β-葡聚糖对鱼类免疫调节作用的研究进展
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摘要
为了维持鱼类健康并为养殖生产带来更好的效益,通常在养殖鱼类饲料中使用免疫增强剂作为饲料添加剂以提高鱼类的生产性能和疾病抵抗力。对多种鱼类的研究发现,饲喂β-葡聚糖能提高其免疫应答、对细菌与病毒的感染和环境压力的抵御力。因此,β-葡聚糖作为免疫增强剂在水产养殖中的应用在集约化养殖的今天显得至关重要。本文就β-葡聚糖的化学结构、β-葡聚糖调节鱼类免疫的机制以及影响β-葡聚糖生物活性的因素等方面进行综述,旨在为后续水产养殖中β-葡聚糖作为饲料添加剂的合理应用提供参考。
In order to maintain the fish health and bring benefits to aquaculture production,immunostimulants have been used as feed additives to improve production and disease resistance of fish. In many species,application of yeast β-glucans through feeding has been found can improve immune response,resistance to bacterial and viral infections and to environmental stress. Thus,the application of β-glucans as immunomodulators appears great importance in intensive aquaculture system. This paper summarized the chemical structure of β-glucans,mechanism of β-glucans on immune regulation for fish,and the influence factors of biological activities of β-glucans,in order to provide profitable reference on the reasonable application of β-glucans as feed additives in subsequent aquaculture.[Chinese Journal of Animal Nutrition,2019,31(7):2971-2980]
In order to maintain the fish health and bring benefits to aquaculture production,immunostimulants have been used as feed additives to improve production and disease resistance of fish. In many species,application of yeast β-glucans through feeding has been found can improve immune response,resistance to bacterial and viral infections and to environmental stress. Thus,the application of β-glucans as immunomodulators appears great importance in intensive aquaculture system. This paper summarized the chemical structure of β-glucans,mechanism of β-glucans on immune regulation for fish,and the influence factors of biological activities of β-glucans,in order to provide profitable reference on the reasonable application of β-glucans as feed additives in subsequent aquaculture.[Chinese Journal of Animal Nutrition,2019,31(7):2971-2980]
引文
[1] YUN C H,ESTRADA A,VAN KESSEL A,et al.β-glucan,extracted from oat,enhances disease resistance against bacterial and parasitic infections[J]. FEMS Immunology&Medical Microbiology,2003,35(1):67-75.
[2] AGUILAR-USCANGA B,FRANOIS J M. A study of the yeast cell wall composition and structure in response to growth conditions and mode of cultivation[J]. Letters in Applied Microbiology,2003,37(3):268-274.
[3] SAMUELSEN A B C,SCHREZENMEIR J,KNUTSEN S H. Effects of orally administered yeast-derived beta-glucans:a review[J].Molecular Nutrition&Food Research,2014,58(1):183-193.
[4] LESAGE G,BUSSEY H. Cell wall assembly in Saccharomyces cerevisiae[J]. Microbiology and Molecular Biology Reviews,2006,70(2):317-343.
[5] WILLIAMS D L,MCNAMEE R B,JONES E L,et al.A method for the solubilization of a(1→3)-β-D-glucan isolated from Saccharomyces cerevisiae[J]. Carbohydrate Research,1991,219:203-213.
[6] LIU Q Y,DUAN B C,XU X J,et al.Progress in rigid polysaccharide-based nanocomposites with therapeutic functions[J].Journal of Materials Chemistry B,2017,5(29):5690-5713.
[7] LKKA G,FALK K,AUSTBL,et al. Uptake of yeast cells in the Atlantic salmon(Salmo salar L.)intestine[J]. Developmental&Comparative Immunology,2014,47(1):77-80.
[8] VALLEJOS-VIDAL E,REYES-LPEZ F,TELES M,et al.The response of fish to immunostimulant diets[J].Fish&Shellfish Immunology,2016,56:34-69.
[9] DENNEHY K M,BROWN G D. The role of theβ-glucan receptor Dectin-1 in control of fungal infection[J]. Journal of Leukocyte Biology,2007,82(2):253-258.
[10] XIA Y,ROSS G D. Generation of recombinant fragments of CD11b expressing the functionalβ-glucanbinding lectin site of CR3(CD11b/CD18)[J]. The Journal of Immunology,1999,162(12):7285-7293.
[11] THOMPSON I J,OYSTON P C,WILLIAMSON D E. Potential of theβ-glucans to enhance innate resistance to biological agents[J]. Expert Reviewof Antiinfective Therapy,2010,8(3):339-352.
[12] GOODRIDGE H S,WOLF A J,UNDERHILL D M.β-glucan recognition by the innate immune system[J].Immunological Reviews,2009,230(1):38-50.
[13] VERESCHAGIN E I,VAN LAMBALGEN A A,DUSHKIN MI,et al. Soluble glucan protects against endotoxin shock in the rat:the role of the scavenger receptor[J].Shock,1998,9(3):193-198.
[14] BENARD E L,ROOBOL S J,SPAINK H P,et al.Phagocytosis of mycobacteria by zebrafish macrophages is dependent on the scavenger receptor Marco,a key control factor of pro-inflammatory signalling[J].Developmental&Comparative Immunology,2014,47(2):223-233.
[15] FINK I R,BENARD E L,HERMSEN T,et al.Molecular and functional characterization of the scavenger receptor CD36 in zebrafish and common carp[J].Molecular Immunology,2015,63(2):381-393.
[16] PIETRETTI D,VERA-JIMENEZ N I,HOOLE D,et al. Oxidative burst and nitric oxide responses in carp macrophages induced by zymosan,MacroGard?and selective Dectin-1 agonists suggest recognition by multiple pattern recognition receptors[J].Fish&Shellfish Immunology,2013,35(3):847-857.
[17] ALCORN S W,MURRAY A L,PASCHO R J.Effects of rearing temperature on immune functions in sockeye salmon(Oncorhynchus nerka)[J]. Fish&Shellfish Immunology,2002,12(4):303-334.
[18] DAZ-ROSALES P,MUOZ-ATIENZA E,TAFALLA C. Role of teleost B cells in viral immunity[J].Fish&Shellfish Immunology,2019,86:135-142.
[19]曹雪涛.医学免疫学[M].6版.北京:人民卫生出版社,2013:119-128.
[20] SILVA MT,CORREIA-NEVES M. Neutrophils and macrophages:the main partners of phagocyte cell systems[J].Frontiers in Immunology,2012,3:174.
[21] PETIT J,WIEGERTJES G F.Long-lived effects of administeringβ-glucans:indications for trained immunity in fish[J]. Developmental&Comparative Immunology,2016,64:93-102.
[22] SNCHEZ-MARTNEZ J G,RBAGO-CASTRO J L,DE LA LUZ VZQUEZ-SAUCEDA M,et al.Effect ofβ-glucan dietary levels on immune response and hematology of channel catfish Ictalurus punctatus juveniles[J]. Latin American Journal of Aquatic Research,2017,45(4):690-698.
[23] GUZMN-VILLANUEVA L T,TOVAR-RAMREZ D,GISBERT E,et al.Dietary administration ofβ-1,3/1,6-glucan and probiotic strain Shewanella putrefaciens,single or combined,on gilthead seabream growth,immune responses and gene expression[J].Fish&Shellfish Immunology,2014,39(1):34-41.
[24] DO HUU H,SANG H M,THANH THUY N T.Dietaryβ-glucan improved growth performance,Vibrio counts,haematological parameters and stress resistance of pompano fish,Trachinotus ovatus Linnaeus,1758[J]. Fish&Shellfish Immunology,2016,54:402-410.
[25] MAGNADTTIR B. Innate immunity of fish(overview)[J]. Fish&Shellfish Immunology,2006,20(2):137-151.
[26] DUNKELBERGER J R,SONG W C. Complement and its role in innate and adaptive immune responses[J].Cell Research,2010,20(1):34-50.
[27] AI Q H,MAI K S,ZHANG L,et al.Effects of dietaryβ-1,3 glucan on innate immune response of large yellowcroaker,Pseudosciaena crocea[J]. Fish&Shellfish Immunology,2007,22(4):394-402.
[28] EL-BOSHY ME,EL-ASHRAMA M,ABDELHAMID F M,et al.Immunomodulatory effect of dietary Saccharomyces cerevisiae,β-glucan and laminaran in mercuric chloride treated Nile tilapia(Oreochromis niloticus)and experimentally infected with Aeromonas hydrophila[J]. Fish&Shellfish Immunology,2010,28(5/6):802-808.
[29] DAWOOD MA O,KOSHIO S,EL-SABAGH M,et al. Changes in the growth,humoral and mucosal immune responses followingβ-glucan and vitamin C administration in red sea bream,Pagrus major[J].Aquaculture,2017,470:214-222.
[30] CHETTRI J K,KANIA P W,BUCHMANN K.Immunomodulation of rainbowtrout(Oncorhynchus mykiss)fry by bath exposure to aβ-glucan from Euglena gracilis[J]. Aquaculture Research,2013,44(9):1407-1415.
[31] PIONNIER N,FALCO A,MIEST J,et al.Dietaryβ-glucan stimulate complement and C-reactive protein acute phase responses in common carp(Cyprinus carpio)during an Aeromonas salmonicida infection[J].Fish&Shellfish Immunology,2013,34(3):819-831.
[32] PIONNIER N,FALCO A,MIEST J J,et al. Feeding common carp Cyprinus carpio withβ-glucan supplemented diet stimulates C-reactive protein and complement immune acute phase responses following PAMPs injection[J]. Fish&Shellfish Immunology,2014,39(2):285-295.
[33] LOKESH J,FERNANDES J MO,KORSNES K,et al.Transcriptional regulation of cytokines in the intestine of Atlantic cod fed yeast derived mannan oligosaccharide orβ-glucan and challenged with Vibrio anguillarum[J]. Fish&Shellfish Immunology,2012,33(3):626-631.
[34] FALCO A,FROST P,MIEST J,et al.Reduced inflammatory response to Aeromonas salmonicida infection in common carp(Cyprinus carpio L.)fed withβ-glucan supplements[J].Fish&Shellfish Immunology,2012,32(6):1051-1057.
[35] FALCO A,MIEST J J,PIONNIER N,et al.β-glucansupplemented diets increase poly(I:C)-induced gene expression of Mx,possibly via TLR3-mediated recognition mechanism in common carp(Cyprinus carpio)[J].Fish&Shellfish Immunology,2014,36(2):494-502.
[36] MIEST J J,ARNDT C,ADAMEK M,et al.Dietaryβ-glucan(MacroGard?)enhances survival of first feeding turbot(Scophthalmus maximus)larvae by altering immunity,metabolism and microbiota[J]. Fish&Shellfish Immunology,2016,48:94-104.
[37] BISWAS G,KORENAGA H,TAKAYAMA H,et al.Cytokine responses in the common carp,Cyprinus carpio L. treated with baker’s yeast extract[J].Aquaculture,2012,356-357:169-175.
[38] SIRIMANAPONG W,THOMPSON K D,OOI E L,et al. The effects of feedingβ-glucan to Pangasianodon hypophthalmus on immune gene expression and resistance to Edwardsiella ictaluri[J]. Fish&Shellfish Immunology,2015,47(1):595-605.
[39] JIANG C Y,WANG P,LI MY,et al.Dietaryβ-glucan enhances the contents of complement component 3and factor B in eggs of zebrafish[J].Developmental&Comparative Immunology,2016,65:107-113.
[40] WANG P,JIANG C Y,LIU S S,et al.Trans-generational enhancement of C-type lysozyme level in eggs of zebrafish by dietaryβ-glucan[J]. Developmental&Comparative Immunology,2017,74:25-31.
[41] LAZADO C C,CAIPANG C MA.Mucosal immunity and probiotics in fish[J]. Fish&Shellfish Immunology,2014,39(1):78-89.
[42] ROMBOUT J H W M,YANG G W,KIRON V. Adaptive immune responses at mucosal surfaces of teleost fish[J]. Fish&Shellfish Immunology,2014,40(2):634-643.
[43] PARRA D,REYES-LOPEZ F E,TORT L. Mucosal immunity and B cells in teleosts:effect of vaccination and stress[J].Frontiers in Immunology,2015,6:354.
[44] DALMO R A,BGWALD J.β-glucans as conductors of immune symphonies[J].Fish&Shellfish Immunology,2008,25(4):384-396.
[45] SCHMITT P,WACYK J,MORALES-LANGE B,et al. Immunomodulatory effect of cathelicidins in response to aβ-glucan in intestinal epithelial cells from rainbowtrout[J]. Developmental&Comparative Immunology,2015,51(1):160-169.
[46] KIRON V,KULKARNI A,DAHLE D,et al.Recognition of purified beta 1,3/1,6 glucan and molecular signalling in the intestine of Atlantic salmon[J]. Developmental&Comparative Immunology,2016,56:57-66.
[47] SWENNEN K,COURTIN C M,DELCOUR J A.Nondigestible oligosaccharides with prebiotic properties[J].Critical Reviews in Food Science and Nutrition,2006,46(6):459-471.
[48] KHLWEIN H,EMERY MJ,RAWLING MD,et al.Effects of a dietaryβ-(1,3)(1,6)-D-glucan supplementation on intestinal microbial communities and intestinal ultrastructure of mirror carp(Cyprinus carpio L.)[J]. Journal of Applied Microbiology,2013,115(5):1091-1106.
[49] JUNG-SCHROERS V,ADAMEK M,JUNG A,et al.Feeding ofβ-1,3/1,6-glucan increases the diversity of the intestinal microflora of carp(Cyprinus carpio)[J].Aquaculture Nutrition,2016,22(5):1026-1039.
[50] CARDA-DIGUEZ M,MIRA A,FOUZ B. Pyrosequencing survey of intestinal microbiota diversity in cultured sea bass(Dicentrarchus labrax)fed functional diets[J]. FEMS Microbiology Ecology,2014,87(2):451-459.
[51] KAWASHIMA T,KOSAKA A,YAN H M,et al.Double-stranded RNA of intestinal commensal but not pathogenic bacteria triggers production of protective interferon-β[J].Immunity,2013,38(6):1187-1197.
[52]石纪奎,李永富,史锋.机械活化酿酒酵母葡聚糖改善其水溶性[J].食品与生物技术学报,2015,34(5):536-541.
[53] PENGKUMSRI N,SIVAMARUTHI B S,SIRILUN S,et al. Extraction ofβ-glucan from Saccharomyces cerevisiae:comparison of different extraction methods and in vivo assessment of immunomodulatory effect in mice[J]. Food Science and Technology,2017,37(1):124-130.
[54] LIU X Y,WANG Q,CUI S W,et al.A newisolation method ofβ-D-glucans from spent yeast Saccharomyces cerevisiae[J]. Food Hydrocolloids,2008,22(2):239-247.
[2] AGUILAR-USCANGA B,FRANOIS J M. A study of the yeast cell wall composition and structure in response to growth conditions and mode of cultivation[J]. Letters in Applied Microbiology,2003,37(3):268-274.
[3] SAMUELSEN A B C,SCHREZENMEIR J,KNUTSEN S H. Effects of orally administered yeast-derived beta-glucans:a review[J].Molecular Nutrition&Food Research,2014,58(1):183-193.
[4] LESAGE G,BUSSEY H. Cell wall assembly in Saccharomyces cerevisiae[J]. Microbiology and Molecular Biology Reviews,2006,70(2):317-343.
[5] WILLIAMS D L,MCNAMEE R B,JONES E L,et al.A method for the solubilization of a(1→3)-β-D-glucan isolated from Saccharomyces cerevisiae[J]. Carbohydrate Research,1991,219:203-213.
[6] LIU Q Y,DUAN B C,XU X J,et al.Progress in rigid polysaccharide-based nanocomposites with therapeutic functions[J].Journal of Materials Chemistry B,2017,5(29):5690-5713.
[7] LKKA G,FALK K,AUSTBL,et al. Uptake of yeast cells in the Atlantic salmon(Salmo salar L.)intestine[J]. Developmental&Comparative Immunology,2014,47(1):77-80.
[8] VALLEJOS-VIDAL E,REYES-LPEZ F,TELES M,et al.The response of fish to immunostimulant diets[J].Fish&Shellfish Immunology,2016,56:34-69.
[9] DENNEHY K M,BROWN G D. The role of theβ-glucan receptor Dectin-1 in control of fungal infection[J]. Journal of Leukocyte Biology,2007,82(2):253-258.
[10] XIA Y,ROSS G D. Generation of recombinant fragments of CD11b expressing the functionalβ-glucanbinding lectin site of CR3(CD11b/CD18)[J]. The Journal of Immunology,1999,162(12):7285-7293.
[11] THOMPSON I J,OYSTON P C,WILLIAMSON D E. Potential of theβ-glucans to enhance innate resistance to biological agents[J]. Expert Reviewof Antiinfective Therapy,2010,8(3):339-352.
[12] GOODRIDGE H S,WOLF A J,UNDERHILL D M.β-glucan recognition by the innate immune system[J].Immunological Reviews,2009,230(1):38-50.
[13] VERESCHAGIN E I,VAN LAMBALGEN A A,DUSHKIN MI,et al. Soluble glucan protects against endotoxin shock in the rat:the role of the scavenger receptor[J].Shock,1998,9(3):193-198.
[14] BENARD E L,ROOBOL S J,SPAINK H P,et al.Phagocytosis of mycobacteria by zebrafish macrophages is dependent on the scavenger receptor Marco,a key control factor of pro-inflammatory signalling[J].Developmental&Comparative Immunology,2014,47(2):223-233.
[15] FINK I R,BENARD E L,HERMSEN T,et al.Molecular and functional characterization of the scavenger receptor CD36 in zebrafish and common carp[J].Molecular Immunology,2015,63(2):381-393.
[16] PIETRETTI D,VERA-JIMENEZ N I,HOOLE D,et al. Oxidative burst and nitric oxide responses in carp macrophages induced by zymosan,MacroGard?and selective Dectin-1 agonists suggest recognition by multiple pattern recognition receptors[J].Fish&Shellfish Immunology,2013,35(3):847-857.
[17] ALCORN S W,MURRAY A L,PASCHO R J.Effects of rearing temperature on immune functions in sockeye salmon(Oncorhynchus nerka)[J]. Fish&Shellfish Immunology,2002,12(4):303-334.
[18] DAZ-ROSALES P,MUOZ-ATIENZA E,TAFALLA C. Role of teleost B cells in viral immunity[J].Fish&Shellfish Immunology,2019,86:135-142.
[19]曹雪涛.医学免疫学[M].6版.北京:人民卫生出版社,2013:119-128.
[20] SILVA MT,CORREIA-NEVES M. Neutrophils and macrophages:the main partners of phagocyte cell systems[J].Frontiers in Immunology,2012,3:174.
[21] PETIT J,WIEGERTJES G F.Long-lived effects of administeringβ-glucans:indications for trained immunity in fish[J]. Developmental&Comparative Immunology,2016,64:93-102.
[22] SNCHEZ-MARTNEZ J G,RBAGO-CASTRO J L,DE LA LUZ VZQUEZ-SAUCEDA M,et al.Effect ofβ-glucan dietary levels on immune response and hematology of channel catfish Ictalurus punctatus juveniles[J]. Latin American Journal of Aquatic Research,2017,45(4):690-698.
[23] GUZMN-VILLANUEVA L T,TOVAR-RAMREZ D,GISBERT E,et al.Dietary administration ofβ-1,3/1,6-glucan and probiotic strain Shewanella putrefaciens,single or combined,on gilthead seabream growth,immune responses and gene expression[J].Fish&Shellfish Immunology,2014,39(1):34-41.
[24] DO HUU H,SANG H M,THANH THUY N T.Dietaryβ-glucan improved growth performance,Vibrio counts,haematological parameters and stress resistance of pompano fish,Trachinotus ovatus Linnaeus,1758[J]. Fish&Shellfish Immunology,2016,54:402-410.
[25] MAGNADTTIR B. Innate immunity of fish(overview)[J]. Fish&Shellfish Immunology,2006,20(2):137-151.
[26] DUNKELBERGER J R,SONG W C. Complement and its role in innate and adaptive immune responses[J].Cell Research,2010,20(1):34-50.
[27] AI Q H,MAI K S,ZHANG L,et al.Effects of dietaryβ-1,3 glucan on innate immune response of large yellowcroaker,Pseudosciaena crocea[J]. Fish&Shellfish Immunology,2007,22(4):394-402.
[28] EL-BOSHY ME,EL-ASHRAMA M,ABDELHAMID F M,et al.Immunomodulatory effect of dietary Saccharomyces cerevisiae,β-glucan and laminaran in mercuric chloride treated Nile tilapia(Oreochromis niloticus)and experimentally infected with Aeromonas hydrophila[J]. Fish&Shellfish Immunology,2010,28(5/6):802-808.
[29] DAWOOD MA O,KOSHIO S,EL-SABAGH M,et al. Changes in the growth,humoral and mucosal immune responses followingβ-glucan and vitamin C administration in red sea bream,Pagrus major[J].Aquaculture,2017,470:214-222.
[30] CHETTRI J K,KANIA P W,BUCHMANN K.Immunomodulation of rainbowtrout(Oncorhynchus mykiss)fry by bath exposure to aβ-glucan from Euglena gracilis[J]. Aquaculture Research,2013,44(9):1407-1415.
[31] PIONNIER N,FALCO A,MIEST J,et al.Dietaryβ-glucan stimulate complement and C-reactive protein acute phase responses in common carp(Cyprinus carpio)during an Aeromonas salmonicida infection[J].Fish&Shellfish Immunology,2013,34(3):819-831.
[32] PIONNIER N,FALCO A,MIEST J J,et al. Feeding common carp Cyprinus carpio withβ-glucan supplemented diet stimulates C-reactive protein and complement immune acute phase responses following PAMPs injection[J]. Fish&Shellfish Immunology,2014,39(2):285-295.
[33] LOKESH J,FERNANDES J MO,KORSNES K,et al.Transcriptional regulation of cytokines in the intestine of Atlantic cod fed yeast derived mannan oligosaccharide orβ-glucan and challenged with Vibrio anguillarum[J]. Fish&Shellfish Immunology,2012,33(3):626-631.
[34] FALCO A,FROST P,MIEST J,et al.Reduced inflammatory response to Aeromonas salmonicida infection in common carp(Cyprinus carpio L.)fed withβ-glucan supplements[J].Fish&Shellfish Immunology,2012,32(6):1051-1057.
[35] FALCO A,MIEST J J,PIONNIER N,et al.β-glucansupplemented diets increase poly(I:C)-induced gene expression of Mx,possibly via TLR3-mediated recognition mechanism in common carp(Cyprinus carpio)[J].Fish&Shellfish Immunology,2014,36(2):494-502.
[36] MIEST J J,ARNDT C,ADAMEK M,et al.Dietaryβ-glucan(MacroGard?)enhances survival of first feeding turbot(Scophthalmus maximus)larvae by altering immunity,metabolism and microbiota[J]. Fish&Shellfish Immunology,2016,48:94-104.
[37] BISWAS G,KORENAGA H,TAKAYAMA H,et al.Cytokine responses in the common carp,Cyprinus carpio L. treated with baker’s yeast extract[J].Aquaculture,2012,356-357:169-175.
[38] SIRIMANAPONG W,THOMPSON K D,OOI E L,et al. The effects of feedingβ-glucan to Pangasianodon hypophthalmus on immune gene expression and resistance to Edwardsiella ictaluri[J]. Fish&Shellfish Immunology,2015,47(1):595-605.
[39] JIANG C Y,WANG P,LI MY,et al.Dietaryβ-glucan enhances the contents of complement component 3and factor B in eggs of zebrafish[J].Developmental&Comparative Immunology,2016,65:107-113.
[40] WANG P,JIANG C Y,LIU S S,et al.Trans-generational enhancement of C-type lysozyme level in eggs of zebrafish by dietaryβ-glucan[J]. Developmental&Comparative Immunology,2017,74:25-31.
[41] LAZADO C C,CAIPANG C MA.Mucosal immunity and probiotics in fish[J]. Fish&Shellfish Immunology,2014,39(1):78-89.
[42] ROMBOUT J H W M,YANG G W,KIRON V. Adaptive immune responses at mucosal surfaces of teleost fish[J]. Fish&Shellfish Immunology,2014,40(2):634-643.
[43] PARRA D,REYES-LOPEZ F E,TORT L. Mucosal immunity and B cells in teleosts:effect of vaccination and stress[J].Frontiers in Immunology,2015,6:354.
[44] DALMO R A,BGWALD J.β-glucans as conductors of immune symphonies[J].Fish&Shellfish Immunology,2008,25(4):384-396.
[45] SCHMITT P,WACYK J,MORALES-LANGE B,et al. Immunomodulatory effect of cathelicidins in response to aβ-glucan in intestinal epithelial cells from rainbowtrout[J]. Developmental&Comparative Immunology,2015,51(1):160-169.
[46] KIRON V,KULKARNI A,DAHLE D,et al.Recognition of purified beta 1,3/1,6 glucan and molecular signalling in the intestine of Atlantic salmon[J]. Developmental&Comparative Immunology,2016,56:57-66.
[47] SWENNEN K,COURTIN C M,DELCOUR J A.Nondigestible oligosaccharides with prebiotic properties[J].Critical Reviews in Food Science and Nutrition,2006,46(6):459-471.
[48] KHLWEIN H,EMERY MJ,RAWLING MD,et al.Effects of a dietaryβ-(1,3)(1,6)-D-glucan supplementation on intestinal microbial communities and intestinal ultrastructure of mirror carp(Cyprinus carpio L.)[J]. Journal of Applied Microbiology,2013,115(5):1091-1106.
[49] JUNG-SCHROERS V,ADAMEK M,JUNG A,et al.Feeding ofβ-1,3/1,6-glucan increases the diversity of the intestinal microflora of carp(Cyprinus carpio)[J].Aquaculture Nutrition,2016,22(5):1026-1039.
[50] CARDA-DIGUEZ M,MIRA A,FOUZ B. Pyrosequencing survey of intestinal microbiota diversity in cultured sea bass(Dicentrarchus labrax)fed functional diets[J]. FEMS Microbiology Ecology,2014,87(2):451-459.
[51] KAWASHIMA T,KOSAKA A,YAN H M,et al.Double-stranded RNA of intestinal commensal but not pathogenic bacteria triggers production of protective interferon-β[J].Immunity,2013,38(6):1187-1197.
[52]石纪奎,李永富,史锋.机械活化酿酒酵母葡聚糖改善其水溶性[J].食品与生物技术学报,2015,34(5):536-541.
[53] PENGKUMSRI N,SIVAMARUTHI B S,SIRILUN S,et al. Extraction ofβ-glucan from Saccharomyces cerevisiae:comparison of different extraction methods and in vivo assessment of immunomodulatory effect in mice[J]. Food Science and Technology,2017,37(1):124-130.
[54] LIU X Y,WANG Q,CUI S W,et al.A newisolation method ofβ-D-glucans from spent yeast Saccharomyces cerevisiae[J]. Food Hydrocolloids,2008,22(2):239-247.