Mycologia
HOME HELP FEEDBACK SUBSCRIPTIONS ARCHIVE SEARCH TABLE OF CONTENTS
DOI: 10.3852/mycologia.98.2.180
This Article
Right arrow Abstract Freely available
Right arrow Full Text (PDF)
Services
Right arrow Similar articles in this journal
Right arrow Alert me to new issues of the journal
Right arrow Download to citation manager
Right arrow reprints & permissions
Citing Articles
Right arrow Citing Articles via Google Scholar
Google Scholar
Right arrow Articles by Pacheco-Sanchez, M.
Right arrow Articles by Tweddell, R. J.
Right arrow Search for Related Content
PubMed
Right arrow Articles by Pacheco-Sanchez, M.
Right arrow Articles by Tweddell, R. J.
Agricola
Right arrow Articles by Pacheco-Sanchez, M.
Right arrow Articles by Tweddell, R. J.
Mycologia, 98(2), 2006, pp. 180-185.
© 2006 by The Mycological Society of America

A bioactive (1->3)-, (1->4)-ß-D-glucan from Collybia dryophila and other mushrooms


Maribel Pacheco-Sanchez

     Centre de recherche en horticulture, Pavillon de l’Envirotron, Université Laval, Québec QC, G1K 7P4 Canada, and Institut des nutraceutiques et des aliments fonctionnels, Université Laval, Québec QC, G1K 7P4 Canada

Yvan Boutin

     Institut des nutraceutiques et des aliments fonctionnels, Université Laval, Québec QC, G1K 7P4 Canada, Transbiotech, Cégep Lévis-Lauzon, 205 route Mgr Bourget, Lévis QC, G6V 6Z9 Canada

Paul Angers

     Institut des nutraceutiques et des aliments fonctionnels, Université Laval, Québec QC, G1K 7P4 Canada

André Gosselin
Russell J. Tweddell 1

     Centre de recherche en horticulture, Pavillon de l’Envirotron, Université Laval, Québec QC, G1K 7P4 Canada, and Institut des nutraceutiques et des aliments fonctionnels, Université Laval, Québec QC, G1K 7P4 Canada

   ABSTRACT
TOP
 ABSTRACT
INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 LITERATURE CITED
 

Polysaccharides from higher Basidiomycete mushrooms, mainly ß-D-glucans, are considered to be potent bioactive fungal compounds. In this study a ß-glucan (1.237 x 106 Da) consisting of (1-> 3) and (1-> 4) glucosidic linkages, named Collybia dryophila polysaccharide (CDP), was extracted from the wild mushroom C. dryophila. CDP was shown to strongly inhibit nitric oxide production in activated macrophages suggesting that this polysaccharide displays a potential anti-inflammatory activity. In addition it was shown that polysaccharides similar to CDP (CDP-like) are present in Lentinus edodes and different wild mushrooms collected in northeastern North America.

Key words: anti-inflammatory activity, higher Basidiomycetes, Lentinus edodes, Marasmius oreades, polysaccharides


   INTRODUCTION
TOP
 ABSTRACT
 INTRODUCTION
MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 LITERATURE CITED
 
Carbohydrates are present in fungi mainly as polysaccharides occurring both exocellularly and intracellularly as insoluble fibrillar and amorphous wall components, soluble gums and storage polysaccharides. As components of the fungal cell wall, ß-linked polysaccharides (chitin, chitosan, ß-glucans) form microcrystalline fibrils within the wall that are responsible for the strength and rigidity of the wall structure (Griffin 1994Go). In addition to water-soluble ß-D-glucans, ß-D-glucans exist with heterosaccharides chains of xylose, mannose, galactose and uronic acids as well as with proteins (Wasser and Weis 1999Go).

Several bioactive (hypoglycemic, immunomodulatory, anti-inflammatory, antitumor, antiviral, antibacterial or antiparasitic activities) compounds, such as polysaccharides, polysaccharides-peptides, nucleosides and triterpenols, have been identified in numerous mushroom species (Wasser and Weis 1999Go). Among these classes of bioactive molecules, polysaccharides are considered to be the most potent active compounds (Borchers et al 1999Go). A recent review (Wasser 2002Go) indicates that at least 651 species and seven intraspecific taxa, representing 182 genera of Basidiomycete mushrooms, contain antitumor or immunomodulatory polysaccharides that have unique structures. Most of these bioactive polysaccharides are (1->3)-ß-D-glucans with (1-> 6)-ß linked side branches (Ooi and Liu 1999Go, Schmid et al 2001Go), such as grifolan from Grifola frondosa (Dick. : Fr.) S.F. Gray, schizophyllan from Schizophyllum commune Fr. and lentinan from Lentinus edodes (Berk.) Sing. (Borchers et al 1999Go, Mizuno 2000Go, Yan et al 2003Go), with the latter being the most widely investigated fungal polysaccharide in medical research (Chihara 1992Go, Jong and Birmingham 1993Go, Liu et al 1999Go).

The principle on which the immunomodulator compounds operate is based primarily on the modulation of different immune cells that may result in several therapeutic properties such as anti-inflammatory activity (Pugh et al 2001Go). In some cases the anti-inflammatory activity is attributable to the inhibition of nitric oxide (NO) production by activated macrophages. Several natural compounds extracted from different plants including 2'-hydroxycinnamal-dehyde from Cinnamomum cassia Nees ex Blume (Lee et al 2005Go), saponins from Platycodon grandiflorum ( Jacq.) A. DC. (Ahn et al 2005Go) and panduratin A from Kaempferia pandurata Roxb. (Yun et al 2003Go) were shown to possess anti-inflammatory properties resulting from their inhibitory effect on NO biosynthesis by macrophages. Among mushrooms, a (1->6)-branched (1->3)-ß-D-glucan (T-5-N) from Dictyophora indusiata (Vent. : Pers.) Fisch (Hara et al 1982Go) as well as different polysaccharides, including MEA, MHA, MCW-A, MCW-N from Auricularia auriculajudea (Bull.) Wettst. and G-A from Ganoderma japonicum (Fr.) Bres., were reported to display anti-inflammatory activity (Ukai et al 1983Go, Ooi and Liu 1999Go).

This paper reports the purification and partial characterization of a polysaccharide with potential anti-inflammatory activity named Collybia dryophila polysaccharide (CDP) that was extracted from Collybia dryophila (Fr.) Kum., a wild mushroom commonly found in northeastern North America. The presence of similar polysaccharides (CDP-like) in other mushrooms including Lentinus edodes also is reported.


   MATERIALS AND METHODS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
RESULTS
 DISCUSSION
 LITERATURE CITED
 
Mushrooms.— – Fruiting bodies of C. dryophila and of 19 different species of wild mushroom (Agaricus arvensis, Amanita muscaria, A. rubescens, Coprinus atramentarius (syn. Coprinopsis atramentaria), C. comatus, Hydnum imbricatum, Lactarius deliciosus, Leccinum aurantiacum, L. subglabripes, Lepiota americana, Lycoperdon pyriforme, Marasmius oreades, Panellus serotinus, Piptoporus betulinus, Polyporus squamosus, Russula variata, Suillus americanus, Tricholoma flavovirens, T. vaccinum) were collected in the Province of Québec (Canada) in 2001. Fruiting bodies of Lentinus edodes were purchased from a commercial grower. Fruiting bodies from different growth stages were pooled, freeze-dried, homogenized and ground into a powder.

Extraction and purification of polysaccharides from mushrooms.— – Each freeze-dried sample (20 g) of the fruiting bodies was extracted with 85%EtOH (350 mL) at 80 C for 2–3 h to remove low molecular mass compounds (Robyt 1998Go) and the residue was extracted with 700 mL of hot water (98 C) for 8 h. The supernatant was recovered after centrifugation and diluted with an equal volume of EtOH. The precipitate was collected after centrifugation and resuspended in 20%trichlor-oacetic acid aqueous solution to remove proteins (Cerning et al 1994Go). Three volumes of 98%EtOH then were added to the filtrate and the precipitate was recovered after centrifugation, dissolved in water, dialyzed (molecular weight cut-off 8000; Spectra/Por, Spectrum Laboratories Inc., Rancho Dominguez, California) against tap water for 72 h at 4 C (Cerning et al 1994Go, Marshall et al 1995Go) and freeze-dried. This powder (the polysaccharide of interest) was weighed and its concentration was expressed as mg/g dry weight of mushrooms.

Size-exclusion chromatography (SEC).— – Purified polysaccharide samples (2 mg) were dissolved in 2 mL of deionized water containing 20 µL of 1%sodium hydroxide, vortexed and filtered through 0.45 µm Whatman filter paper. Homogeneity and molecular weight (Mr) of the polysaccharides were determined by HPLC-SEC (Millipore, Milford, Michigan) with a TSK G-6000 PWXL column (7.8 x 300 mm; TosoHaas, Montgomeryville, Pennsylvania). Elution was performed at 25 C with 0.1%NaNO3 aqueous solution at a flow rate of 0.5 mL/min. Peaks were detected with a differential refractive index (RI-150; Millipore). Mr was estimated by comparison with retention time of high-molecular-weight dextran standards (9.7 x 104, 1.46 x 105, 3.26 x 105, 5.74 x 105, 8.48 x 105, 2.45 x 106 and 3.8 x 106 Da). The calibration curve was Log Mr '0.4453Rt µ13.86, where Rt represents the retention time (Netopilìk 2003Go).

Enzymatic hydrolysis.— – Fifty µL of endo-ß-glucanase (1.25 units) from Bacillus subtilis, (1->3)/(1->3)-, (1->4)-ß-D-glucan 3(4)-glucanohydrolase (EC 3.2.1.6 [EC] ) (Sigma-Aldrich, St Louis, Missouri), were incubated at 37 C with 2 mg/mL of each purified polysaccharide, laminarin (Sigma), oat (1->3)/(1->4)-ß-glucan (Sigma) or pachyman (Megazyme, Wicklow, Ireland) in 0.5 mL of sodium acetate buffer (0.1 M, pH 5.6). After 24 h the reaction was stopped by boiling (10 min). The supernatant was recovered by centrifugation (13 000 rpm, 10 min, room temperature), filtered through 0.45 µm Whatman filter paper to remove impurities and subjected to ion-exchange chromatography on a Supercosil C-116 column (250 x 4.6 mm, Sigma). Elution was performed at 60 C with NaOH (1 x 10'4 N) aqueous solution at a constant flow of 0.5 mL/min. Peaks were detected with a differential refractive index (RI-150). Laminaridextrins, cellobiose, gentobiose and glucose (Sigma) were used as standards. Degree of hydrolysis was estimated as the ratio of reducing sugar in the supernatant to total polysaccharide content of the suspension. Experiments were performed in triplicate.

Nitrite assay.— – The test used to assess the potential anti-inflammatory activity of molecules consisted of evaluating their capacity to inhibit NO production in activated macrophages (Yun et al 2003Go, Lee et al 2005Go). Murine macrophages (cell line RAW 264.7, 2 x 105 cells/mL) were cultured in phenol red-free Dulbecco’s modified Eagle medium (DMEM; Invitrogen Inc., Burlington, Ontario, Canada) containing 100 U/mL penicillin, 100 µg/mL streptomycin, 2 mM glutamine and 5%heat-inactivated fetal bovine serum (FBS; Bio Media Canada Inc., Drummondville, Québec, Canada) and activated with lipopolysaccharide (LPS, 0.05 µg/mL; Sigma) and gamma interferon (IFN{gamma}, 0.1 ng/mL; Sigma). Activated macrophages were incubated with purified polysaccharides (0 [control], 50 and 250 µg/ mL) for 48 h at 37 C in a humidified atmosphere (ca. 95 %) containing 5%CO2, without agitation. After incubation nitrite concentration was used as an indicator of NO released in the medium and was measured with the Griess reaction (Green et al 1982Go). Briefly, 150 µL of culture medium were mixed with 150 µL of Griess reagent (1%sulphanilamide and 0.1%naphthyl-ethylenediamide in 5%phosphoric acid). After 30 min at room temperature, the optical density was measured at 548 nm with an automated plate reader (Tecan Genios, Gröding/Salzburg, Austria). Calibration curves were made with NaNO2. The experiment was performed in triplicate. Results were expressed in percentage of inhibition of NO production: (Nitrite concentration [control] ' Nitrite concentration [purified polysaccharides])/Nitrite concentration [control] x 100.


   RESULTS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
DISCUSSION
 LITERATURE CITED
 
A water-soluble polysaccharide, named CDP, was extracted and purified from the fruiting body of C. dryophila. After concentration, dialysis and lyophilization, the yield of CDP was 35.1 mg/g dry weight. Purified CDP was only slightly soluble in hot water but freely soluble in a 1%sodium hydroxide solution. Chromatographic analysis of purified CDP was performed with HPLC-SEC. Purified CDP gave a single peak with a Mr estimated at 1.237 x 106 Da (FIG. 1Go), based on the calibration curve of the retention time of dextran standards.


Figure 1
View larger version (5K):
[in this window]
[in a new window]
  		FIG. 1. SEC chromatogram of CDP in 0.1%NaNO3 aqueous solution at 25 C using TSK G-6000 PWXL column (7.8 x 300 mm) and RI detector.

 
Following the same procedure water-soluble polysaccharides were extracted and purified from A. rubescens, C. comatus, L. deliciosus, L. edodes, M. oreades, P. serotinus and S. americanus. The polysaccharides showed a homogeneous weight distribution (a single narrow peak) and a Mr similar to CDP, ranging from 6.099 x 105 to 1.550 x 106 (TABLE IGo). These polysaccharides were named CDP-like. A. arvensis, A. muscaria, C. atramentarius, H. imbricatum, L. aurantiacum, L. subglabripes, L. americana, L. pyriforme, P. betulinus, P. squamosus, R. variata, T. flavovirens and T. vaccinum did not contain CDP-like.


View this table:
[in this window]
[in a new window]
  		TABLE I. CDP-like yield and weight distribution in Lentinus edodes 2nd in different wild mushrooms collected in the Province of Québec (Canada)

 
To further characterize CDP and CDP-like, polysaccharide hydrolysis products were analyzed after exhaustive digestion with an endo-ß-glucanase from B. subtilis. This enzyme first was tested against glucans including pachyman, laminarin and oat glucan. The enzyme was shown not to hydrolyze pachyman (data not shown) and to weakly hydrolyze laminarin releasing oligosaccharides with a degree of polymerization higher than 6 (FIG. 2AGo) whereas it strongly hydrolyzed oat glucan (FIG. 2BGo). In the case of oat glucan main products of hydrolysis were laminaritriose and laminaritetraose and no cellobiose was observed as products of hydrolysis. Incubation of CDP with B. subtilis endo-ß-glucanase resulted in a strong hydrolysis of the polysaccharide (degree of hydrolysis of 69%). The major products of hydrolysis were laminaritriose, laminaribiose and glucose (FIG. 2CGo) in a 2 : 1 : 1 ratio. Hydrolysis profiles of CDP-like extracted from A. rubescens (FIG. 3AGo), L. edodes (FIG. 3BGo), M. oreades (FIG. 3CGo), as well as from other CDP-like containing mushrooms (data not shown), were comparable to that displayed by CDP.


Figure 2
View larger version (7K):
[in this window]
[in a new window]
  		FIG. 2. Fractionation by ion-exchanger chromatography using Supercosil C-116 column and RI detector of saccharides released from laminarin (A), oat glucan (B), and CDP from Collybia dryophila (C) after exhaustive hydrolysis with Bacillus subtilis ß-glucanase. Glucose (1), Laminaribiose (2), Laminaritriose (3), Laminaritetraose (4), Oligosaccharides with a degree of polymerization ' 6 (5). Peak not identified (P).

 

Figure 3
View larger version (7K):
[in this window]
[in a new window]
  		FIG. 3. Fractionation by ion-exchanger chromatography using Supercosil C-116 column and RI detector of saccharides released from CDP-like from Amanita rubescens (A), Lentinus edodes (B) and Marasmius oreades (C) after exhaustive hydrolysis with Bacillus subtilis ß-glucanase. Glucose (1), Laminaribiose (2), Laminaritriose (3), Oligosaccharides with a degree of polymerization '6 (4). Peak not identified (P).

 
The capacity to inhibit NO production in activated macrophages by CDP and CDP-like in RAW 264.7 activated macrophages was evaluated to assess their potential anti-inflammatory activity. The results showed a marked inhibition of NO production, measured as nitrite formation, when activated macrophages were cultured in the presence of CDP and CDP-like (FIG. 4Go), particularly when a concentration of 250 µ g/mL was used. Indeed at this concentration CDP from C. dryophila and CDP-like from L. edodes and M. oreades caused a 93.3%, 90.5%and 81.6%inhibition of NO production respectively; this inhibition appeared to be dose dependent.


Figure 4
View larger version (14K):
[in this window]
[in a new window]
  		FIG. 4. Effect of different concentrations of CDP from Collybia dryophila ({square}) and CDP-like from Lentinus edodes ({blacksquare}) and Marasmius oreades ({blacksquare}) on NO production in macrophages RAW 264.7 activated with LPS and IFN{gamma}. Percentage of inhibition of NO production was calculated thusly: (nitrite concentration [control] ' Nitrite concentration [CDP or CDP-like])/nitrite concentration [control] x 100. Each value represents the mean of three replicates x SD. The numbers in brackets above the bars indicate the concentration of nitrite released (µ M) in the culture medium.

 

   DISCUSSION
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
LITERATURE CITED
 
Higher Basidiomycete mushrooms have been used in folk medicine throughout the world since ancient times. In the past decade several medicinally active Basidiomycetes were developed commercially (Wasser and Weis 1999Go) and it is now well established that mushrooms represent a source for the development of drugs (Mizuno 2000Go). In this study a water-soluble polysaccharide with an estimated Mr of 1.237 x 106 Da was purified from C. dryophila and named CDP. Chromatographic analysis of CDP revealed a single peak indicating homogeneous molecular weight or narrow weight distribution of the polysaccharide and appropriate isolation and purification procedures.

Following the procedure used for C. dryophila CDP extraction and purification, polysaccharides were extracted and purified from L. edodes and different wild mushrooms collected in Québec. Polysaccharides showing a homogeneous weight distribution (a single narrow peak) and a Mr comparable to that of C. dryophila CDP were considered to be CDP-like. According to this premise, several mushrooms were shown to contain CDP-like. Among the wild mushrooms tested, C. comatus, A. rubescens, P. serotinus and M. oreades showed the highest contents of CDP-like, whereas CDP-like content in the commercially grown mushroom L. edodes was estimated to be 11.0 mg/g dry weight. Several wild mushrooms were shown to contain no CDP-like.

Further characterization of CDP by enzymatic hydrolysis with B. subtilis endo-ß-glucanase was performed to determine the type of glucosidic linkage within the polysaccharide. Initially endo-ß-glucanase hydrolysis of known polysaccharides was accomplished to confirm the reported selective hydrolysis of (1->3)-or (1->4)-linkages when glucose residue is substituted at C-3 (IUBMB 1992Go, Sakurai 2000Go). Results from the strong hydrolysis of oat glycan ([1->3], [1->4]-ß-glucan), the weak hydrolysis of laminarin (essentially a linear [1->3]-ß-glucan) as well as the lack of hydrolysis of pachyman ([1->3]-ß-glucan with [1->6]-ß ramifications) suggests that the enzyme hydrolyzed almost exclusively glucans containing (1->3)-ß and (1->4)-ß glucosidic linkages and hydrolyzed preferentially (1->4)-ß linkages in (1->3), (1->4)-ß-glucan. Incubation of CDP with endo-ß-glucanase resulted in a strong hydrolysis of the polysaccharide and the major products of CDP hydrolysis were laminaritriose, laminaribiose and glucose. These results strongly suggest CDP structure as a ß-glucan consisting of (1-> 3) and (1->4) linkages.

For mushrooms containing CDP-like, the hydrolysis profiles of these polysaccharides are comparable to that displayed by CDP extracted from C. dryophila, indicating that CDP-like from these mushrooms are also (1->3)-, (1->4)-ß-D-glucans. Few fungi are reported to contain (1->3)-, (1->4)-ß-D-glucans. Indeed to our knowledge only polysaccharides extracted from Achlya ambisexualis Raper, Monilia fructicola (G. Wint.) Honey and Armillaria mellea (Fr.) Kum. were reported to contain (1->3)- and (1->4)-ß linked glucose groups (Ruiz-Herrera 1992Go). Of particular interest, this study reports for the first time to our knowledge the presence of such a polysaccharide in L. edodes, a mushroom extensively investigated for its therapeutic effects.

CDP from C. dryophila and CDP-like from L. edodes and M. oreades were shown to inhibit NO production in macrophages activated by LPS and IFN{gamma}, suggesting a potential anti-inflammatory effect. Hara et al (1982)Go, Ukai et al (1983)Go and Ooi and Liu (1999)Go reported (1->3), (1->6) and other uncharacterized polysaccharides from Basidiomycete mushrooms displaying anti-inflammatory activity but did not demonstrate inhibition of NO production. In contrast lentinan, the most studied fungal (1->3)-, (1->6)-ß-D-glucan, has been reported to increase NO production in mouse macrophages (Mizuno 2000Go, Murata et al 2002Go). The type of glucosidic linkage (i.e. [1->3], [1->4] vs [1->3], [1->6]) may be partially responsible for the differential effect of CDP and CDP-like on NO production when compared to lentinan, although further structure/activity analysis is required to confirm this hypothesis.

Continuous elevated production of NO by macrophages may account for several disorders associated with chronic inflammatory diseases (Scuro et al 2004Go). Hence molecules, such as CDP and CDP-like, with the capability to decrease NO production by macrophages are of interest (Scarfi et al 1999Go, Aktan 2004Go).

Overall, results presented herein report for the first time the presence of rare (1->3)-, (1->4)-ß-D-glucans in different mushrooms, including L. edodes, that could find practical application as anti-inflammatory agents. Research in our laboratory will further characterize CDP to provide additional evidence of CDP anti-inflammatory activity using animal models and to elucidate its mechanism of action in the inflammatory response.


   ACKNOWLEDGMENTS
 
We thank Maurice Thibault for collecting the wild mushrooms, Dr Carole Martinez for her assistance and Dr Tyler Avis for critical review of the manuscript. This research was supported by Conseil de Recherche en Agriculture, Pêche et Alimentation du Québec (CORPAQ).


   FOOTNOTES
 
Accepted for publication December 2, 2005.

1 Corresponding author. E-mail: russell.tweddell{at}crh.ulaval.ca


   LITERATURE CITED
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 LITERATURE CITED
 
Ahn KS, Noh EJ, Zhao HL, Jung SH, Kang SS, Kim YS. 2005. Inhibition of inducible nitric oxide synthase and cyclooxygenase II by Platycodon grandiflorum saponins via suppression of nuclear factor-{kappa} B activation in RAW 264.7 cells. Life Sci 76:2315–2328.[CrossRef][Medline]

Aktan F. 2004. iNOS-mediated nitric oxide production and its regulation. Life Sci 75:639–653.[CrossRef][Medline]

Borchers AT, Stern JS, Hackman RM, Keen CL, Greshwin ME. 1999. Mushrooms, tumors and immunity. Proc Soc Exp Biol Med 221:281–293.[Abstract]

Cerning J, Renard CMGC, Thibault JF, Bouilanne C, Landon M, Desmazeaud M, Topisirovic L. 1994. Carbon source requirements for exopolysaccharide production by Lactobacillus casei CG11 and partial structure analysis of the polymer. Appl Environ Microbiol 60:3914–3919.[Abstract/Free Full Text]

Chihara G. 1992. Recent progress in immunopharmacology and therapeutic effects of polysaccharides. Develop Biol Stand 77:191–197.

Green LC, Wagner DA, Glogowski J, Skipper PL, Wishnok JS, Tannenbaum SR. 1982. Analysis of nitrate, nitrite and [15N] mitrate in biological fluids. Analyt Biochem 126:131–138.

Griffin DH. 1994. Fungal Physiology. New York: Wiley-Liss Inc.

Hara C, Kiho T, Tanaka T, Ukai S. 1982. Anti-inflammatory activity and conformational behavior of a branched (1 leads to 3)-beta-D-glucan from an alkaline extract of Dictyophora indusiata Fisch. Carbohydr Res 110:77–87.

International Union of Biochemistry and Molecular Biology. 1992. Enzyme Nomenclature. New York: Academic Press Inc.

Jong SC, Birmingham JM. 1993. Medicinal and therapeutic value of the shiitake mushrooms. Adv Appl Microbiol 39:153–184.[Medline]

Lee SH, Lee SY, Son DJ, Lee H, Yoo HS, Song S, Oh KW, Han DC, Kwon BM, Hong JT. 2005. Inhibitory effect of 2' hydroxycinnamaldehyde on nitric oxide production through inhibition of NF-{kappa} B activation in RAW 264.7 cells. Biochem Pharmacol 69:791–799.[CrossRef][Medline]

Liu F, Ooi VEC, Fung MC. 1999. Analysis of immunomodulating cytokine mRNAs in the mouse induced by mushroom polysaccharides. Life Sci 64:1005–1011.[CrossRef][Medline]

Marshall VM, Cowie EN, Moreton RS. 1995. Analysis and production of two exopolysaccharides from Lactococcus lactis subsp. cremoris LC330. J Dairy Res 62:621–628.

Mizuno M. 2000. Anti-tumor polysaccharides from mushrooms during storage. BioFactors 12:275–281.[Medline]

Murata Y, Shimamura T, Tagami T, Takatsuki F, Hamuro J. 2002. The skewing to Th1 induced by lentinan is directed through the distinctive cytokine production by macrophages with elevated intracellular glutathione content. Int Immunopharmacol 2:673–689.[CrossRef][Medline]

Netopilìk M. 2003. Problems connected with band-broadening in size-exclusion chromatography with dual detection. J Biochem Biophys Method 56:79–93.[CrossRef][Medline]

Ooi VEC, Liu F. 1999. A review of pharmacological activities of mushroom polysaccharides. Int J Medic Mush 1:195–206.

Pugh N, Ross SA, ElSohly MA, Pasco DS. 2001. Characterization of aloeride, a new high-molecular-weight polysaccharide from Aloe vera with potent immunostimulatory activity. J Ag Food Chem 49:1030–1034.[CrossRef]

Robyt JF. 1998. Essentials of Carbohydrate Chemistry. New York: Springer-Verlag Inc.

Ruiz-Herrera J. 1992. Fungal Cell Wall: structure, synthesis and assembly. Boca Raton Florida: CRC Press Inc.

Sakurai N. 2000. ß-glucohydrolase and ß-glucanohydrolase acting on 1,3;1,4-ß-glucan of cereal plants. In: Ohnishi M., ed. Glycoenzymes. Tokyo, Japan: Japan Scientific Societies Press. p 115–121.

Scarfi S, Giovine M, Gasparini A, Damonte G, Millo E, Pozzolini M, Benatti U. 1999. Modified peptide nucleic acids are internalized in mouse macrophages Raw 264.7 and inhibit inducible nitric oxide synthase. FEBS Lett 451:264–268.[CrossRef][Medline]

Schmid F, Stone BA, McDougall BM, Bacic A, Martin KL, Brownlee RTC, Chai E, Seviour RJ. 2001. Structure of epiglucan, a highly side-chain/branched (1->3;1->6)-ß-glucan from the micro fungus Epicoccum nigrum Ehrenb. ex Schlecht. Carbohydr Res 331:163–171.

Scuro LS, Simioni PU, Gabriel DL, Saviani EE, Modolo LV, Tamashiro WMSC, Salgado I. 2004. Suppression of nitric oxide production in mouse macrophages by soybean flavonoids accumulated in response to nitroprusside and fungal elicitation. BMC Biochem 5:5.[CrossRef][Medline]

Ukai S, Kiho T, Hara C, Kuruma I, Tanaka Y. 1983. Polysaccharides in fungi XIV. Anti-inflammatory effect of the polysaccharides from the fruit bodies of several fungi. J Pharmacobiodyn 6:983–990.[Medline]

Wasser SP, Weis AL. 1999. Therapeutic effects of substances occurring in higher Basidiomycetes mushrooms: a modern perspective. Crit Rev Immunol 19:65–96.[Medline]

———. 2002. Medicinal mushrooms as a source of antitumor and immunomodulating polysaccharides. Appl Microbiol Biotechnol 60:258–274.[CrossRef][Medline]

Yan J, Zong H, Shen A, Chen S, Yin X, Shen X, Liu W, Gu X, Gu J. 2003. The ß-(1-6)-branched ß-(1-3) glucohexaose and its analogues containing an {alpha}-(1–3)-linked bond have similar stimulatory effects on the mouse spleen as lentinan. Int Immunopharmacol 3:1861–1871.[CrossRef][Medline]

Yun JM, Kwon H, Hwang JK. 2003. In vitro anti-inflammatory activity of panduratin A isolated from Kaempferia pandurata in RAW 264.7 cells. Planta Med 69: 1102–1108.[CrossRef][Medline]





This Article
Right arrow Abstract Freely available
Right arrow Full Text (PDF)
Services
Right arrow Similar articles in this journal
Right arrow Alert me to new issues of the journal
Right arrow Download to citation manager
Right arrow reprints & permissions
Citing Articles
Right arrow Citing Articles via Google Scholar
Google Scholar
Right arrow Articles by Pacheco-Sanchez, M.
Right arrow Articles by Tweddell, R. J.
Right arrow Search for Related Content
PubMed
Right arrow Articles by Pacheco-Sanchez, M.
Right arrow Articles by Tweddell, R. J.
Agricola
Right arrow Articles by Pacheco-Sanchez, M.
Right arrow Articles by Tweddell, R. J.

HOME HELP FEEDBACK SUBSCRIPTIONS ARCHIVE SEARCH TABLE OF CONTENTS