研究者詳細 - 三谷　奈見季

論文 - 三谷　奈見季

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Silicon efflux transporters isolated from two pumpkin cultivars contrasting in Si uptake 査読

Namiki Mitani-Ueno, Naoki Yamaji, Jian Feng Ma

Plant Signaling and Behavior 6 ( 7 ) 73 - 76 2011年

記述言語：英語掲載種別：研究論文（学術雑誌）

The accumulation of silicon (Si) differs greatly with plant species and cultivars due to different ability of the roots to take up Si. In Si accumulating plants such as rice, barley and maize, Si uptake is mediated by the influx (Lsi1) and efflux (Lsi2) transporters. Here we report isolation and functional analysis of two Si efflux transporters (CmLsi2-1 and CmLsi2-2) from two pumpkin (Cucurbita moschata Duch.) cultivars contrasting in Si uptake. These cultivars are used for rootstocks of bloom and bloomless cucumber, respectively. Different from mutations in the Si influx transporter CmLsi1, there was no difference in the sequence of either CmLsi2 between two cultivars. Both CmLsi2-1 and CmLsi2-2 showed an efflux transport activity for Si and they were expressed in both the roots and shoots. These results confirm our previous finding that mutation in CmLsi1, but not in CmLsi2-1 and CmLsi2-2 are responsible for bloomless phenotype resulting from low Si uptake. © 2011 Landes Bioscience.

DOI： 10.4161/psb.6.7.15462

Transport of silicon from roots to panicles in plants 査読

Jian Feng Ma, Naoki Yamaji, Namiki Mitani-Ueno

Proceedings of the Japan Academy Series B: Physical and Biological Sciences 87 ( 7 ) 377 - 385 2011年

記述言語：英語

Silicon (Si) is the most abundant minerals in soil and exerts beneficial effects on plant growth by alleviating various stresses. The transport of Si from soil to the panicles is mediated by different transporters. Lsi1, belonging to a NIP group of the aquaporin family, is responsible for the uptake of Si from soil into the root cells in both dicots and monocots although its expression patterns and cellular localization differ with plant species. The subsequent transport of Si out of the root cells towards the stele is medicated by an active efflux transporter, Lsi2. Lsi1 and Lsi2 are polarly localized at the distal and proximal sides, respectively, of both exodermis and endodermis in rice root. Silicon in the xylem sap is presented in the form of monosilicic acid and is unloaded by Lsi6, a homolog of Lsi1 in rice. Lsi6 is also involved in the inter-vascular transfer of Si at the node, which is necessary for preferential Si distribution to the panicles. © 2011 The Japan Academy.

DOI： 10.2183/pjab.87.377

Involvement of Silicon Influx Transporter OsNIP2;1 in Selenite Uptake in Rice 査読

Xue Qiang Zhao, Namiki Mitani, Naoki Yamaji, Ren Fang Shen, Jian Feng Ma

PLANT PHYSIOLOGY 153 ( 4 ) 1871 - 1877 2010年8月

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

Rice (Oryza sativa) as a staple food, provides a major source of dietary selenium (Se) for humans, which essentially requires Se, however, the molecular mechanism for Se uptake is still poorly understood. Herein, we show evidence that the uptake of selenite, a main bioavailable form of Se in paddy soils, is mediated by a silicon (Si) influx transporter Lsi1 (OsNIP2;1) in rice. Defect of OsNIP2;1 resulted in a significant decrease in the Se concentration of the shoots and xylem sap when selenite was given. However, there was no difference in the Se concentration between the wild-type rice and mutant of OsNIP2;1 when selenate was supplied. A short-term uptake experiment showed that selenite uptake greatly increased with decreasing pH in the external solution. Si as silicic acid did not inhibit the Se uptake from selenite in both rice and yeast (Saccharomyces cerevisiae) at low pHs. Expression of OsNIP2;1 in yeast enhanced the selenite uptake at pH 3.5 and 5.5 but not at pH 7.5. On the other hand, defect of Si efflux transporter Lsi2 did not affect the uptake of Se either from selenite or selenate. Taken together, our results indicate that Si influx transporter OsNIP2;1 is permeable to selenite.

DOI： 10.1104/pp.110.157867

The role of the rice aquaporin Lsi1 in arsenite efflux from roots 査読

Fang-Jie Zhao, Yukiko Ago, Namiki Mitani, Ren-Ying Li, Yu-Hong Su, Naoki Yamaji, Steve P. McGrath, Jian Feng Ma

NEW PHYTOLOGIST 186 ( 2 ) 392 - 399 2010年

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-BLACKWELL PUBLISHING, INC

When supplied with arsenate (As(V)), plant roots extrude a substantial amount of arsenite (As(III)) to the external medium through as yet unidentified pathways. The rice (Oryza sativa) silicon transporter Lsi1 (OsNIP2;1, an aquaporin channel) is the major entry route of arsenite into rice roots. Whether Lsi1 also mediates arsenite efflux was investigated.
Expression of Lsi1 in Xenopus laevis oocytes enhanced arsenite efflux, indicating that Lsi1 facilitates arsenite transport bidirectionally.
Arsenite was the predominant arsenic species in arsenate-exposed rice plants. During 24-h exposure to 5 mu M arsenate, rice roots extruded arsenite to the external medium rapidly, accounting for 60-90% of the arsenate uptake. A rice mutant defective in Lsi1 (lsi1) extruded significantly less arsenite than the wild-type rice and, as a result, accumulated more arsenite in the roots. By contrast, Lsi2 mutation had little effect on arsenite efflux to the external medium.
We conclude that Lsi1 plays a role in arsenite efflux in rice roots exposed to arsenate. However, this pathway accounts for only 15-20% of the total efflux, suggesting the existence of other efflux transporters.

DOI： 10.1111/j.1469-8137.2010.03192.x

The Rice Aquaporin Lsi1 Mediates Uptake of Methylated Arsenic Species 査読

Ren-Ying Li, Yukiko Ago, Wen-Ju Liu, Namiki Mitani, Joerg Feldmann, Steve P. McGrath, Jian Feng Ma, Fang-Jie Zhao

PLANT PHYSIOLOGY 150 ( 4 ) 2071 - 2080 2009年8月

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

Pentavalent methylated arsenic (As) species such as monomethylarsonic acid [MMA(V)] and dimethylarsinic acid [DMA(V)] are used as herbicides or pesticides, and can also be synthesized by soil microorganisms or algae through As methylation. The mechanism of MMA(V) and DMA(V) uptake remains unknown. Recent studies have shown that arsenite is taken up by rice (Oryza sativa) roots through two silicon transporters, Lsi1 (the aquaporin NIP2;1) and Lsi2 (an efflux carrier). Here we investigated whether these two transporters also mediate the uptake of MMA(V) and DMA(V). MMA(V) was partly reduced to trivalent MMA(III) in rice roots, but only MMA(V) was translocated to shoots. DMA(V) was stable in plants. The rice lsi1 mutant lost about 80% and 50% of the uptake capacity for MMA(V) and DMA(V), respectively, compared with the wild-type rice, whereas Lsi2 mutation had little effect. The short-term uptake kinetics of MMA(V) can be described by a Michaelis-Menten plus linear model, with the wild type having 3.5-fold higher V(max) than the lsi1 mutant. The uptake kinetics of DMA(V) were linear with the slope being 2.8-fold higher in the wild type than the lsi1 mutant. Heterologous expression of Lsi1 in Xenopus laevis oocytes significantly increased the uptake of MMA(V) but not DMA(V), possibly because of a very limited uptake of the latter. Uptake of MMA(V) and DMA(V) by wild-type rice was increased as the pH of the medium decreased, consistent with an increasing proportion of the undissociated species. The results demonstrate that Lsi1 mediates the uptake of undissociated methylated As in rice roots.

DOI： 10.1104/pp.109.140350

Identification and Characterization of Maize and Barley Lsi2-Like Silicon Efflux Transporters Reveals a Distinct Silicon Uptake System from That in Rice 査読

Namiki Mitani, Yukako Chiba, Naoki Yamaji, Jian Feng Ma

PLANT CELL 21 ( 7 ) 2133 - 2142 2009年7月

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

Silicon (Si) uptake has been extensively examined in rice (Oryza sativa), but it is poorly understood in other gramineous crops. We identified Low Silicon Rice 2 (Lsi2)-like Si efflux transporters from two important gramineous crops: maize (Zea mays) and barley (Hordeum vulgare). Both maize and barley Lsi2 expressed in Xenopus laevis oocytes showed Si efflux transport activity. Furthermore, barley Lsi2 was able to recover Si uptake in a rice mutant defective in Si efflux. Maize and barley Lsi2 were only expressed in the roots. Expression of maize and barley Lsi2 was downregulated in response to exogenously applied Si. Moreover, there was a significant positive correlation between the ability of roots to absorb Si and the expression levels of Lsi2 in eight barley cultivars, suggesting that Lsi2 is a key Si transporter in barley. Immunostaining showed that maize and barley Lsi2 localized only at the endodermis, with no polarity. Protein gel blot analysis indicated that maize and barley Lsi2 localized on the plasma membrane. The unique features of maize and barley Si influx and efflux transporters, including their cell-type specificity and the lack of polarity of their localization in Lsi2, indicate that these crops have a different Si uptake system from that in rice.

DOI： 10.1105/tpc.109.067884

HvLsi1 is a silicon influx transporter in barley. 査読

Chiba Y, Mitani N, Yamaji N, Ma JF

The Plant journal : for cell and molecular biology 57 ( 5 ) 810 - 818 2009年3月

A Bacterial-Type ABC Transporter Is Involved in Aluminum Tolerance in Rice 査読

Chao Feng Huang, Naoki Yamaji, Namiki Mitani, Masahiro Yano, Yoshiaki Nagamura, Jian Feng Ma

PLANT CELL 21 ( 2 ) 655 - 667 2009年2月

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

Aluminum (Al) toxicity is a major factor limiting crop production in acidic soil, but the molecular mechanisms of Al tolerance are poorly understood. Here, we report that two genes, STAR1 (for sensitive to Al rhizotoxicity1) and STAR2, are responsible for Al tolerance in rice. STAR1 encodes a nucleotide binding domain, while STAR2 encodes a transmembrane domain, of a bacterial-type ATP binding cassette (ABC) transporter. Disruption of either gene resulted in hypersensitivity to aluminum toxicity. Both STAR1 and STAR2 are expressed mainly in the roots and are specifically induced by Al exposure. Expression in onion epidermal cells, rice protoplasts, and yeast showed that STAR1 interacts with STAR2 to form a complex that localizes to the vesicle membranes of all root cells, except for those in the epidermal layer of the mature zone. When expressed together in Xenopus laevis oocytes, STAR1/2 shows efflux transport activity specific for UDP-glucose. Furthermore, addition of exogenous UDP-glucose rescued root growth in the star1 mutant exposed to Al. These results indicate that STAR1 and STAR2 form a complex that functions as an ABC transporter, which is required for detoxification of Al in rice. The ABC transporter transports UDP-glucose, which may be used to modify the cell wall.

DOI： 10.1105/tpc.108.064543

OsFRDL1 Is a Citrate Transporter Required for Efficient Translocation of Iron in Rice 査読

Kengo Yokosho, Naoki Yamaji, Daisei Ueno, Namiki Mitani, Jian Feng Ma

PLANT PHYSIOLOGY 149 ( 1 ) 297 - 305 2009年1月

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

Multidrug and toxic compound extrusion (MATE) transporters represent a large family in plants, but their functions are poorly understood. Here, we report the function of a rice (Oryza sativa) MATE gene (Os03g0216700, OsFRDL1), the closest homolog of barley (Hordeum vulgare) HvAACT1 (aluminum [Al]-activated citrate transporter 1), in terms of metal stress (iron [Fe] deficiency and Al toxicity). This gene was mainly expressed in the roots and the expression level was not affected by either Fe deficiency or Al toxicity. Knockout of this gene resulted in leaf chlorosis, lower leaf Fe concentration, higher accumulation of zinc and manganese concentration in the leaves, and precipitation of Fe in the root's stele. The concentration of citrate and ferric iron in the xylem sap was lower in the knockout line compared to the wild-type rice. Heterologous expression of OsFRDL1 in Xenopus oocytes showed transport activity for citrate. Immunostaining showed that OsFRDL1 was localized at the pericycle cells of the roots. On the other hand, there was no difference in the Al-induced secretion of citrate from the roots between the knockout line and the wild-type rice. Taken together, our results indicate that OsFRDL1 is a citrate transporter localized at the pericycle cells, which is necessary for efficient translocation of Fe to the shoot as a Fe-citrate complex.

DOI： 10.1104/pp.108.128132

Identification of Maize Silicon Influx Transporters 査読

Namiki Mitani, Naoki Yamaji, Jian Feng Ma

PLANT AND CELL PHYSIOLOGY 50 ( 1 ) 5 - 12 2009年1月

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

Maize (Zea mays L.) shows a high accumulation of silicon (Si), but transporters involved in the uptake and distribution have not been identified. In the present study, we isolated two genes (ZmLsi1 and ZmLsi6), which are homologous to rice influx Si transporter OsLsi1. Heterologous expression in Xenopus laevis oocytes showed that both ZmLsi1 and ZmLsi6 are permeable to silicic acid. ZmLsi1 was mainly expressed in the roots. By contrast, ZmLsi6 was expressed more in the leaf sheaths and blades. Different from OsLsi1, the expression level of both ZmLsi1 and ZmLsi6 was unaffected by Si supply. Immunostaining showed that ZmLsi1 was localized on the plasma membrane of the distal side of root epidermal and hypodermal cells in the seminal and crown roots, and also in cortex cells in lateral roots. In the shoots, ZmLsi6 was found in the xylem parenchyma cells that are adjacent to the vessels in both leaf sheaths and leaf blades. ZmLsi6 in the leaf sheaths and blades also exhibited polar localization on the side facing towards the vessel. Taken together, it can be concluded that ZmLsi1 is an influx transporter of Si, which is responsible for the transport of Si from the external solution to the root cells and that ZmLsi6 mainly functions as a Si transporter for xylem unloading.

DOI： 10.1093/pcp/pcn110

NIP1;1, an Aquaporin Homolog, Determines the Arsenite Sensitivity of Arabidopsis thaliana 査読

Takehiro Kamiya, Mayuki Tanaka, Namiki Mitani, Jian Feng Ma, Masayoshi Maeshima, Toru Fujiwara

JOURNAL OF BIOLOGICAL CHEMISTRY 284 ( 4 ) 2114 - 2120 2009年1月

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 BIOCHEMISTRY MOLECULAR BIOLOGY INC

Arsenite [As(III)] is highly toxic to organisms, including plants. Very recently, transporters in rice responsible for As(III) transport have been described (Ma, J. F., Yamaji, N., Mitani, N., Xu, X. Y., Su, Y. H., McGrath, S. P., and Zhao, F. J. (2008) Proc. Natl. Acad. Sci. U. S. A. 105, 9931-9935), but little is known about As(III) tolerance. In this study, three independent As(III)-tolerant mutants were isolated from ethyl methanesulfonate-mutagenized M2 seeds of Arabidopsis thaliana. All three mutants carried independent mutations in Nodulin 26-like intrinsic protein 1;1 (NIP1;1), a homolog of an aquaporin. Two independent transgenic lines carrying T-DNA in NIP1; 1 were highly tolerant to As(III), establishing that NIP1;1 is the causal gene of As(III) tolerance. Because an aquaglyceroporin is able to transport As(III), we measured As(III) transport activity. When expressed in Xenopus oocytes, NIP1; 1 was capable of transporting As(III). As content in the mutant plants was 30% lower than in wild-type plants. Promoter beta-glucuronidase and real-time PCR analysis showed that NIP1; 1 is highly expressed in roots, and GFP-NIP1;1 is localized to the plasma membrane. These data show that NIP1; 1 is involved in As(III) uptake into roots and that disruption of NIP1; 1 function confers As(III) tolerance to plants. NIP1;2 and NIP5;1, closely related homologs of NIP1;1, were also permeable to As(III). Although the disruption of these genes reduced the As content in plants, As(III) tolerance was not observed in nip1;2 and nip5;1 mutants. This indicates that As(III) tolerance cannot be simply explained by decreased As contents in plants.

DOI： 10.1074/jbc.M806881200

Transporters of arsenite in rice and their role in arsenic accumulation in rice grain 査読

Jian Feng Ma, Naoki Yamaji, Namiki Mitani, Xiao-Yan Xu, Yu-Hong Su, Steve P. McGrath, Fang-Jie Zhao

PROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICA 105 ( 29 ) 9931 - 9935 2008年7月

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

Arsenic poisoning affects millions of people worldwide. Human arsenic intake from rice consumption can be substantial because rice is particularly efficient in assimilating arsenic from paddy soils, although the mechanism has not been elucidated. Here we report that two different types of transporters mediate transport of arsenite, the predominant form of arsenic in paddy soil, from the external medium to the xylem. Transporters belonging to the NIP subfamily of aquaporins in rice are permeable to arsenite but not to arsenate. Mutation in OsNIP2;1 (Lsi1, a silicon influx transporter) significantly decreases arsenite uptake. Furthermore, in the rice mutants defective in the silicon efflux transporter Lsi2, arsenite transport to the xylem and accumulation in shoots and grain decreased greatly. Mutation in Lsi2 had a much greater impact on arsenic accumulation in shoots and grain in field-grown rice than Lsi1. Arsenite transport in rice roots therefore shares the same highly efficient pathway as silicon, which explains why rice is efficient in arsenic accumulation. Our results provide insight into the uptake mechanism of arsenite in rice and strategies for reducing arsenic accumulation in grain for enhanced food safety.

DOI： 10.1073/pnas.0802361105

Characterization of substrate specificity of a rice silicon transporter, Lsi1 査読

Namiki Mitani, Naoki Yamaji, Jian Feng Ma

Pflugers Archiv European Journal of Physiology 456 ( 4 ) 679 - 686 2008年7月

記述言語：英語

Lsi1 (OsNIP2
1) is the first silicon (silicic acid) transporter identified in plant, which belongs to the nodulin 26-like intrinsic membrane protein (NIP) subfamily. In this study, we characterized the function of this transporter by using the Xenopus laevis oocyte expression system. The transport activity of Lsi1 for silicic acid was significantly inhibited by HgCl2 but not by low temperature. Lsi1 also showed an efflux transport activity for silicic acid. The substrate specificity study showed that Lsi1 was able to transport urea and boric acid
however, the transport activity for silicic acid was not affected by the presence of equimolar urea and was decreased only slightly by boric acid. Furthermore, among the NIPs subgroup, OsNIP2
2 showed transport activity for silicic acid, whereas OsNIP1
1 and OsNIP3
1 did not. We propose that Lsi1 and its close homologues form a unique subgroup of NIP with a distinct ar/R selectivity filter, which is located in the narrowest region on the extra-membrane mouth and govern the substrate specificity of the pore. © 2007 Springer-Verlag.

DOI： 10.1007/s00424-007-0408-y

Transcriptome analysis of the silicon-Magnaporthe grisea interaction

A. M. Brunings, L. E. Datnoff, J. F. Ma, N. Mitani, Y. Nagamura, B. Rathinasabapathi

PHYTOPATHOLOGY 98 ( 6 ) S28 - S28 2008年6月

Genotypic difference in silicon uptake and expression of silicon transporter genes in rice 査読

Jian Feng Ma, Naoki Yamaji, Kazunori Tamai, Namiki Mitani

PLANT PHYSIOLOGY 145 ( 3 ) 919 - 924 2007年11月

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

Rice (Oryza sativa) is a highly silicon (Si)-accumulating species that shows genotypic differences in Si accumulation. We investigated the physiological and molecular mechanisms involved in the genotypic difference in Si uptake between the japonica var. Nipponbare and the indica var. Kasalath. Both the Si concentration in the shoot and the Si uptake per root dry weight were higher in Nipponbare than in Kasalath grown in either soil or nutrient solution. The Si uptake by a single root was also higher in Nipponbare than in Kasalath. A kinetics study showed that Nipponbare and Kasalath had a similar K-m value, whereas the V-max was higher in Nipponbare. The expression of two Si transporter genes (Low silicon rice 1 [Lsi1] and Lsi2) investigated using real-time reverse transcription polymerase chain reaction revealed higher expression of both genes in Nipponbare than in Kasalath. Immunostaining with Lsi1 and Lsi2 antibodies revealed a similar pattern of subcellular localization of these two Si transporters in both varieties; Lsi1 and Lsi2 were localized at the distal and proximal sides, respectively, of both exodermis and endodermis of the roots. These results revealed that the genotypic difference in the Si accumulation results from the difference in abundance of Si transporters in rice roots.

DOI： 10.1104/pp.107.107599

An aluminum-activated citrate transporter in barley. 査読

Jun Furukawa, Naoki Yamaji, Hua Wang, Namiki Mitani, Yoshiko Murata, Kazuhiro Sato, Maki Katsuhara, Kazuyoshi Takeda, Jian Feng Ma

Plant & cell physiology 48 ( 8 ) 1081 - 91 2007年8月

記述言語：英語

Soluble ionic aluminum (Al) inhibits root growth and reduces crop production on acid soils. Al-resistant cultivars of barley (Hordeum vulgare L.) detoxify Al by secreting citrate from the roots, but the responsible gene has not been identified yet. Here, we identified a gene (HvAACT1) responsible for the Al-activated citrate secretion by fine mapping combined with microarray analysis, using an Al-resistant cultivar, Murasakimochi, and an Al-sensitive cultivar, Morex. This gene belongs to the multidrug and toxic compound extrusion (MATE) family and was constitutively expressed mainly in the roots of the Al-resistant barley cultivar. Heterologous expression of HvAACT1 in Xenopus oocytes showed efflux activity for (14)C-labeled citrate, but not for malate. Two-electrode voltage clamp analysis also showed transport activity of citrate in the HvAACT1-expressing oocytes in the presence of Al. Overexpression of this gene in tobacco enhanced citrate secretion and Al resistance compared with the wild-type plants. Transiently expressed green fluorescent protein-tagged HvAACT1 was localized at the plasma membrane of the onion epidermal cells, and immunostaining showed that HvAACT1 was localized in the epidermal cells of the barley root tips. A good correlation was found between the expression of HvAACT1 and citrate secretion in 10 barley cultivars differing in Al resistance. Taken together, our results demonstrate that HvAACT1 is an Al-activated citrate transporter responsible for Al resistance in barley.

DOI： 10.1093/pcp/pcm091

An efflux transporter of silicon in rice 査読

Jian Feng Ma, Naoki Yamaji, Namiki Mitani, Kazunori Tamai, Saeko Konishi, Toru Fujiwara, Maki Katsuhara, Masahiro Yano

NATURE 448 ( 7150 ) 209 - U12 2007年7月

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

Silicon is an important nutrient for the optimal growth and sustainable production of rice(1-4). Rice accumulates up to 10% silicon in the shoot, and this high accumulation is required to protect the plant from multiple abiotic and biotic stresses(1-5). A gene, Lsi1, that encodes a silicon influx transporter has been identified in rice(6). Here we describe a previously uncharacterized gene, low silicon rice 2 (Lsi2), which has no similarity to Lsi1. This gene is constitutively expressed in the roots. The protein encoded by this gene is localized, like Lsi1, on the plasma membrane of cells in both the exodermis and the endodermis, but in contrast to Lsi1, which is localized on the distal side, Lsi2 is localized on the proximal side of the same cells. Expression of Lsi2 in Xenopus oocytes did not result in influx transport activity for silicon, but preloading of the oocytes with silicon resulted in a release of silicon, indicating that Lsi2 is a silicon efflux transporter. The identification of this silicon transporter revealed a unique mechanism of nutrient transport in plants: having an influx transporter on one side and an efflux transporter on the other side of the cell to permit the effective transcellular transport of the nutrients.

DOI： 10.1038/nature05964

Characterization of silicon permeability of NIP in gramineous plants 査読

Namiki Mitani, Naoki Yamaji, Maki Katsuhara, Jian Feng Ma

PLANT AND CELL PHYSIOLOGY 48 S30 - S30 2007年

Functional analysis of a rice Si transporter homolog Lsi6. 査読

Naoki Yamaji, Namiki Mitani, Jian Feng Ma

PLANT AND CELL PHYSIOLOGY 48 S30 - S30 2007年

A silicon transporter in rice. 査読

Ma JF, Tamai K, Yamaji N, Mitani N, Konishi S, Katsuhara M, Ishiguro M, Murata Y, Yano M

Nature 440 ( 7084 ) 688 - 691 2006年3月

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