, resulting in a single S locus. Furthermore, the S loci in the current Rosaceae species might have evolved independently from the duplicated S loci, which could explain the presence of genus-specific SI recognition mechanisms in the Rosaceae. The results of the present study should be valuable for the future development of artificial SI control and for self-compatible breeding in rosaceous horticultural plant species.

DOI： 10.2503/hortj.MI-060

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記述言語：英語   掲載種別：研究発表ペーパー・要旨（国際会議）   出版者・発行元：AMER SOC HORTICULTURAL SCIENCE  

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会議種別：口頭発表（招待・特別）  

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会議種別：公開講演，セミナー，チュートリアル，講習，講義等  

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会議種別：公開講演，セミナー，チュートリアル，講習，講義等  

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会議種別：公開講演，セミナー，チュートリアル，講習，講義等  

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会議種別：シンポジウム・ワークショップ パネル（指名）  

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会議種別：口頭発表（招待・特別）  

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会議種別：公開講演，セミナー，チュートリアル，講習，講義等  

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会議種別：口頭発表（基調）  

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会議種別：公開講演，セミナー，チュートリアル，講習，講義等  

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会議種別：シンポジウム・ワークショップ パネル（公募）  

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会議種別：公開講演，セミナー，チュートリアル，講習，講義等  

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会議種別：シンポジウム・ワークショップ パネル（指名）  

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会議種別：公開講演，セミナー，チュートリアル，講習，講義等  

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会議種別：シンポジウム・ワークショップ パネル（指名）  

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会議種別：公開講演，セミナー，チュートリアル，講習，講義等  

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会議種別：公開講演，セミナー，チュートリアル，講習，講義等  

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会議種別：シンポジウム・ワークショップ パネル（指名）  

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会議種別：公開講演，セミナー，チュートリアル，講習，講義等  

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会議種別：公開講演，セミナー，チュートリアル，講習，講義等  

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会議種別：公開講演，セミナー，チュートリアル，講習，講義等  

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会議種別：口頭発表（基調）  

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会議種別：シンポジウム・ワークショップ パネル（指名）  

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会議種別：シンポジウム・ワークショップ パネル（指名）  

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記述言語：英語   会議種別：口頭発表（基調）  

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会議種別：口頭発表（基調）  

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記述言語：英語   会議種別：口頭発表（招待・特別）  

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記述言語：英語   会議種別：口頭発表（招待・特別）  

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記述言語：英語   会議種別：口頭発表（招待・特別）  

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記述言語：英語   会議種別：口頭発表（招待・特別）  

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記述言語：英語   会議種別：口頭発表（招待・特別）  

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会議種別：口頭発表（招待・特別）  

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会議種別：口頭発表（招待・特別）  

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記述言語：英語   会議種別：口頭発表（招待・特別）  

researchmap

会議種別：口頭発表（招待・特別）  

researchmap

会議種別：シンポジウム・ワークショップ パネル（指名）  

researchmap

会議種別：口頭発表（招待・特別）  

researchmap

記述言語：英語   会議種別：口頭発表（招待・特別）  

researchmap

会議種別：口頭発表（招待・特別）  

researchmap

会議種別：口頭発表（基調）  

researchmap

会議種別：公開講演，セミナー，チュートリアル，講習，講義等  

researchmap