JP2010057395A - Lactic acid bacterium having enteric canal immunomodulating action - Google Patents
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本発明は、腸管免疫を賦活させる作用を有するラクトコッカス・ラクティス(Lactococcus lactis)に属する乳酸菌の新規株に関する。 The present invention relates to a novel strain of lactic acid bacteria belonging to Lactococcus lactis having an action of stimulating intestinal immunity.
ヒトの腸管は、摂取した食物を消化・吸収する器官であると同時に、有害な細菌や病原菌、ウイルスなどが経口的に体内へ侵入するのを防ぐ免疫組織である。腸管で誘導される主要な免疫応答の1つが、免疫グロブリンの一種であるイムノグロブリンA(IgA)抗体の分泌である。IgA抗体は、病原菌の腸管粘膜からの侵入阻止、毒素の中和、アレルゲンの侵入阻止などに働き、その分泌増加は、腸管免疫の賦活化を意味する。腸管におけるIgA抗体産生で重要な役割を果たす組織の1つがパイエル板である。また、IgA抗体の分泌を促進する細胞間伝達物質(サイトカイン)として、インターロイキン(IL)−5、IL−6、IL−10が知られている。 The human intestinal tract is an organ that digests and absorbs ingested food, and at the same time is an immune system that prevents harmful bacteria, pathogens, viruses, etc. from entering the body orally. One of the major immune responses induced in the intestine is the secretion of immunoglobulin A (IgA) antibody, a type of immunoglobulin. IgA antibodies act to prevent invasion of pathogenic bacteria from the intestinal mucosa, neutralize toxins, prevent invasion of allergens, etc. Increased secretion means activation of intestinal immunity. One tissue that plays an important role in IgA antibody production in the intestine is the Peyer's patch. Further, interleukin (IL) -5, IL-6, and IL-10 are known as intercellular transmitters (cytokines) that promote secretion of IgA antibodies.
特に乳酸菌は、プロバイオティクス即ち「腸管フローラを改善することによって、宿主に有益な作用をもたらす生きた微生物」として、その整腸作用や免疫賦活作用が注目されている。例えば、パイエル板細胞におけるIgA産生を強力に誘導できるビフィズス菌の菌株及びそれを含有する免疫賦活用組成物が知られている(特許文献1を参照)。また、しば漬けに生息する植物性乳酸菌の中から、免疫調節作用を有するラクトバチルス・ペントーサスに属する新規株が分離されている(特許文献2を参照)。更に、すんき漬などから分離した、植物由来のラクトバチルス属に属する乳酸菌、植物由来のロイコノストック属に属する乳酸菌、及び植物由来のペディオコッカス属に属する乳酸菌のうち少なくとも1以上の乳酸菌の生菌、死菌又はその菌体処理物を有効成分とする腸管免疫活性化剤が知られている(特許文献3を参照)。これら乳酸菌の生菌、死菌又はその菌体処理物は、動物由来の動物性乳酸菌に比べて十分な腸管免疫活性化作用やパイエル板細胞のIgA抗体産生促進作用を有することが認められた。 In particular, lactic acid bacteria are attracting attention as probiotics, ie, “living microorganisms that bring about beneficial effects on the host by improving the intestinal flora”, and their intestinal action and immunostimulatory action. For example, a strain of bifidobacteria that can strongly induce IgA production in Peyer's patch cells and an immunostimulatory composition containing the same are known (see Patent Document 1). In addition, a novel strain belonging to Lactobacillus pentosas having an immunomodulating action has been isolated from plant lactic acid bacteria that inhabit shibazuke (see Patent Document 2). Furthermore, at least one or more of lactic acid bacteria belonging to the genus Lactobacillus derived from plants, lactic acid bacteria belonging to the genus Leuconostoc derived from plants, and lactic acid bacteria belonging to the genus Pediococcus derived from plants isolated from pickles An intestinal immunity activator is known which contains live bacteria, dead bacteria or treated cells thereof as an active ingredient (see Patent Document 3). It was confirmed that these live and killed bacteria of lactic acid bacteria or treated cells thereof had a sufficient intestinal immunity activation effect and an acceleration of IgA antibody production of Peyer's patch cells compared to animal-derived lactic acid bacteria.
清酒醸造に使用する生もと系の山廃酒母には、清酒酵母以外に、硝酸還元菌や乳酸菌、その他様々な微生物の変遷が見られる。山廃酒母に生息する乳酸菌は、ロイコノストック・メセンテロイデス(Leuconostoc
mesenteroides)とラクトバチルス・サケ(Lactobacillus sake)の2種類がよく知られている。しかしながら、山廃酒母の詳細な乳酸菌叢は、少なくとも本願発明者らが知る限り、今日まで十分に明らかにされていない。山廃酒母から分離した乳酸菌の機能、効能についても、十分な研究がなされていない。
In addition to sake yeast, there are changes in nitrate-reducing bacteria, lactic acid bacteria, and various other microorganisms in the raw mountain waste liquor used for sake brewing. Lactic acid bacteria inhabiting the mountain waste liquor are Leuconostoc mesenteroides (Leuconostoc)
Two types are well known: mesenteroides) and Lactobacillus sake. However, the detailed lactic acid bacteria flora of the Yamawaku Sake Mother has not been sufficiently clarified to date, at least as far as the present inventors know. There has not been enough research on the function and efficacy of lactic acid bacteria isolated from Yamawaku Sake.
そこで、本発明の目的は、山廃酒母から分離される乳酸菌の中から、腸管免疫の調節作用などの有用な生理活性を有する菌株を提供することにある。 Then, the objective of this invention is providing the strain which has useful physiological activities, such as a regulation effect | action of an intestinal tract immunity, from the lactic acid bacteria isolate | separated from a mountain waste liquor.
本願発明者らは、山廃酒母から132株の乳酸菌を分離し、それらの腸管免疫調節作用を研究した。その結果、これら乳酸菌の中から、ラクトコッカス・ラクティスに属する菌株などに高い腸管免疫賦活作用があることを見出し、本発明に至ったものである。 The inventors of the present application isolated 132 lactic acid bacteria from Yamabo Sake and studied their intestinal immunity-modulating action. As a result, among these lactic acid bacteria, a strain belonging to Lactococcus lactis has been found to have a high intestinal immunity stimulating action, and the present invention has been achieved.
本発明によれば、山廃酒母から分離した、腸管免疫調節作用を有するラクトコッカス・ラクティス(Lactococcus lactis)に属する乳酸菌が提供される。 According to the present invention, there is provided a lactic acid bacterium belonging to Lactococcus lactis having an intestinal immunity-modulating action, which is isolated from a mountain waste liquor.
本発明のラクトコッカス・ラクティスに属する乳酸菌は、腸管のパイエル板細胞からのIgA抗体分泌を促進させる作用を有する。従って、本乳酸菌を生菌、死菌又は菌体処理物などのいかなる形態でも経口的に摂取すれば、腸管免疫を活性化させることができ、それにより、生体に悪影響を及ぼし得る病原菌などの体内への侵入を抑制し、様々な疾病予防にも寄与するという効果が得られる。 The lactic acid bacteria belonging to Lactococcus lactis of the present invention have an action of promoting IgA antibody secretion from Peyer's patch cells of the intestinal tract. Therefore, ingestion of intestinal tract immunity can be activated by ingesting the lactic acid bacterium orally in any form such as live bacteria, dead bacteria, or treated bacterial cells, thereby causing the body of pathogenic bacteria or the like that can adversely affect the living body. The effect of suppressing the invasion into the body and contributing to the prevention of various diseases can be obtained.
本発明の乳酸菌は、マンニトールを唯一の炭素源とする培地を用いることによって、山廃酒母から効果的に単離することができる。 The lactic acid bacteria of the present invention can be effectively isolated from a mountain waste liquor by using a medium containing mannitol as the sole carbon source.
より具体的に、本発明の乳酸菌は、独立行政法人産業技術総合研究所・特許生物寄託センターに受託番号NITE P−603で寄託された、ラクトコッカス・ラクティスKLB−M8株である。このラクトコッカス・ラクティスKLB−M8株は、高い腸管免疫賦活活性を有するので好ましい。 More specifically, the lactic acid bacterium of the present invention is Lactococcus lactis KLB-M8 strain deposited at the National Institute of Advanced Industrial Science and Technology, Patent Biological Deposit Center under the deposit number NITE P-603. This Lactococcus lactis KLB-M8 strain is preferable because it has high intestinal immunity activation activity.
以下に、添付図面を参照しつつ、本発明の好適な実施例を詳細に説明する。 Hereinafter, preferred embodiments of the present invention will be described in detail with reference to the accompanying drawings.
本発明の乳酸菌は、清酒醸造に用いる山廃酒母に生息する乳酸菌から所望の菌株を分離することによって得られる。清酒醸造における仕込み8日目頃又は14日目頃の山廃酒母は、特に乳酸菌が増殖しているので好ましい。乳酸菌株の分離は、公知の方法を用いて行うことができる。例えば、山廃酒母を乳酸菌分離用寒天培地に添加して培養し、コロニーを形成したものを選択する。 The lactic acid bacteria of the present invention can be obtained by isolating a desired strain from lactic acid bacteria that inhabit mountain waste liquor used for sake brewing. The mountain waste liquor mothers around the 8th day or the 14th day in the sake brewing are particularly preferred because lactic acid bacteria are growing. Separation of the lactic acid strain can be performed using a known method. For example, a mountain waste liquor is added to an agar medium for separating lactic acid bacteria and cultured to select a colony.
乳酸菌単離用の培地には、乳酸菌検出用の中川培地、マンニトール又はソルビトールを唯一の炭素源とする乳酸菌分離用培地(MRS培地)を使用することができる。特に高い腸管免疫賦活活性を有するラクトコッカス・ラクティスに属する乳酸菌は、マンニトールを唯一の炭素源とするMRS培地を用いることによって、より選択的に分離される。 As the medium for isolating lactic acid bacteria, Nakagawa medium for detecting lactic acid bacteria, or a medium for separating lactic acid bacteria (MRS medium) using mannitol or sorbitol as the sole carbon source can be used. Lactic acid bacteria belonging to Lactococcus lactis having particularly high intestinal immunity activation activity are more selectively separated by using an MRS medium containing mannitol as the sole carbon source.
分離した乳酸菌の種類は、16SrRNAの塩基配列を解析することによって同定される。中川培地で単離したものには、ロイコノストック・シトレウム(Leuconostoc
citreum)、ラクトバチルス・サケが含まれていた。マンニトールを唯一の炭素源とするMRS培地から単離したものには、ラクトコッカス・ラクティスが含まれていた。ソルビトールを唯一の炭素源とするMRS培地には、ラクトバチルス・プランタラム(Lactobacillus
plantarum)が含まれていた。これら乳酸菌の中でラクトバチルス・サケ以外のものは、これまで山廃酒母からの検出が報告されていなかったものである。
The kind of the separated lactic acid bacteria is identified by analyzing the base sequence of 16S rRNA. For those isolated on Nakagawa medium, Leuconostoc
citreum) and Lactobacillus salmon. Those isolated from MRS medium with mannitol as the sole carbon source contained Lactococcus lactis. The MRS medium with sorbitol as the only carbon source includes Lactobacillus (Lactobacillus).
plantarum). Among these lactic acid bacteria, those other than Lactobacillus salmon have not been reported to be detected from mountain waste liquors.
本発明の乳酸菌は、腸管のパイエル板細胞からのIgA抗体分泌を促進させることにより、腸管免疫を調節する作用を有する。その中で特に高い腸管免疫賦活活性を有するものは、ラクトコッカス・ラクティスKLB−M8株であり、上述したようにマンニトールを唯一の炭素源とするMRS培地を用いて山廃酒母から分離される。この菌株は、独立行政法人産業技術総合研究所・特許生物寄託センターに受託番号NITE P−603で寄託されている。 The lactic acid bacterium of the present invention has an action of regulating intestinal immunity by promoting the secretion of IgA antibody from Peyer's patch cells of the intestinal tract. Among them, the Lactococcus lactis KLB-M8 strain has a particularly high intestinal immunity activation activity, and is isolated from a mountain waste liquor using an MRS medium containing mannitol as a sole carbon source as described above. This strain is deposited under the accession number NITE P-603 at the National Institute of Advanced Industrial Science and Technology (AIST).
これら本発明の乳酸菌は、生菌、死菌、又は有機溶媒などで抽出したような菌体処理物のいずれかの形態であっても良い。本発明の乳酸菌をいかなる形態でも経口的に摂取することによって、腸管免疫を活性化させることができる。これにより、生体に悪影響を及ぼし得る病原菌やウィルスなどが体内へ侵入するのを抑制し、様々な疾病予防にも寄与するという効果が得られる。 These lactic acid bacteria of the present invention may be in the form of any one of viable bacteria, killed bacteria, or a treated cell product extracted with an organic solvent. Intestinal immunity can be activated by orally ingesting the lactic acid bacteria of the present invention in any form. As a result, it is possible to suppress the invasion of pathogenic bacteria or viruses that may adversely affect the living body into the body, thereby contributing to the prevention of various diseases.
(実施例1)
清酒醸造用の山廃酒母から上述した方法を用いて分離した132種の乳酸菌について、マウスのパイエル板細胞からのIgA抗体の分泌を促進させる菌株を、以下のin
vitro試験により探索した。分離した前記132株の乳酸菌は、MRS培地で1日培養した後、集菌し、生理食塩水で洗浄し、ホルムアルデヒドを加えて一晩30℃でインキュベートし、更に生理食塩水で洗浄して、試験に供した。
Example 1
About 132 types of lactic acid bacteria isolated from the waste sake mother for sake brewing using the method described above, a strain that promotes the secretion of IgA antibody from mouse Peyer's patch cells is expressed in the following in
Searched by in vitro tests. The isolated 132 strains of lactic acid bacteria were cultured in MRS medium for 1 day, collected, washed with physiological saline, added formaldehyde, incubated overnight at 30 ° C., further washed with physiological saline, It used for the test.
6週齢のオスBALB/cマウスの解剖を行い、小腸のパイエル板を摘出した。パイエル板内の免疫担当細胞を抽出し、1×106個の細胞を96穴プレートに播きこんだ。そこに、上述したようにホルムアルデヒド処理した前記132株の乳酸菌の死菌体を1×108cells/mlとなるように添加し、10%FBS(牛胎児血清)を含むRPMI(Roswell
Park Memorial Institute)培地で培養した。
Six-week-old male BALB / c mice were dissected and the small intestine Peyer's patches were extracted. The immunocompetent cells in the Peyer's patches were extracted and 1 × 10 6 cells were seeded in a 96-well plate. The dead cells of the 132 strains of lactic acid bacteria treated with formaldehyde as described above were added to 1 × 10 8 cells / ml, and RPMI (Roswell containing 10% FBS (fetal calf serum)) was added.
Cultured in Park Memorial Institute) medium.
培養開始から72時間後、培養上清を回収した。前記培養上清中にパイエル板細胞から分泌されたIgA抗体、IL(インターロイキン)−5、IL−6をELISA法で測定した。対照として、前記乳酸菌を添加しないもの(control)と、LPS(リポポリサッカライド)を0.25mg/mlとなるように添加したもの(LPS)とについても、同様の測定を行った。前記132株の中から比較的高い免疫賦活活性を示す乳酸菌(M1〜M15)について、それらの結果を前記対照と共に図1〜図3に示す。 After 72 hours from the start of culture, the culture supernatant was collected. IgA antibodies, IL (interleukin) -5, and IL-6 secreted from Peyer's patch cells in the culture supernatant were measured by ELISA. As a control, the same measurement was performed on the control without addition of the lactic acid bacteria (control) and the addition of LPS (lipopolysaccharide) at 0.25 mg / ml (LPS). The results of lactic acid bacteria (M1 to M15) exhibiting relatively high immunostimulatory activity among the 132 strains are shown in FIGS. 1 to 3 together with the control.
図1に示すように、乳酸菌を添加しない対照(control)と比べてIgA抗体の濃度を有意に(約1.4倍)増加させた菌株は、M7、M8、M15であった。また、有意とまで言えないが、M1、M9、M10、M11、M12の菌株も、IgA抗体の分泌を増加させる傾向があった。 As shown in FIG. 1, the strains that significantly increased the IgA antibody concentration (about 1.4 times) compared to the control without addition of lactic acid bacteria were M7, M8, and M15. Although not significant, the M1, M9, M10, M11, and M12 strains also tended to increase the secretion of IgA antibodies.
次に、IgA抗体の分泌誘導機構を調べるために、IgA抗体の誘導物質であるIL−5とIL−6とを、上述したように各菌株M1〜M15及び両対照についてELISA法で測定した。図2は、IL−5分泌量の測定結果を示している。同図に示すように、IL−5は、LPSの添加によって対照(control)の12倍以上に増加した。ところが、M1〜M15の中に、その添加によって対照(control)から有意に増加した菌株は無かった。しかしながら、M7、M8、M9株はIL−5を増加させる傾向を示した。 Next, in order to examine the secretory induction mechanism of IgA antibody, IL-5 and IL-6, which are inducers of IgA antibody, were measured by ELISA for each strain M1 to M15 and both controls as described above. FIG. 2 shows the measurement results of IL-5 secretion. As shown in the figure, IL-5 was increased to 12 times or more of the control by addition of LPS. However, none of the strains M1 to M15 significantly increased from the control by the addition. However, M7, M8 and M9 strains showed a tendency to increase IL-5.
図3は、IL−6分泌量の測定結果を示している。同図に示すように、IL−6は、LPS及びすべての菌株M1〜M15の添加によって対照(control)から増加した。特にM8株は、その添加によるIL−6の分泌促進が顕著であった。M8株は、IgA抗体分泌量及びIL−5分泌量の増加においても高い値を示しており、特に腸管免疫の賦活に有用な菌株と判断される。 FIG. 3 shows the measurement results of IL-6 secretion. As shown in the figure, IL-6 was increased from the control by addition of LPS and all strains M1-M15. In particular, the M8 strain was markedly enhanced in the secretion of IL-6 by its addition. The M8 strain also shows high values in the increase in IgA antibody secretion amount and IL-5 secretion amount, and is judged to be a particularly useful strain for stimulating intestinal immunity.
本実施例の結果から、山廃酒母から分離した乳酸菌の中に、腸管免疫を司るパイエル板細胞からのIgA抗体の分泌を促進させることにより腸管免疫賦活作用を示す菌株が含まれることは明らかである。そのような菌株では、図1〜図3を対比すると、IgA抗体の分泌量とIL−5、IL−6の分泌量との間に高い相関性が認められる。これから、乳酸菌によるパイエル板細胞からのIL−5、IL−6の分泌増加がIgA抗体の分泌を促進するという、免疫賦活の作用機構も明らかとなった。 From the results of the present example, it is clear that the lactic acid bacteria isolated from the mountain waste liquor include strains that show intestinal immunity activation by promoting secretion of IgA antibodies from Peyer's patch cells that control intestinal immunity. is there. In such a strain, when FIGS. 1 to 3 are compared, a high correlation is observed between the secretion amount of IgA antibody and the secretion amounts of IL-5 and IL-6. From this, the mechanism of action of immunostimulation that the increase in secretion of IL-5 and IL-6 from Peyer's patch cells by lactic acid bacteria promotes the secretion of IgA antibody was also clarified.
(実施例2)
実施例1において特に高い免疫賦活活性を示したM8株を用いて、以下のin vivo試験を行った。このM8株が、上述したように独立行政法人産業技術総合研究所・特許生物寄託センターに寄託したラクトコッカス・ラクティスKLB−M8株(以下、KLB−M8株と称す)である。
(Example 2)
The following in vivo tests were conducted using the M8 strain that showed particularly high immunostimulatory activity in Example 1. This M8 strain is the Lactococcus lactis KLB-M8 strain (hereinafter referred to as the KLB-M8 strain) deposited at the National Institute of Advanced Industrial Science and Technology, as described above.
5週齢のオスBALB/cマウスを無作為に4匹ずつ対照群と乳酸菌摂取群との2群に分け、1週間予備飼育した。予備飼育期間中、前記両群のマウスには、標準的なラボダイエットと飲用水とを自由に摂取させた。その後、対照群には、2%スキムミルクを含む飲用水を自由に摂取させた。乳酸菌摂取群には、2%スキムミルクを含む飲用水にKLB−M8株の加熱死菌体を1×108cells/mlとなるように加えたものを自由に摂取させた。尚、ラボダイエットは、予備飼育後の試験期間中も引き続き、乳酸菌摂取群と対照群共に同様に自由に摂取させた。 Four 5-week-old male BALB / c mice were randomly divided into two groups, a control group and a lactic acid bacterium-intake group, and pre-bred for one week. During the pre-breeding period, the mice in both groups were given free access to a standard lab diet and drinking water. Thereafter, the control group was allowed to freely drink drinking water containing 2% skim milk. The lactic acid bacteria ingestion group was allowed to freely ingest drinking water containing 2% skim milk with KLB-M8 strain heat-killed cells added to 1 × 10 8 cells / ml. The lab diet was continuously ingested in the same manner for both the lactic acid bacteria ingestion group and the control group during the test period after the preliminary breeding.
乳酸菌を与え始めてから7日後に、前記両群のマウスを解剖した。各個体毎に小腸のパイエル板を摘出し、パイエル板内の免疫担当細胞を抽出した。この細胞を、それぞれLPSを0,1,5,20μg/ml添加したRPMI−1640培地(10%FBS含有)で72時間培養し、培養上清を回収した。ELISA法を用いて、培養上清中にパイエル板細胞から分泌されたIgA抗体量を測定した。その結果を図4に、LPS濃度毎に棒グラフで示す。 Seven days after the start of feeding lactic acid bacteria, the mice in both groups were dissected. For each individual, the Peyer's patch of the small intestine was excised and the immunocompetent cells in the Peyer's patch were extracted. The cells were cultured in RPMI-1640 medium (containing 10% FBS) supplemented with 0, 1, 5, 20 μg / ml of LPS for 72 hours, and the culture supernatant was collected. Using the ELISA method, the amount of IgA antibody secreted from Peyer's patch cells into the culture supernatant was measured. The results are shown in FIG. 4 as bar graphs for each LPS concentration.
同図に示すように、乳酸菌摂取群のIgA抗体量は、特にLPS濃度が0または1μg/mlの場合、対照群の1.3倍以上であった。本実施例では、全体として、乳酸菌摂取群のIgA抗体量と対照群のそれとの間に、有意と言える程度の差は認められなかった。しかしながら、いずれのLPS濃度でも、乳酸菌摂取群のIgA抗体量が対照群を上回った。 As shown in the figure, the amount of IgA antibody in the lactic acid bacteria intake group was 1.3 times or more that in the control group, particularly when the LPS concentration was 0 or 1 μg / ml. In this example, as a whole, no significant difference was observed between the IgA antibody amount in the lactic acid bacteria intake group and that in the control group. However, at any LPS concentration, the amount of IgA antibody in the lactic acid bacteria intake group exceeded the control group.
本実施例の結果から、KLB−M8株の摂取によって、小腸のパイエル板免疫担当細胞が刺激され、IgA抗体を産生しやすい状態へ穏やかに変化したと言うことができる。このようにIgA抗体産生能力が増加したということは、腸管免疫の賦活化を表している。従って、山廃酒母由来の乳酸菌ラクトコッカス・ラクティスKLB−M8株は、in
vivo刺激に対しても、その腸管免疫賦活作用が証明された。
From the results of this Example, it can be said that the ingestion of the KLB-M8 strain stimulated Peyer's patch immunocompetent cells in the small intestine and gently changed it to a state in which IgA antibodies can be easily produced. The increased ability to produce IgA antibody in this way represents activation of intestinal immunity. Therefore, the lactic acid bacterium Lactococcus lactis KLB-M8 strain derived from a mountain waste liquor
The intestinal immunity stimulating action was proved also against in vivo stimulation.
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Cited By (4)
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WO2012029367A1 (en) | 2010-08-31 | 2012-03-08 | 国立大学法人新潟大学 | Intestine immunomodulator |
WO2014129599A1 (en) * | 2013-02-22 | 2014-08-28 | 日東薬品工業株式会社 | Immunostimulation agent |
JP5879254B2 (en) * | 2010-03-04 | 2016-03-08 | 株式会社ロッテ | Immunoglobulin A secretion promoter |
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JP5879254B2 (en) * | 2010-03-04 | 2016-03-08 | 株式会社ロッテ | Immunoglobulin A secretion promoter |
WO2012029367A1 (en) | 2010-08-31 | 2012-03-08 | 国立大学法人新潟大学 | Intestine immunomodulator |
US8980289B2 (en) | 2010-08-31 | 2015-03-17 | Niigata University | Intestine immunomodulator |
US10570366B2 (en) | 2011-12-28 | 2020-02-25 | Yamada Bee Company Inc. | Lactic acid bacterium having IgA production promoting activity, and use thereof |
WO2014129599A1 (en) * | 2013-02-22 | 2014-08-28 | 日東薬品工業株式会社 | Immunostimulation agent |
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JPWO2014129599A1 (en) * | 2013-02-22 | 2017-02-02 | 日東薬品工業株式会社 | Immunostimulator |
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