研究者総覧

佐藤 康 (サトウ ヤスシ)

  • 大学院理工学研究科 環境機能科学専攻 教授
Last Updated :2020/11/10

研究者情報

学位

  • 博士(理学)(東北大学)

ホームページURL

J-Global ID

研究キーワード

  • UDP-N-アセチルグルコサミン (UDP-GlcNAc)   環境応答   形態形成   細胞壁   管状要素分化   ラッカーゼ   ペルオキシダーゼ   リグニン合成   environmental response   morphogenesis   cell wall   tracheary element differentiation   laccase   peroxidase   lignin synthesis   

研究分野

  • ライフサイエンス / 植物分子、生理科学

経歴

  • 2020年10月 - 現在  愛媛大学大学院理工学研究科 環境機能科学専攻教授
  • 2007年04月 - 2020年09月  愛媛大学大学院理工学研究科 環境機能科学専攻准教授
  • 2006年04月 - 2007年03月  愛媛大学大学院理工学研究科 環境機能科学専攻助手
  • 2005年04月 - 2006年03月  愛媛大学理学部生物学科特任講師
  • 1996年04月 - 2005年03月  愛媛大学理学部生物地球圏科学科助手
  • 1995年06月 - 1996年03月  愛媛大学理学部生物学科助手

学歴

  • 1994年04月 - 1995年05月   東北大学大学院   理学研究科   生物学専攻 研究生
  • 1991年04月 - 1994年03月   東北大学大学院   理学研究科 博士後期課程   生物学専攻
  • 1989年04月 - 1991年03月   東北大学大学院   理学研究科 博士前期課程   生物学専攻
  • 1985年04月 - 1989年03月   東北大学   理学部   生物学科

所属学協会

  • American Society of Plant Biologists   日本植物学会   日本植物生理学会   中国四国植物学会   日本植物バイオテクノロジー学会   

研究活動情報

論文

  • Masato Nakamura, Tomoaki Kamehama, Yasushi Sato
    Plant Biotechnology 37 1 105 - 109 2020年03月 [査読有り]
     研究論文(学術雑誌)
  • Rebecca A. Smith, Mathias Schuetz, Steven D. Karlen, David Bird, Naohito Tokunaga, Yasushi Sato, Shawn D. Mansfield, John Ralph, A. Lacey Samuels
    Plant Physiology 174 2 1028 - 1036 2017年06月 [査読有り]
     研究論文(学術雑誌) 
    © 2017 American Society of Plant Biologists. All rights reserved. Many land plants evolved tall and sturdy growth habits due to specialized cells with thick lignified cell walls: tracheary elements that function in water transport and fibers that function in structural support. The objective of this study was to define how and when diverse cell populations contribute lignin precursors, monolignols, to secondary cell walls during lignification of the Arabidopsis (Arabidopsis thaliana) inflorescence stem. Previous work demonstrated that, when lignin biosynthesis is suppressed in fiber and tracheary element cells with thickened walls, fibers become lignin-depleted while vascular bundles still lignify, suggesting that nonlignifying neighboring xylem cells are contributing to lignification. In this work, we dissect the contributions of different cell types, specifically xylary parenchyma and fiber cells, to lignification of the stem using cell-type-specific promoters to either knock down an essential monolignol biosynthetic gene or to introduce novel monolignol conjugates. Analysis of either reductions in lignin in knockdown lines, or the addition of novel monolignol conjugates, directly identifies the xylary parenchyma and fiber cell populations that contribute to the stem lignification and the developmental timing at which each contribution is most important.
  • Sato Y, Goto S, Teraoka S, Takagaki K, Takehara A, Sano S, Sakakibara M
    Environments 4 2 40  2017年06月 [査読有り]
     研究論文(学術雑誌)
  • Nino A. Espinas, Koichi Kobayashi, Yasushi Sato, Nobuyoshi Mochizuki, Kaori Takahashi, Ryouichi Tanaka, Tatsuru Masuda
    FRONTIERS IN PLANT SCIENCE 7 1326  2016年08月 [査読有り]
     研究論文(学術雑誌) 
    Heme is involved in various biological processes as a cofactor of hemoproteins located in various organelles. In plant cells, heme is synthesized by two isoforms of plastid-localized ferrochelatase. FC1 and FC2. In this study, by characterizing Arabidopsis T-DNA insertional mutants, we showed that the allocation of heme is differentially regulated by ferrochelatase isoforms in plant cells. Analyses of weak (fc1-1) and null (fc1-2) mutants suggest that FC1-producing heme is required for initial growth of seedling development. In contrast, weak (fc2-1) and null (fc2-2) mutants of FC2 showed pale green leaves and retarded growth, indicating that FC2-producing heme is necessary for chloroplast development. During the initial growth stage, FC2 deficiency caused reduction of plastid cytochromes. In addition, although FC2 deficiency marginally affected the assembly of photosynthetic reaction center complexes, it caused relatively larger but insufficient light-harvesting antenna to reaction centers, resulting in lower efficiency of photosynthesis. In the later vegetative growth, however, fc2-2 recovered photosynthetic growth, showing that FC1-producing heme may complement the FC2 deficiency. On the other hand, reduced level of cytochromes in microsomal fraction was discovered in fc1-1, suggesting that FC1-producing heme is mainly allocated to extraplastidic organelles. Furthermore, the expression of FC1 is induced by the treatment of an elicitor flg22 while that of FC2 was reduced, and fo1-abolished the flg22-dependent induction of FC1 expression and peroxidase activity. Consequently, our results clarified that FC2 produces heme for the photosynthetic machinery in the chloroplast, while FC1 is the housekeeping enzyme providing heme cofactor to the entire cell. In addition, FC1 can partly complement FC2 deficiency and is also involved in defense against stressful conditions.
  • Keisuke Furo, Mamoru Nozaki, Hiroshi Murashige, Yasushi Sato
    FEBS LETTERS 589 21 3258 - 3262 2015年10月 [査読有り]
     研究論文(学術雑誌) 
    Uridine diphosphate-N-acetylglucosamine (UDP-GlcNAc) donates GlcNAc for various glycans and glycoconjugates. We previously found that GlcNAc supplementation increases the UDP-GlcNAc content in Arabidopsis; however, the metabolic pathway was undefined. Here, we show that the homolog of human GlcNAc kinase (GNK) is conserved in land plants. Enzymatic assays of the Arabidopsis homologous protein (AtGNK) revealed kinase activity that was highly specific for GlcNAc. We also demonstrate the role of AtGNK in plants by using its knockout mutant, which presents lower UDP-GlcNAc contents and is insensitive to GlcNAc supplementation. Moreover, our results demonstrate the presence of a GlcNAc salvage pathway in plants. (C) 2015 Federation of European Biochemical Societies. Published by Elsevier B.V. All rights reserved.
  • 榊原 正幸, 佐野 栄, 久保田 有紀, 佐藤康
    環境放射能除染学会誌 = Journal of the Society for Remediation of Radioactive Contamination in the Environment 2 1 13 - 18 環境放射能除染学会 2014年03月 [査読有り]
  • Chisato Takeuchi, Kouji Nagatani, Yasushi Sato
    JOURNAL OF PLANT RESEARCH 126 6 811 - 821 2013年11月 [査読有り]
     研究論文(学術雑誌) 
    We investigated the effect of elicitors on xylem differentiation and lignification using a Zinnia elegans xylogenic culture system. Water-soluble chitosan and a fungal elicitor derived from Botrytis cinerea were used as elicitors. Elicitor addition at the start of culturing inhibited tracheary element (TE) differentiation in a concentration-dependent manner, and 30 mu g mL(-1) of chitosan or 16.7 mu g mL(-1) of the fungal elicitor strikingly inhibited TE differentiation and lignification. Addition of chitosan (at 50 mu g mL(-1)) or the fungal elicitor (at 16.7 mu g mL(-1)) during the culturing period also inhibited TE differentiation without inhibiting cell division, except for immature TEs undergoing secondary wall thickening. Elicitor addition after immature TE appearance also caused the accumulation of an extracellular lignin-like substance. It appears that elicitor addition at the start of culturing inhibits the process by which dedifferentiated cells differentiate into xylem cell precursors. Elicitor addition during culturing also appears to inhibit the transition from xylem cell precursors to immature TEs, and induces xylem cell precursors or xylem parenchyma cells to produce an extracellular stress lignin-like substance.
  • Mamoru Nozaki, Munetaka Sugiyama, Jun Duan, Hiroshi Uematsu, Tatsuya Genda, Yasushi Sato
    PLANT CELL 24 8 3366 - 3379 2012年08月 [査読有り]
     研究論文(学術雑誌) 
    To study the regulatory mechanisms underlying lignin biosynthesis, we isolated and characterized lignescens (lig), a previously undescribed temperature-sensitive mutant of Arabidopsis thaliana that exhibits ectopic lignin deposition and growth defects under high-temperature conditions. The lig mutation was identified as a single base transition in GNA1 encoding glucosamine-6-phosphate N-acetyltransferase (GNA), a critical enzyme of UDP-N-acetylglucosamine (UDP-GlcNAc) biosynthesis. lig harbors a glycine-to-serine substitution at residue 68 (G68S) of GNA1. Enzyme activity assays of the mutant protein (GNA1(G68S)) showed its thermolability relative to the wild-type protein. The lig mutant exposed to the restrictive temperature contained a significantly smaller amount of UDP-GlcNAc than did the wild type. The growth defects and ectopic lignification of lig were suppressed by the addition of UDP-GlcNAc. Since UDP-GlcNAc is an initial sugar donor of N-glycan synthesis and impaired N-glycan synthesis is known to induce the unfolded protein response (UPR), we examined possible relationships between N-glycan synthesis, UPR, and the lig phenotype. N-glycans were reduced and LUMINAL BINDING PROTEIN3, a typical UPR gene, was expressed in lig at the restrictive temperature. Furthermore, treatment with UPR-inducing reagents phenocopied the lig mutant. Our data collectively suggest that impairment of N-glycan synthesis due to a shortage of UDP-GlcNAc leads to ectopic lignin accumulation, mostly through the UPR.
  • Yasushi Sato, Youko Yajima, Naohito Tokunaga, Ross Whetten
    BIOLOGIA 66 1 88 - 95 2011年02月 [査読有り]
     研究論文(学術雑誌) 
    Lignin is synthesized not only during morphogenesis of vascular plants but also in response to various stresses. Isolated Zinnia elegans mesophyll cells can differentiate into tracheary elements (TEs), and deposit lignin into cell walls in TE-inductive medium (D medium). Meanwhile isolated mesophyll cells cultured in hormone-free medium (Co medium) accumulate stress lignin-like substance during culture. Therefore this culture system is suitable for study of lignin and lignin-like substance formation. In D medium lignin was deposited in TEs, but in Co medium, extracellular lignin-like substance accumulated. Analysis of the culture media indicated the presence of dilignols in D culture, but not in Co culture. To investigate the fate of lignin precursors, we added coniferyl alcohol (CA) in each culture. In Co medium, CA was polymerized into dilignols rapidly but they were present only temporarily, and in D medium CA was polymerized into dilignols relatively slowly but their content increased continually. Meanwhile, in Co culture, peroxidase activity in the medium was much higher than the peroxidase activity bound ionically to the cell walls. In D culture, ionically bound peroxidase activity was higher than that in the medium. These results may suggest that lignin deposition in TEs is related to ionically bound peroxidases in D culture, and lignin-like substance deposition in the medium is related to peroxidases in the medium in Co culture.
  • Asahi Hiraga, Tsuyoshi Kaneta, Yasushi Sato, Seiichi Sato
    CELL BIOLOGY INTERNATIONAL 34 2 189 - 196 2010年02月 [査読有り]
     研究論文(学術雑誌) 
    Evans Blue staining indicated that actively growing tobacco BY-2 cells in the exponential phase died more rapidly than quiescent cells in the stationary phase when the cells cultured under agitation were placed under still conditions. Fifty percent cell death was induced at about 18, 26, 80 and 140 h for early, mid, late exponential- and stationary-phase cells, respectively. Actively growing cells became TUNEL (transferase-mediated dUTP nick end labelling)-positive more rapidly than quiescent cells, suggesting that the cell death evaluated by Evans Blue is accompanied by DNA cleavages. Electrophoresis of genomic DNA showed a typical 'DNA laddering' pattern formed by multiples of about 200 bp internucleosomal units. Chromatin condensation was first detected at least within 24 h by light microscopy, and then cell shrinkage followed. These findings suggest that the death of BY-2 cells induced by still conditions is POD (programmed cell death).
  • Naohito Tokunaga, Tsuyoshi Kaneta, Seiichi Sato, Yasushi Sato
    PHYSIOLOGIA PLANTARUM 136 2 237 - 249 2009年06月 [査読有り]
     研究論文(学術雑誌) 
    We have investigated the mechanism of lignification during tracheary element (TE) differentiation using a Zinnia elegans xylogenic culture. In the process, we isolated ZPO-C, a peroxidase gene of Z. elegans that is expressed specifically in differentiating TEs. ZPO-C is suggested to be involved in lignification of Z. elegans TEs in vivo and in vitro. Furthermore, a peroxidase gene of Arabidopsis thaliana (AtPrx66), which is homologous to ZPO-C, was identified. The expression profile and functions of the gene in planta remain to be investigated. In this study, we performed promoter::beta-glucuronidase (GUS) assays to investigate the expression profiles and functions of the ZPO-C-like peroxidases in A. thaliana. We generated transgenic A. thaliana lines carrying AtPrx66, AtPrx47 or AtPrx64 (peroxidases showing high sequence similarity to AtPrx66) promoter::GUS reporter gene fusions. The GUS activities of AtPrx66, AtPrx47 and AtPrx64 promoter::GUS lines were arranged concentrically from the center to the periphery in the roots of seedlings. Furthermore, histochemical GUS assays using inflorescence stems showed that AtPrx66, AtPrx47 and AtPrx64 promoter-driven GUS were mainly expressed in the differentiating vessels, xylem parenchyma and sclerenchyma, respectively. These results suggest that the gene expressions of these three peroxidases, which showed high sequence similarity to one another, are differentially regulated in various tissues and organs. In addition, our results suggest that while AtPrx66 and AtPrx47 are associated with lignification of vessels, AtPrx64 is associated with lignification of sclerenchyma.
  • Y. Makimoto, H. Yano, T. Kaneta, Y. Sato, S. Sato
    PROTOPLASMA 229 1 53 - 62 2006年11月 [査読有り]
     研究論文(学術雑誌) 
    Fibrillarin is known to play an important role in precursor ribosomal RNA processing and ribosome assembly. The present study describes a fibrillarin homolog gene isolated from tobacco BY-2 cells and its expression during the cell cycle. The cDNA for a fibrillarin homolog, named NtFib1, was first cloned in Nicotiana tabacum with degenerate primers. It encodes 314 amino acids and the deduced amino acid sequence has some highly conserved functional domains, such as the glycine and arginine-rich (GAR) domain for nucleolar localization and the RNA-binding motif. The C-terminal region is highly conserved and has 7 beta-sheets and 7 alpha-helices which are peculiar to fibrillarin. Thus, it is suggested that the fibrillarin homolog of this plant species functions in the same way as the fibrillarin already known from human and yeast cells. Northern blot analysis of BY-2 cells synchronized with aphidicolin or a combination of aphidicolin and propyzamide showed that the histone H4 gene was specifically expressed in the S phase but NtFib1 mRNA remained at high levels during the cell cycle. Examination of the localization of NtFib1 protein tagged with green-fluorescent protein (GFP) suggested that some persisting in the mitotic apparatus was eventually incorporated into reconstructed nucleoli in late telophase. Newly synthesized GFP-tagged NtFib1 protein in the cytoplasm was added to the recycled protein in early mitosis. Highly concentrated actinomycin D completely inhibited the transcription of genes coding for rRNA (rDNA) but did not significantly suppress the amount of either NtFib1 mRNA or protein, although the NtFib1 protein was reversibly dislocated from nucleoli. Although hypoxic shock completely prohibited rDNA transcription, NtFib1 mRNA remained at the same level as in the control experiment, even after the 4 h treatment. These results indicate that the transcription of NtFib1 mRNA is not related to rDNA transcription and NtFib1 mRNA is resistant to disrupting factors during the cell cycle.
  • N. Kuya, M. Kato, Y. Sato, T. Kaneta, S. Sato
    PROTOPLASMA 229 1 83 - 91 2006年11月 [査読有り]
     研究論文(学術雑誌) 
    The cellular structures of statocytes implicated in gravisensing in primary and lateral roots of Vigna angularis were compared. The statocytes of lateral roots already had small amyloplasts immediately after they emerged from the primary root. Although these amyloplasts sedimented, the lateral roots showed much weaker gravitropism than primary roots, at least until they reached a length of about 30 mm. The nuclei were usually positioned in the upper end of the statocytes in both types of roots. Electron microscopic surveys showed that many tubular elements of endoplasmic reticulum (ER) were frequently localized in the lower end of the statocyte and they sometimes diverged or curved, suggesting that the ER forms a large reticulate complex. It is worth noting that statocytes with a large ER complex were found much more frequently in primary roots than in lateral roots. The amyloplasts were not always settled on this complex but were very frequently under it, especially in the primary roots. In lateral roots, they were usually localized under the ER complex when they were present. Thus, it is suggested that the differential development and organization of the amyloplast-ER complex system is involved in the differential gravitropism of the two types of roots.
  • Yasushi Sato, Ross W. Whetten
    JOURNAL OF PLANT RESEARCH 119 6 581 - 588 2006年11月 [査読有り]
     研究論文(学術雑誌) 
    We previously showed that eight laccase genes (Lac 1-Lac 8) are preferentially expressed in differentiating xylem and are associated with lignification in loblolly pine (Pinus taeda) [Sato et al. (2001) J Plant Res 114:147-155]. In this study we generated transgenic tobacco suspension cell cultures that express the pine Lac 1 and Lac 2 proteins, and characterized the abilities of these proteins to oxidize monolignols. Lac 1 and Lac 2 enzymatic activities were detected only in the cell walls of transgenic tobacco cells, and could be extracted with high salt. The optimum pH for laccase activity with coniferyl alcohol as substrate was 5.0 for Lac 1 and between 5.0 and 6.0 for Lac 2. The activities of Lac 1 and Lac 2 increased as the concentration of CuSO4 in the reaction mixtures increased in the range from 1 to 100 mu M. Both enzymes were able to oxidize coniferyl alcohol and to produce dimers of coniferyl alcohol. These results are consistent with the hypothesis that Lac 1 and Lac 2 are involved in lignification in differentiating xylem of loblolly pine.
  • Naohito Tokunaga, Nami Uchimura, Yasushi Sato
    PROTOPLASMA 228 4 179 - 187 2006年09月 [査読有り]
     研究論文(学術雑誌) 
    Gibberellin (GA) is considered an important growth regulator involved in many aspects of plant development. However, little is known about the relationship between GA and lignification. In this study, we analyzed the role of GA in tracheary element (TE) differentiation and lignification using a Zinnia elegans xylogenic culture. When gibberellic acid-3 (GA(3)) was exogenously supplied, a slight increase in the frequency of TE differentiation and a remarkable increase in lignin content were observed. Computer image analysis of individual TEs showed that the lignification level of each TE was significantly increased in the culture treated with GA(3) compared with those of the control. In contrast, suppression of TE differentiation and lignification was observed when GA biosynthesis was inhibited by ancymidol, paclobutrazol, or uniconazole. This suppression was restored by the addition of GA(3). These results suggest that GA plays an important role in TE differentiation, and even more so in lignification. When conditioned medium obtained after 120 h of control culture was analyzed by high-performance liquid chromatography, many lignin precursors were detected. However, these lignin precursors were greatly reduced in the GA-treated culture. This result suggests that GA promotes lignification by activating the polymerization of lignin precursors. Gibberellin (GA) is considered an important growth regulator involved in many aspects of plant development. However, little is known about the relationship between GA and lignification. In this study, we analyzed the role of GA in tracheary element (TE) differentiation and lignification using a Zinnia elegans xylogenic culture. When gibberellic acid-3 (GA(3)) was exogenously supplied, a slight increase in the frequency of TE differentiation and a remarkable increase in lignin content were observed. Computer image analysis of individual TEs showed that the lignification level of each TE was significantly increased in the culture treated with GA(3) compared with those of the control. In contrast, suppression of TE differentiation and lignification was observed when GA biosynthesis was inhibited by ancymidol, paclobutrazol, or uniconazole. This suppression was restored by the addition of GA(3). These results suggest that GA plays an important role in TE differentiation, and even more so in lignification. When conditioned medium obtained after 120 h of control culture was analyzed by high-performance liquid chromatography, many lignin precursors were detected. However, these lignin precursors were greatly reduced in the GA-treated culture. This result suggests that GA promotes lignification by activating the polymerization of lignin precursors.
  • Y Sato, T Demura, K Yamawaki, Y Inoue, S Sato, M Sugiyama, H Fukuda
    PLANT AND CELL PHYSIOLOGY 47 4 493 - 503 2006年04月 [査読有り]
     研究論文(学術雑誌) 
    In an attempt to elucidate the regulatory mechanism of vessel lignification, we isolated ZPO-C, a novel peroxidase gene of Zinnia elegans that is expressed specifically in differentiating tracheary elements (TEs). The ZPO-C transcript was shown to accumulate transiently at the time of secondary wall thickening of TEs in xylogenic culture of Zinnia cells. In situ hybridization indicated specific accumulation of the ZPO-C transcript in immature vessels in Zinnia seedlings. Immunohistochemical analysis using anti-ZPO-C antibody showed that the ZPO-C protein is abundant in TEs, especially at their secondary walls. For enzymatic characterization of ZPO-C, 6xHis-tagged ZPO-C was produced in tobacco cultured cells and purified. The ZPOC:6xHis protein had a peroxidase activity preferring sinapyl alcohol as well as coniferyl alcohol as a substrate, with a narrow pH optimum around 5.25. The peroxidase activity required calcium ion and was elevated by increasing Ca2+ concentration in the range of 0-10 mM. An Arabidopsis homolog of ZPO-C, At5g51890, was examined for expression patterns with transgenic plants carrying a yellow fluorescent protein (YFP) gene under the control of the A15g51890 promoter. The YFP fluorescence localization demonstrated vessel-specific expression of A15g51890 in the Arabidopsis roots. Taken collectively, our results strongly suggest that ZPO-C and its homologs play an important role in lignifcation of secondary cell walls in differentiating TEs.
  • S Sato, H Yano, Y Makimoto, T Kaneta, Y Sato
    JOURNAL OF PLANT RESEARCH 118 2 71 - 81 2005年04月 [査読有り]
     
    The nucleolus is the most obvious structure in the eukaryotic nucleus. It is known to be a ribosome-producing apparatus where ribosomal (r) DNA is transcribed and the primary rRNA transcripts are processed to produce three of the four rRNA species. Electron microscopy has shown that the nucleolus consists of three major components, a dense fibrillar component (DFC), a granular component (GC) and a fibrillar center (FC). The DFC and FCs are integrated into a fundamental nucleolar substructure called the nucleolonema. The DFC corresponds to the matrix of the nucleolonema, and the FC is an electron microscopic counterpart of argyrophobic lacunae localized in the nucleolonema. The spherical FCs are intermittently arranged along the length of the nucleolonema in actively growing cells but are fused with each other to form tubular FCs when rDNA transcription is hampered. The RNase-gold complex does not bind to the FC but to the DFC and the GC, suggesting that rDNA transcription does not occur in the FC although both fluorescence in situ hybridization (FISH) and electron microscopic in situ hybridization reveal that the rDNA is specifically localized in the FCs. Immunogold-labeling after bromo-UTP (BrUTP) incorporation shows that rDNA transcription takes place in the boundary region between the FC and the DFC, and primary rRNA transcripts are expected to be processed outward within the DFC. Data have accumulated suggesting that the nucleolonema is a fundamental substructure of the nucleolus, and its skeleton is the tandem arrangement of the FCs, which are resting harbors or storages of rDNA. Ibis paper proposes that the transversal structural organization of the nucleolonema is centrifugally built up by several structural and functional domains: condensed and/or loosened rDNA, rDNA transcription zone, and transcript processing and ribosome assembly zones.
  • N Tokunaga, N Sakakibara, T Umezawa, Y Ito, H Fukuda, Y Sato
    PLANT AND CELL PHYSIOLOGY 46 1 224 - 232 2005年01月 [査読有り]
     研究論文(学術雑誌) 
    During differentiation of isolated Zinnia mesophyll cells into tracheary elements (TEs), lignification on TEs progresses by supply of monolignols not only from TEs themselves but also from surrounding xylem parenchymalike cells through the culture medium. However, how lignin polymerizes from the secreted monolignols has not been resolved. In this study, we analyzed phenol compounds in culture medium with reversed-phase HPLC, gas chromatography-mass spectrometry and nuclear magnetic resonance spectrometry, and found 12 phenolic compounds including coniferyl alcohol and four dilignols, i.e. erythroguaiacylglycerol-beta-coniferyl ether, threo-guaiacylglycerol-beta-coniferyl ether, dehydrodiconiferyl alcohol and pinoresinol, in the medium in which TEs were developing. Coniferyl alcohol applied to TE-inductive cultures during TE formation rapidly disappeared from the medium, and caused a sudden increase in dilignols. Addition of the dilignols promoted lignification of TEs in which monolignol biosynthesis was blocked by an inhibitor of phenylalanine anmmonia-lyase (PAL), L-alpha-aminooxy-beta-phenylpropionic acid (AOPP). These results suggested that dilignols can act as intermediates of lignin polymerization.
  • Yasuko Ito, Naohito Tokunaga, Yasushi Sato, Hiroo Fukuda
    Plant Biotechnology 21 3 205 - 213 2004年09月 [査読有り]
     研究論文(学術雑誌) 
    In order to understand cell-cell interactions involved in xylem differentiation, we studied intercellular molecules in an in vitro Zinnia xylogenic culture system, where single mesophyll cells transdifferentiate into tracheary elements (TEs) and xylem parenchyma cells. We found that UV-absorbing substances accumulated predominantly in xylogenesis - inducing medium and kept increasing even after the TEs died. This accumulation was inhibited by L- α -aminooxy- β - phenylpropionic acid (AOPP), an inhibitor of phenylalanine ammonia-lyase, and also by brefeldin A, an inhibitor of vesicle transport. These results indicated that living non-TE cells, probably xylem parenchyma cells, secrete some kinds of phenylpropanoids via a vesicle transport system. Further experiment showed that inhibition of brassinosteroid biosynthesis by uniconazole suppressed TE differentiation, but not the secretion of UV-absorbing substances into the medium, implying that differentiation of xylem parenchyma cells might not be strongly affected by the depletion of endogenous brassinosteroids.
  • M Hosokawa, S Suzuki, T Umezawa, Y Sato
    PLANT AND CELL PHYSIOLOGY 42 9 959 - 968 2001年09月 [査読有り]
     研究論文(学術雑誌) 
    Tracheary element (TE) differentiation is a typical example of programmed cell death (PCD) in higher plants, and maturation of TEs is completed by degradation of all cell contents. However, lignification of TEs progresses even after PCD. We investigated how and whence monolignols are supplied to TEs which have undergone PCD during differentiation of isolated Zinnia mesophyll cells into TEs. Higher densities of cell culture induced greater lignification of TEs. Whereas the continuous exchanging of culture medium suppressed lignification of TEs, further addition of coniferyl alcohol into the exchanging medium reduced the suppression of lignification. Analysis of the culture medium by HPLC and GC-MS showed that coniferyl alcohol, coniferaldehyde, and sinapyl alcohol accumulated in TE inductive culture. The concentration of coniferyl alcohol peaked at the beginning of secondary wall thickening, decreased rapidly during secondary wall thickening, then increased again. These results indicated that lignification on TEs progresses by supply of monolignols from not only TEs themselves but also surrounding xylem parenchyma-like cells through medium in vitro.
  • Y Sato, WL Bao, R Sederoff, R Whetten
    JOURNAL OF PLANT RESEARCH 114 1114 147 - 155 2001年06月 [査読有り]
     研究論文(学術雑誌) 
    Eight cDNA clones for laccase (Lac 1-Lac 8) were isolated and characterized for analysis of the role of laccases in xylem formation by molecular and genetic approaches. Five of them (Lac 1-5) were isolated from cDNA libraries derived from differentiating xylem through a cDNA sequencing project in loblolly pine xylem (Allona of al. 1998) and other three (Lac 6-8) were isolated using a probe from a tobacco blue copper oxidase. Analysis of the cDNAs revealed that the proteins predicted had N-terminal signal sequences, 8-21 glycosylation sites, and four copper binding sites. The putative mature laccases range between 59.2 and 61.7 kD. The predicted isoelectric points vary between 7.3 and 9.9. Phylogenetic analysis shows these laccase cDNAs form three clusters in the cationic plant laccase group. Northern analysis indicated that all eight of the laccase transcripts are most abundant in differentiating xylem of the six tissues and organs tested. The Lad transcript is also detectable in immature pollen cones, and the Lac 7 transcript is detectable in several organs tested. The predominance of transcript abundance in differentiating xylem suggests that laccases play an important role in pine xylem development, consistent with previous hypotheses that laccases are involved in lignin synthesis in differentiating xylem.
  • Y Sato, T Watanabe, A Komamine, T Hibino, D Shibata, M Sugiyama, H Fukuda
    PLANT PHYSIOLOGY 113 2 425 - 430 1997年02月 [査読有り]
     研究論文(学術雑誌) 
    Changes in the enzymatic activity of cinnamyl alcohol dehydrogenase (CAD) and in the expression of a gene for CAD during tracheary element (TE) differentiation were investigated in cultures of single cells isolated from the mesophyll of zinnia (Zinnia elegans). In cultures in which TE differentiation was induced (TE-inductive cultures), CAD activity increased from h 36 after the start of culture (12 h before the start of thickening of the secondary cell wall) and peaked at h 72, when lignin was actively being deposited. In control cultures in which TE differentiation was not induced, CAD activity remained at a very low level for 5 d. Some isoforms of CAD were detected only in the TE-inductive cultures by native gel electrophoresis and subsequent staining for CAD activity. A cDNA clone for CAD, ZCAD1, was isolated from Z. elegans using a cDNA clone for CAD from Aralia cordata as the probe. RNA gel-blot analysis revealed that in the TE-inductive cultures the level of ZCAD1 mRNA increased from h 36 and peaked at h 38 to 60. No such increases were observed in control cultures. These results indicated that both the gene expression and the activity of CAD are strictly regulated, in association with lignification, during TE differentiation in Z. elegans.
  • Y Sato, M Sugiyama, T Takagi, H Fukuda
    JOURNAL OF PLANT RESEARCH 108 1092 463 - 468 1995年12月 [査読有り]
     研究論文(学術雑誌) 
    Cell wall-bound and tracheary element-specific peroxidase isoenzymes, designated P5A and P5B, were shown previously to be associated with lignification during the differentiation into tracheary elements of single cells isolated from the mesophyll of Zinnia elegans (Sate et al. Planta 189: 584-589, 1993; Planta 196: 141-147, 1995). Isoenzymes corresponding to P5 (RP5A and RP5B) were present at a relatively high level in the roots of Zinnia elegans. These isoenzymes were purified from the Zinnia roots by several column-chromatographic steps. Both RP5A and RP5B had molecular masses of 35 kDa. purified RP5A and RP5B were cleaved by CNBr and the partial amino acid sequences of these isoenzymes were determined.
  • Yasushi Sato, Munetaka Sugiyama, Atsushi Komamine, Hiroo Fukuda
    Planta: An International Journal of Plant Biology 196 1 141 - 147 1995年 [査読有り]
     研究論文(学術雑誌) 
    Cell wall-bound peroxidase (EC 1.11.1.7) isoenzymes (P1-P5) from cells of Zinnia elegans L. that were differentiating into tracheary elements were separated and characterized to obtain information about the relationships between these isoenzymes and the biosynthesis of lignin. Fractionation of Zinnia cells by centrifugation in solutions of Percoll revealed that P1, P2, and P5 were present in differentiated tracheary elements. These peroxidase isoenzymes were separated by several column-chromatographic steps. During hydrophobic chromatography on Phenyl Superose, P5 activity was separated into activities P5A and P5B. Enzymatically pure preparations of P1, P3, P5A, and P5B were finally obtained and used for the characterization of each isoenzyme. The optimum pH was 5.5–6.0 for P1, 5.0–7.5 for P3, 5.0 for P5A, and 4.0 for P5B. Each of the isoenzymes oxidized coniferyl alcohol efficiently, whereas p-coumaryl alcohol and sinapyl alcohol were poor substrates for all the isoenzymes. An absolute requirement for Ca2+ ions was demonstrated for P3. Based on these results, possible roles of peroxidase isoenzymes in the formation of lignin during the differentiation of tracheary elements are discussed. © 1995, Springer-Verlag. All rights reserved.
  • H FUKUDA, Y SATO, T YOSHIMURA, T DEMURA
    JOURNAL OF PLANT RESEARCH Special Issue 3 97 - 107 1993年07月 [査読有り]
     研究論文(学術雑誌)
  • Yasushi Sato, Munetaka Sugiyama, Ryszard J. Górecki, Hiroo Fukuda, Atsushi Komamine
    Planta 189 4 584 - 589 1993年04月 [査読有り]
     研究論文(学術雑誌) 
    In a culture system in which single cells isolated from the mesophyll of Zinnia elegans L. differentiate to tracheary elements (TEs), two inhibitors of phenylalanine ammonia-lyase (EC 4.3.1.5), L-α-aminooxy-β-phenylpropionic acid (AOPP) at 10 μM inhibited lignification without reducing the number of TEs formed. These inhibitors caused intracellular changes in peroxidase (EC 1.11.1.7) activities. The inhibitors increased the activity of peroxidases bound to the cell walls and especially the activity of peroxidase bound ionically to the cell walls. In contrast, the activity of extracellular peroxidase decreased. There were five isoenzymes, P1-P5, in the ionically bound peroxidase of cultured Zinnia cells. Among the isoenzymes, P4 and P5 appeared to be specific for TE differentation. Treatment with AOPP and AIP resulted in increases in the activities of P2, P4 and P5 isoenzymes, with the most prominent increase in P5 activity. The addition of lignin precursors, including coniferyl alcohol, to the AOPP-treated cells restored lignification, and suppressed the alteration of peroxidase isoenzyme patterns caused by AOPP. The relationship between the wall-bound peroxidases and lignification during TE differentiation is discussed in the light of these results. © 1993 Springer-Verlag.

書籍

  • 福島 和彦, 船田 良, 杉山 淳司, 高部 圭司, 梅澤 俊明, 山本 浩之 (担当:分担執筆, 範囲:細胞の分化とリグニン形成の分子機構)
    海青社 2011年10月 ISBN: 4860992520 321-337
  • Immunoelectron Microscopy. Methods and Protocols.
    佐藤 成一, 佐藤 康 (担当:分担執筆, 範囲:Localization of rDNA at nucleolar structural components by immunoelectron microscopy.)
    Springer 2010年 ISBN: 9781607617839 287-296
  • 福島 和彦, 船田 良, 杉山 淳司, 高部 圭司, 梅澤 俊明, 山本 浩之 (担当:分担執筆, 範囲:細胞の分化とリグニン形成の分子機構)
    海青社 2003年03月 ISBN: 4860992024 208-218
  • Molecular Breeding of Woody Plants
    佐藤 康 (担当:分担執筆, 範囲:Spatial and temporal regulation of lignifcation during tracheary element differentiation.)
    Elsevier 2001年 ISBN: 9780080536750 19-28

講演・口頭発表等

  • ヘキソサミン経路の植物特異的な機能解明に向けた,ホスホアセチルグルコサミンムターゼの酵素学的解析  [通常講演]
    作田 悠太郎, 佐藤 康
    日本植物学会第84回大会 2020年09月 ポスター発表
  • Analysis of physiological functions of N-acetylglucosamine transporter in Arabidopsis thaliana
    Eka Nurhangga, 佐藤 康
    日本植物学会第84回大会 2020年09月 ポスター発表
  • シロイヌナズナにおけるヘキソサミン経路が環境ストレス耐性に果たす役割の解析  [通常講演]
    的場 洋佑, 足立 楓, 佐藤 康
    日本植物学会第84回大会 2020年09月 口頭発表(一般)
  • シロイヌナズナにおけるヘキソサミン経路が環境ストレス耐性に果たす役割  [通常講演]
    的場 洋佑, 足立 楓, 佐藤 康
    第61回 日本植物生理学会年会 2020年03月
  • Analysis of physiological functions of N-acetylglucosamine (GlcNAc) transporter in Arabidopsis  [通常講演]
    Eka Nurhangga, 佐藤 康
    日本植物学会第83回大会 2019年09月 ポスター発表
  • シロイヌナズナGlcNAc キナーゼ破壊変異体を用いた,GlcNAc 再生経路が植物の生存に果たす役割の解明  [通常講演]
    財津 良多, 佐藤 康
    日本植物学会第83回大会 2019年09月 ポスター発表
  • シロイヌナズナのヘキソサミン経路に関わるホスホアセチルグルコサミンムーゼの酵素学的解析  [通常講演]
    作田 悠太郎, 佐藤 康
    日本植物学会第83回大会 2019年09月 口頭発表(一般)
  • Functions of GlcNAc salvage pathway for survival of plants – Physiological analysis using a knock-out mutant of GlcNAc kinase of Arabidopsis –  [通常講演]
    財津 良多, 佐藤 康
    IUFRO Tree Biotechnology 2019 2019年06月 ポスター発表
  • シロイヌナズナのヘキソサミン経路に関わるホスホアセチルグルコサミンムターゼの酵素学的・機能学的解析  [通常講演]
    作田悠太郎, 佐藤 康
    生物系三学会中国四国支部大会広島大会 2019年05月 口頭発表(一般)
  • Analysis of the function of N-acetylglucosamine (GlcNAc) transporters in Arabidopsis thaliana  [通常講演]
    Eka Nurhangga, 越澤 慶将, 佐藤 康
    生物系三学会中国四国支部大会広島大会 2019年05月 ポスター発表
  • シロイヌナズナにおけるヘキソサミン経路の生理機能 〜UDP-GalNAcの生成及びストレス応答リグニン蓄積の解析〜  [通常講演]
    亀浜 朝瑛, 中村 優斗, 佐藤 康
    生物系三学会中国四国支部大会広島大会 2019年05月 ポスター発表
  • GlcNAc再生経路が植物の生存に果たす役割 〜シロイヌナズナGlcNAcキナーゼ破壊変異体を用いた生理学的解析〜  [通常講演]
    財津 良多, 佐藤 康
    生物系三学会中国四国支部大会広島大会 2019年05月 ポスター発表
  • 小胞体ストレスによるシロイヌナズナの根の伸長抑制及びリグニン異常蓄積へのTHESEUS1の役割  [通常講演]
    中村優斗, 佐藤 康
    日本植物学会第82回大会 2018年09月 ポスター発表
  • GlcNAc再生経路が担う生存への役割 -シロイヌナズナGlcNAcキナーゼ破壊変異体を用いた生理学的解析-  [通常講演]
    財津良多, 佐藤 康
    日本植物学会第82回大会 2018年09月 口頭発表(一般)
  • シロイヌナズナのlig変異体における変異型GNA1の熱不安定性に起因する温度依存的小胞体ストレス応答とDMSOの抑圧効果  [通常講演]
    野崎守, 中村優斗, 佐藤康, 杉山宗隆
    Biothermology Workshop 2017 2017年12月 シンポジウム・ワークショップパネル(公募)
  • 植物におけるGlcNAcキナーゼの生理機能の解明-シロイヌナズナGlcNAcキナーゼ破壊変異体の表現型解析-  [通常講演]
    財津良多, 佐藤 康
    日本植物学会第81回大会 2017年09月 口頭発表(一般)
  • UDP-GlcNAc再生に関わるシロイヌナズナGlcNAcキナーゼの生理機能の解析  [通常講演]
    財津良多, 佐藤 康
    生物系三学会中国四国支部大会高知大会 2017年05月 口頭発表(一般)
  • タバコ培養細胞BY-2の静置培養誘導PCDにおける液胞および小胞体ストレスとの関連性  [通常講演]
    平賀旭, 佐藤 康
    日本植物学会第80回大会 2016年09月 ポスター発表
  • UDP-GlcNAc再生に関わるシロイヌナズナN-アセチルグルコサミンキナーゼの生理機能の解析  [通常講演]
    佐藤 康, 田中昌樹, 財津良多
    日本植物学会第80回大会 2016年09月 ポスター発表
  • シロイヌナズナの側根原基形成における非対称細胞分裂の終結制御とミトコンドリア機能および温度との関係  [通常講演]
    間宮章仁, 大塚蔵嵩, 山本荷葉子, 八木祐介, 中村崇裕, 野崎 守, 佐藤 康, 上田貴志, 蜂谷卓士, 野口 航, 平山 隆志, 杉山 宗隆
    第38回 日本分子生物学会年会 2015年12月 口頭発表(一般)
  • UDP-GlcNAc欠乏により誘導される根の成長阻害及びリグニン異常蓄積機構の解析  [通常講演]
    中妻直登, 佐藤 康
    日本植物学会第79会大会 2015年09月 ポスター発表
  • UDP-GlcNAcの再生経路に関わるシロイヌナズナN-アセチルグルコサミンキナーゼの機能解析  [通常講演]
    田中昌樹, 村重紘志, 佐藤 康
    日本植物学会第79回大会 2015年09月 ポスター発表
  • シロイヌナズナの側根帯化とミトコンドリアmRNAの編集・代謝との関係  [通常講演]
    間宮章仁, 大塚 蔵嵩, 山本 荷葉子, 野崎 守, 佐藤 康, 八木 祐介, 中村 崇裕, 平山 隆志, 杉山 宗隆
    日本植物学会第79回大会 2015年09月
  • GlcNAc及びGalNAcの再利用経路に関わるシロイヌナズナ突然変異体の単離と解析  [通常講演]
    小松淳平, 佐藤 康
    日本植物学会第79回大会 2015年09月
  • GlcNAc及びGalNAcの再利用経路に関わるシロイヌナズナ突然変異体の単離  [通常講演]
    小松淳平, 佐藤 康
    中国四国植物学会 第72回大会 2015年05月
  • シロイヌナズナの側根帯化変異体の解析から見えてきたミトコンドリアmRNAのポリA依存的代謝と編集の機能的連関  [通常講演]
    大塚蔵嵩, 山本荷葉子, 間宮章仁, 斉藤真人, 有田真規, 玉置裕章, 八木祐介, 中村崇裕, 野崎 守, 佐藤 康, 上田貴志, 平山隆志, 杉山宗隆
    第56回日本植物生理学会年会 2015年03月 口頭発表(一般)
  • シロイヌナズナTDF変異体の解析から見えてきたミトコンドリアmRNAのポリA依存的代謝と編集の密接な関係  [通常講演]
    大塚蔵嵩, 山本荷葉子, 間宮章仁, 斉藤真人, 有田真規, 玉置裕章, 八木祐介, 中村崇裕, 野崎 守, 佐藤康, 上田貴志, 平山隆志, 杉山宗隆
    第4回 植物RNA研究者ネットワークシンポジウム 2015年01月 口頭発表(一般)
  • UDP-GlcNAc再生に関わるシロイヌナズナN-アセチルグルコサミンキナーゼの特性および機能解析  [通常講演]
    村重紘志, 風呂圭祐, 野崎 守, 佐藤 康
    日本植物学会第78会大会 2014年09月 口頭発表(一般)
  • カヤツリグサ科マツバイの高増殖無菌培養条件の検討及びCs吸収・蓄積機構の解析  [通常講演]
    後藤慎平, 寺岡翔也, 竹原明成, 榊原正幸, 佐野 栄, 佐藤 康
    日本植物学会第78会大会 2014年09月 ポスター発表
  • シロイヌナズナにおけるストレス応答リグニン蓄積と小胞体ストレスとの関係の解析  [通常講演]
    中妻直登, 野崎 守, 佐藤 康
    第55回日本植物生理学会年会 2014年03月 ポスター発表
  • 植物におけるN-アセチルグルコサミンの再利用に関わるN-アセチルグルコサミンキナーゼの同定と解析  [通常講演]
    風呂圭祐, 野崎 守, 佐藤 康
    日本植物学会第77会大会 2013年09月 口頭発表(一般)
  • シロイヌナズナにおける2つのフェロキラターゼアイソフォームによるヘム分配機構の解析  [通常講演]
    増田 健, N. A. Espinas, 小林康一, 佐藤 康, 高橋香織, 田中亮一, 望月伸悦
    日本植物学会第77会大会 2013年09月 ポスター発表
  • カヤツリグサ科ハリイ属チャボイによる重金属汚染のファイトレメディエーションに関する基礎的研究  [通常講演]
    榊原正幸, 向井 菫, 佐藤 康, 佐野 栄
    第19回地下水・土壌汚染とその防止対策に関する研究集会 2013年06月 口頭発表(一般)
  • 側根原基形成時の細胞分裂制御におけるミトコンドリアmRNA代謝の役割  [通常講演]
    大塚蔵嵩, 野崎 守, 佐藤 康, 蜂谷卓士, 野口 航, 上田貴志, 平山隆志, 杉山宗隆
    第54回日本植物生理学会年会 2013年03月 口頭発表(一般)
  • Allocation of heme is differentially regulated by ferrochelatasse isoforms in Arabidopsis cells  [通常講演]
    N. Espinas, K. Kobayashi, Y. Sato, N. Mochizuki, T. Masuda
    第54回日本植物生理学会年会 2013年03月 ポスター発表
  • シロイヌナズナ温度感受性突然変異体lignescensにおける異常なリグニン蓄積について  [通常講演]
    野崎 守, 杉山 宗隆, 佐藤 康
    日本植物学会第76回大会 2012年09月 ポスター発表
  • タバコBY-2細胞の静置培養により誘導される細胞死におよぼす細胞周期の影響  [通常講演]
    平賀 旭, 金田 剛史, 佐藤 康, 佐藤 成一
    日本植物学会第76回大会 2012年09月 ポスター発表
  • キトサンシグナル伝達系解明のためのシロイヌナズナ根成長阻害誘導実験系の確立  [通常講演]
    木村 友哉, 大谷 周平, 佐藤 康
    日本植物学会第76回大会 2012年09月 ポスター発表
  • What factors affect the induction of cell death in exponential- and stationary- phase BY-2 cells under still conditions?  [通常講演]
    A. Hiraga, T. Kaneta, Y. Sato, S. Sato
    XVIII International botanical congress, Melbourne 2011年07月 ポスター発表
  • Analysis of the regulatory mechanisms of wound induced lignification in Arabidopsis thaliana  [通常講演]
    Y. Sato, F. Katsuno, T. Ohmori
    XVIII International botanical congress, Melbourne 2011年07月 ポスター発表
  • Analysis of lignescens, a novel temperature-sensitive mutant of Arabidopsis exhibiting ectopic lignin deposition  [通常講演]
    M. Nozaki, M. Sugiyama, Y. Sato
    XVIII International botanical congress, Melbourne 2011年07月 ポスター発表
  • タバコBY-2 細胞の細胞周期と静置培養により誘導される細胞死との関係  [通常講演]
    平賀 旭, 金田 剛史, 佐藤 康, 佐藤 成一
    日本植物学会第74回大会 2010年09月 ポスター発表
  • アズキ側根におけるオーキシン排出担体 VaPIN の発現の組織特異性の解析  [通常講演]
    久家 徳之, 金田 剛史, 佐藤 康, 佐藤 成一
    日本植物学会第74回大会 2010年09月 ポスター発表
  • 様々なシソにおける紫外線耐性機構の解析  [通常講演]
    王 麗林, 齋藤 和樹, 佐藤 康
    日本植物学会第74回大会 2010年09月 ポスター発表
  • 制限温度下で異所性リグニン沈着を示すシロイヌナズナ温度感受性突然変異体 lignescens の原因遺伝子  [通常講演]
    野崎 守, 杉山 宗隆, 佐藤 康
    日本植物学会第74回大会 2010年09月 ポスター発表
  • タバコBY-2細胞の静置により誘導される視細胞の割合に及ぼす要因について  [通常講演]
    平賀 旭, 金田 剛史, 佐藤 康, 佐藤 成一
    日本植物学会中国四国支部第67回大会 2010年05月

MISC

共同研究・競争的資金等の研究課題

  • 植物におけるUDP-N-アセチルグルコサミン生合成の生理的意義
    日本学術振興会:科研費:基盤研究(C)
    研究期間 : 2013年04月 -2016年03月 
    代表者 : 佐藤 康
  • ヒャクニチソウ管状要素分化実験系を用いたリグニン合成ペルオキシダーゼ遺伝子の解析
    日本学術振興会:科研費:奨励研究(A)
    研究期間 : 1997年04月 -1999年03月 
    代表者 : 佐藤 康

社会貢献活動

  • 親子で楽しむ科学実験
    期間 : 2019年08月24日 - 2019年08月25日
    役割 : 企画
    主催者・発行元 : 愛媛大学理学部
  • 親子で楽しむ科学実験
    期間 : 2016年08月20日 - 2016年08月21日
    役割 : 企画
    主催者・発行元 : 愛媛大学理学部
  • 親子で楽しむ科学実験
    期間 : 2015年08月22日 - 2015年08月23日
    役割 : 企画
    主催者・発行元 : 愛媛大学理学部

愛媛大学教員活動実績

教育活動(B)

担当授業科目(B01)

  • 2019, 前期, 学部, 生物学ゼミナールⅠ
  • 2019, 前期, 学部, 生物学特別演習Ⅰ
  • 2019, 前期, 学部, 卒業研究Ⅰ
  • 2019, 前期, 学部, 基礎生物学実験
  • 2019, 前期, 学部, 生物学実験III
  • 2019, 前期, 学部, 基礎生物学実験
  • 2019, 前期, 学部, 植物分子生理学
  • 2019, 前期, 修士, 細胞機能構造学
  • 2019, 前期, 修士, 生物学ゼミナールI
  • 2019, 前期, 修士, 生物学ゼミナールⅢ
  • 2019, 前期, 修士, 生物学課題実験I
  • 2019, 前期, 修士, 生物学課題実験Ⅲ
  • 2019, 前期, 学部, 基礎生物学実験


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