Implications of cell wall immunocytochemical profiles on the structural and functional traits of root and stem galls induced by Eriosoma lanigerum on Malus domestica.

Alterations in cell wall composition imply in new structural and functional traits in gall developmental sites, even when the inducer is a sucking exophytophagous insect with strict feeding sites as the aphid associated to Malus domestica Borkh. This host plant is an economically important, fruit-bearing species, susceptible to gall induction by the sucking aphid Eriosoma lanigerum Hausmann, 1802. Herein, the immunocytochemical detection of arabinogalactan-proteins (AGPs), pectins, and hemicelluloses using monoclonal antibodies was performed in samples of non-galled roots and stems, and of root and stem galls on M. domestica. The dynamics of these cell wall components was discussed under the structural and functional traits of the galls proximal, median, and distal regions, according to the proximity of E. lanigerum colony feeding site. In the proximal region, the epitopes of AGPs and homogalacturonans (HGs) are related to cell growth and divisions, which result in the overproduction of parenchyma cells both in root and stem galls. In the proximal and median regions, the co-occurrence of HGs and arabinans in the cell walls of parenchyma and secondary tissues favors the nutrient flow and water-holding capacity, while the xylogalacturonans and hemicelluloses may function as additional carbohydrate resources to E. lanigerum. The immunocytochemical profile of the cell walls support the feeding activity of E. lanigerum mainly in the gall proximal region. The similarity of the cell wall components of the gall distal region and the non-galled portions, both in roots and stems, relates to the decrease of the cecidogenetic field the more distant the E. lanigerum colony is.

Molecular weight-dependent antitumor effects of prunes-derived type I arabinogalactan on human and murine triple wild-type melanomas.

The regulation of metastasis-related cellular aspects of two structurally similar AGIs from prunes tea infusion, with different molar masses, was studied in vitro against Triple Wild-Type metastatic melanoma (TWM) from murine and human origin. The higher molar mass AGI (AGI-78KDa) induced TWMs cells death and, in murine cell line, it decreased some metastasis-related cellular processes: invasiveness capacity, cell-extracellular matrix interaction, and colonies sizes. The lower molar mass AGI (AGI-12KDa) did not induce cell death but decreased TWMs proliferation rate and, in murine cell line, it decreased cell adhesion and colonies sizes. Both AGIs alter the clonogenic capacity of human cell line. In spite to understand why we saw so many differences between AGIs effects on murine and human cell lines we performed in silico analysis that demonstrated differential gene expression profiles between them. Complementary network topological predictions suggested that AGIs can modulate multiple pathways in a specie-dependent manner, which explain differential results obtained in vitro between cell lines. Our results pointed to therapeutic potential of AGIs from prunes tea against TWMs and showed that molecular weight of AGIs may influence their antitumor effects.

Water-soluble polysaccharides from Piper regnellii (Pariparoba) leaves: Structural characterization and antinociceptive and anti-inflammatory activities.

Piper regnellii is a plant popularly known as "Pariparoba" and it is widely used in folk medicine to treat pain, inflammation, among others. This work presents the extraction, purification and characterization of polysaccharides present in the plant leaves and evaluation of their anti-inflammatory and antinociceptive activities. From the crude aqueous extract of P. regnellii leaves, a polysaccharide fraction named PR30R, predominantly constituted of arabinose, galactose and galacturonic acid monosaccharide units, was obtained. Methylation and NMR analysis showed that the main polysaccharides of PR30R are a type II arabinogalactan, formed by a ß-D-Galp-(1 â†’ 3) main chain, substituted at O-6 by side chains of ß-D-Galp-(1 â†’ 6), which are substituted at O-3 by non-reducing α-L-Araf ends, and a homogalacturonan, formed by →4)-α-D-GalpA-(1→ units. Intraperitoneal administration of the crude polysaccharide fraction PRSF reduced significantly nociception induced by acetic acid in mice at the doses tested, and the PR30R fraction, derived from PRSF, presented antinociceptive and anti-inflammatory effects at a dose of 0.1096 mg/kg (PRSF ED50). These data support the use of the plant leaves in folk medicine as an herbal tea to treat pain and inflammation.

Investigation of the chemical structure and analgesic and anti-inflammatory properties of polysaccharides that constitute the dietary fibers of soursop (Annona muricata) fruit.

Soursop fruits are widely used in the folk medicine to treat a variety of health conditions. Once the chemical structure of dietary fibers from fruits is closely related to its biological functions in the human body, we aimed to explore structural features and biological activity of dietary fibers from soursop. Polysaccharides that constitute the soluble and insoluble fibers were extracted and further analyzed using monosaccharide composition, methylation, molecular weight determination and 13C NMR data. Soursop soluble fibers (SWa fraction) were characterized as having type II arabinogalactan and a highly methyl esterified homogalacturonan, while non-cellulosic insoluble fibers (SSKa fraction) were mainly composed by a pectic arabinan, a xylan-xyloglucan complex and a glucuronoxylan. The oral pre-treatment with SWa and SSKa promoted antinociception in mice writhing test, reducing the number of pain-like behaviors (in 84.2 % and 46.9 %, respectively, at 10 mg/kg) and peritoneal leucocyte migration (55.4 % and 59.1 %, at 10 mg/kg), effects possibly associated with the pectins present in fruit pulp extractions. SWa also significantly inhibited the plasmatic extravasation of Evans blue dye in 39.6 % at 10 mg/kg. This paper describes for the first time the structural features of soursop dietary fibers that may be of biological significance in future.

The Role of Arabinogalactan Type II Degradation in Plant-Microbe Interactions.

Arabinogalactans (AGs) are structural polysaccharides of the plant cell wall. A small proportion of the AGs are associated with hemicellulose and pectin. Furthermore, AGs are associated with proteins forming the so-called arabinogalactan proteins (AGPs), which can be found in the plant cell wall or attached through a glycosylphosphatidylinositol (GPI) anchor to the plasma membrane. AGPs are a family of highly glycosylated proteins grouped with cell wall proteins rich in hydroxyproline. These glycoproteins have important and diverse functions in plants, such as growth, cellular differentiation, signaling, and microbe-plant interactions, and several reports suggest that carbohydrate components are crucial for AGP functions. In beneficial plant-microbe interactions, AGPs attract symbiotic species of fungi or bacteria, promote the development of infectious structures and the colonization of root tips, and furthermore, these interactions can activate plant defense mechanisms. On the other hand, plants secrete and accumulate AGPs at infection sites, creating cross-links with pectin. As part of the plant cell wall degradation machinery, beneficial and pathogenic fungi and bacteria can produce the enzymes necessary for the complete depolymerization of AGs including endo-ß-(1,3), ß-(1,4) and ß-(1,6)-galactanases, ß-(1,3/1,6) galactanases, α-L-arabinofuranosidases, ß-L-arabinopyranosidases, and ß-D-glucuronidases. These hydrolytic enzymes are secreted during plant-pathogen interactions and could have implications for the function of AGPs. It has been proposed that AGPs could prevent infection by pathogenic microorganisms because their degradation products generated by hydrolytic enzymes of pathogens function as damage-associated molecular patterns (DAMPs) eliciting the plant defense response. In this review, we describe the structure and function of AGs and AGPs as components of the plant cell wall. Additionally, we describe the set of enzymes secreted by microorganisms to degrade AGs from AGPs and its possible implication for plant-microbe interactions.

Mucilages of cacti from Brazilian biodiversity: Extraction, physicochemical and technological properties.

Brazilian cacti have the high potential to be used as a source of mucilages with different technological functions that can be widely explored. The physicochemical, structural and technological properties of mucilages extracted from seven cladodes of cacti native from the semi-arid region of the Brazil were investigated. The mucilages exhibited high yields (8.9 to 21.54%), high contents of carbohydrates (39.77 to 87.68%), proteins (4.27 to 14.76%), and minerals, specifically Ca (2.90 to 15.65%). All species evaluated presented a diverse monosaccharide composition, mainly constituted of galactose and arabinose, suggesting the presence of arabinogalactans as the main polysaccharides. Both showed peaks and bands characteristic of polysaccharides with semi-crystalline structure (14.50-38.60%), due to the high concentration of calcium. From the results, mucilages functional properties showed good water and oil retention capacities and a high emulsification capacity (100%) with stability above 83%. Foaming capacity was above 58% with stability greater than 95%. This study highlights the potential of using cacti as a source of functional mucilages for use in food applications.

A polysaccharide fraction from Handroanthus albus (yellow ipê) leaves with antinociceptive and anti-inflammatory activities.

Handroanthus albus, commonly known as yellow ipê, is a native and widely distributed tree in Brazil. An aqueous soluble polysaccharide fraction (HASP) was obtained from its leaves, and monosaccharide composition, glycosidic linkage analysis by methylation and NMR spectroscopy indicated that HASP is mainly composed of a type II arabinogalactan, and suggested that other polysaccharides could also be present in a smaller proportion. HASP was able to promote antinociception in formalin-induced (second phase) and on glutamate-induced nociception tests, besides reducing the number of abdominal contortions induced by acetic acid in mice. Moreover, HASP reduced acetic acid-induced leukocyte infiltration in the peritoneal cavity and showed anti-edematogenic activity, decreasing mechanical allodynia and myeloperoxidase activity in the carrageenan-induced paw edema model. These results showed that the polysaccharide fraction HASP from H. albus leaves has interesting antinociceptive and anti-inflammatory activities.

Alkaline conditions better extract anti-inflammatory polysaccharides from winemaking by-products.

Winemaking generates large amounts of by-products, a well recognized source of phenolic compounds. However, less attention has been paid to the polysaccharide-rich fraction (PRF) and effects of fractionation techniques on its potential bioactivity. Therefore, PRFs from Syrah and Tempranillo winemaking by-products were extracted under aqueous (neutral pH conditions), acidic and alkaline conditions. PRFs were screened for their monosaccharide composition, uronic acid content, homogeneity and molecular weight. Anti-inflammatory activity of PRFs were evaluated on stimulated RAW 264.7 macrophages. PRF obtained in water and/or under acidic conditions showed heterogeneous profiles. As like as in the others, a heterogeneous and complex profile was detected in extracts procured under alkaline conditions. A high content of uronic acid was found in aqueous extracts, thus indicating the presence of pectin. Pectin and hemicellulose were present in PRFs procured under acidic conditions. Alkaline conditions rendered extracts containing a complex mixture of monosaccharides, mainly xylose. This latter PRF was the only one exhibiting anti-inflammatory potential (at 100 µg/mL) by reducing the release of TNF-α and activation of NF-κB in LPS-activated RAW 264.7 macrophages, with no effect on cell viability. Regardless of the grape variety, PRFs obtained under alkaline conditions were the best option to obtain bioactive polysaccharides with potential application as a source of anti-inflammatory compounds. A complex mixture of polymers may be responsible for the anti-inflammatory effects. Finally, according to results procured by NMR, it is possible to suggest that bioactive fractions are composed of a chain of α-L-Araf-(1 → 3) linked, ß-D-Xylp- (1 → 4), α-D-Glcp-(1 → 4) linked, α-D-GalpA-(1 → 4), α-D-Gal-(1 → 2) forming possible RG I and RG II and xylan chains.

A cell surface arabinogalactan-peptide influences root hair cell fate.

Root hairs (RHs) develop from specialized epidermal trichoblast cells, whereas epidermal cells that lack RHs are known as atrichoblasts. The mechanism controlling RH cell fate is only partially understood. RH cell fate is regulated by a transcription factor complex that promotes the expression of the homeodomain protein GLABRA 2 (GL2), which blocks RH development by inhibiting ROOT HAIR DEFECTIVE 6 (RHD6). Suppression of GL2 expression activates RHD6, a series of downstream TFs including ROOT HAIR DEFECTIVE 6 LIKE-4 (RSL4) and their target genes, and causes epidermal cells to develop into RHs. Brassinosteroids (BRs) influence RH cell fate. In the absence of BRs, phosphorylated BIN2 (a Type-II GSK3-like kinase) inhibits a protein complex that regulates GL2 expression. Perturbation of the arabinogalactan peptide (AGP21) in Arabidopsis thaliana triggers aberrant RH development, similar to that observed in plants with defective BR signaling. We reveal that an O-glycosylated AGP21 peptide, which is positively regulated by BZR1, a transcription factor activated by BR signaling, affects RH cell fate by altering GL2 expression in a BIN2-dependent manner. Changes in cell surface AGP disrupts BR responses and inhibits the downstream effect of BIN2 on the RH repressor GL2 in root epidermis.

A polysaccharide fraction from "ipê-roxo" (Handroanthus heptaphyllus) leaves with gastroprotective activity.

A polysaccharide fraction from Handroanthus heptaphyllus leaves was obtained with a simple and quick purification method. Methylation analysis and NMR spectroscopy indicated the presence of a complex polysaccharide fraction mainly constituted by a type II arabinogalactan. This is the first report in literature on structural elucidation of polysaccharides of species from genus Handroanthus. Oral and intraperitoneal administration of the polysaccharide fraction from Handroanthus heptaphyllus (HHSF) protected the gastric mucosa in an acute model of gastric lesion induced by ethanol, preserving gastric mucus. Furthermore, in the indomethacin model, HHSF reduced wounded area and inhibited mucus and GSH depletion. HHSF also accelerated gastric ulcer healing, accompanied by the maintenance of GSH levels. In addition, in an oxidative stress model with human epithelial cell line (Caco-2), HHSF was able to preserve GSH levels and was not toxic to cells. Collectively, these results showed that HHSF has an interesting antiulcerogenic activity and could constitute an interesting option for the treatment of gastric ulcer.

Pectins from cashew apple fruit (Anacardium occidentale): Extraction and chemical characterization.

The cashew tree (Anacardium occidentale) is a tropical evergreen tree largely cultivated in Brazil, which produces the cashew apple, a peduncle rich in carbohydrates and considered an industrial waste of the nut production. Until now, there were no data available about the chemical structure of cell wall polysaccharides found in cashew apple. In this work, its pectic polysaccharides have been characterized through monosaccharide composition, HPSEC, methylation and 13C and 1H/13C HSQC-DEPT-NMR analyses. Highly methyl esterified homogalacturonan with a DE of 76% mixed with arabinogalactan were found. This latter was purified and presented a highly branched type II arabinogalactan (AG II) and small amounts of a type I rhamnogalacturonan in which the AG II could be anchored. These findings about the chemical structure of cashew apple pectins could contribute to develop future nutritional, biotechnological and pharmacological uses for this industrial waste from the cashew nut production.

Chemical characterization of fructooligosaccharides, inulin and structurally diverse polysaccharides from chamomile tea.

Chamomile is one of most known species of medicinal plants. It has valuable pharmacological properties that produce positive effects in many therapeutical uses. Some of these properties are attributed to the presence of secondary metabolites but is already known that primary metabolites can also produce positive effects. In this study we elucidate the fine chemical structure of polysaccharides present in the infusion of chamomile flower chapters. After ethanolic precipitation, polysaccharides were obtained from the tea (fraction MRW, 3.2% yield), purified and characterized as an inulin type fructan, a highly methyl esterified and acetylated homogalacturonan (DE = 87% and DA = 19%), and a type II arabinogalactan. From ethanolic supernatant (20.2% yield), fructooligosaccharides (FOS) ranging from GF2 (m/z 543) to GF10 (m/z 1839) were detected. Inulin and FOS are well-established prebiotics, as well as the pectic polysaccharides. Thus, chamomile could be a source of structurally diverse dietary fibers with potential prebiotic, gastrointestinal and immunological functions.

Arabinogalactan from edible jambo fruit induces different responses on cytokine secretion by THP-1 macrophages in the absence and presence of proinflammatory stimulus.

Syzygium jambos is an Indo-Malaian found in many tropical countries and it is mainly composed of carbohydrates. Fraction PF-WSP and SF-WSP were obtained by aqueous extraction followed by Fehling's treatment. Monosaccharide analysis showed that fraction PF-WSP has a high content of uronic acids (90%) and fraction SF-WSP presented mainly galactose (39.1%) and arabinose (34.2%), as neutral sugars and 9% of galacturonic acid. The presence of type II arabinogalactan in SF-WSP was evidenced by methylation analysis and 13C/1H HSQC NMR experiments. Immunomodulating properties of SF-WSP was investigated. It decreases THP-1 macrophage viability at the highest concentration tested (200µg/mL). We then tested non-cytotoxic concentrations of SF-WSP on THP-1 cytokine production in the presence and absence of LPS. The results showed that SF-WSP increased TNF-α, IL-1ß and IL-10 secretion in a concentration-dependent manner as well as attenuated the inflammatory response induced by LPS at the highest concentration (100µg/mL). These results contribute to elucidate the effects of fruit pectic polysaccharides on immune cells.

New findings on green sweet pepper (Capsicum annum) pectins: Rhamnogalacturonan and type I and II arabinogalactans.

Polysaccharides were extracted from sweet pepper (Capsicum annum) with hot water and named ANW (9% yield). Starch was precipitated by freeze-thaw treatment, while pectic polysaccharides (8% yield) remained soluble and consisted of GalA (67.0%), Rha (1.6%), Ara (6.4%), Xyl (0.3%), Gal (6.7%) and Glc (4.4%). A highly methoxylated homogalacturonan (HG, degree of methylesterification of 85% and degree of acetylation of 5%), and type I and type II arabinogalactans (AG-I and AG-II) were observed in NMR analyses. These were fractionated with Fehling's solution to give HG (5.5% yield) and AG fractions (0.6% yield). AG-I and AG-II were further separated by ultrafiltration. AG-II (0.2% yield) consisted of Ara (17.1%), Gal (36.0%), Rha (5.6%) and GalA (12.0%), had a molecular weight of 5.3×104g/mol and methylation and 1H/13C HSQC-DEPT-NMR analyses showed that it was anchored in type I rhamnogalacturonan. This is the first study that reports the presence of AG-I and AG-II in sweet pepper fruits.

Characterization of a water soluble, hyperbranched arabinogalactan from yacon (Smallanthus sonchifolius) roots.

Yacon (Smallanthus sonchifolius Poepp. & Endl.) roots are largely grown in Andean countries and have attracted recent interest due to their antioxidant and prebiotic effects. Yacon is typically consumed as a fruit due to its sweet taste and juiciness. The macromolecular properties of an aqueous extract of yacon are investigated using asymmetric flow field-flow fractionation (AF4) coupled to UV, multiangle light scattering (MALS) and differential refractive index (dRI) detection. The method allows for determination of molar mass and size over the size distribution. Three major populations were found of which one strongly dominates in concentration. Through collection of fractions from AF4, carbohydrate composition and glycosidic linkage analysis for the dominating population was performed. The results show that the dominating population consists of a highly branched arabinogalactan (type 2) with a molar mass of approximately 1-2·105g/mol, a hydrodynamic radius of approximately 6-10nm and a relatively high apparent density (approx. 70-150kg/m3).

Extraction and characterization of pectins from primary cell walls of edible açaí (Euterpe oleraceae) berries, fruits of a monocotyledon palm.

Açaí berries (Euterpe oleracea) are greatly consumed in Brazil and exported to other countries as a nutritional supplement, due to health benefits attributed to its consumption. However, the complete chemical structure of bioactive polysaccharides was not fully elucidated yet. In this work, we characterize pectic polysaccharides from açaí berries through monosaccharide composition, HPSEC, methylation and 13C and 1H/13C HSQC-DEPT-NMR analyses. A highly methoxylated homogalacturonan with a DM of 88% and Mw of 22kDa together with small amounts of a mannoglucan were found. Moreover, a type II arabinogalactan (Mw=45kDa) containing a backbone with high portions of 6-O-linked and 3,6-O-linked Galp chains rather than 3-O-linked Galp was also isolated and structurally characterized. The type II arabinogalactan was found as a side chain of a type I rhamnogalacturonan. These findings contribute to correlate the fine chemical structure with the previously reported action of açaí polysaccharides on innate immune response. Moreover, from the taxonomic point of view, the results bring new information about polysaccharide composition of primary cell walls of palms (Arecaceae), that despite being commelinid monocots, have a distinct cell wall composition.

Polysaccharides from Arctium lappa L.: Chemical structure and biological activity.

The plant Arctium lappa L. is popularly used to relieve symptoms of inflammatory disorders. A crude polysaccharide fraction (SAA) resulting of aqueous extraction of A. lappa leaves showed a dose dependent anti-edematogenic activity on carrageenan-induced paw edema, which persisted for up to 48h. Sequential fractionation by ultrafiltration at 50kDa and 30kDa cut-off membranes yielded three fractions, namely RF50, RF30, and EF30. All these maintained the anti-edematogenic effect, but RF30 showed a more potent action, inhibiting 57% of the paw edema at a dose of 4.9mg/kg. The polysaccharide RF30 contained galacturonic acid, galactose, arabinose, rhamnose, glucose, and mannose in a 7:4:2:1:2:1 ratio and had a Mw of 91,000g/mol. Methylation analysis and NMR spectroscopy indicated that RF30 is mainly constituted by a type I rhamnogalacturonan branched by side chains of types I and II arabinogalactans, and arabinan.

A comparative study of mucilage and pulp polysaccharides from tamarillo fruit (Solanum betaceum Cav.).

A comparative study of mucilage (locular tissue) and pulp polysaccharides from ripe tamarillo fruits (Solanum betaceum Cav.) was carried out. After aqueous and alkaline extractions and various purification steps (freeze-thaw and α-amylase - EC 3.2.1.1 treatments, Fehling precipitation and ultrafiltration through 50 kDa cut-off membrane), the obtained fractions from mucilage were analyzed by sugar composition, HPSEC, and NMR spectroscopy analyses. The results showed that the mucilage of tamarillo contains a highly methoxylated homogalacturonans mixed with type I arabinogalactans, a linear (1 â†’ 5)-linked α-L-arabinan, and a linear (1 â†’ 4)-ß-D-xylan. A comparison with polysaccharides extracted from the pulp revealed that differences were observed in the yield and in the ratio of extracted polysaccharides. Moreover, structural differences between pulp and mucilage polysaccharides were also observed, such as in the length of side chains of the pectins, and in the degree of branching of the xylans.

Pectinas de plantas medicinais: características estruturais e atividades imunomoduladoras / Pectins of medicinal plants: structural characteristics and immunomodulatory activities

RESUMO As plantas medicinais apresentam várias propriedades terapêuticas, as quais estão relacionadas com a presença de compostos bioativos. Dentre os compostos, destacam-se as pectinas, que compreendem um grupo de polissacarídeos ácidos de relevante importância medicinal e nutracêutica. As pectinas são formadas por unidades de ácido galacturônico, unidas por ligação do tipo α-(1→4), sendo classificadas em homogalacturonanas e ramnogalacturonanas tipo I (RG-I) e tipo II (RG-II). Outros polissacarídeos constituídos por arabinose e/ou galactose têm sido isolados em associação com polissacarídeos pécticos, como as arabinogalactanas (AG) (tipo I e tipo II). As AG-II podem estar associadas a proteínas, denominadas de arabinogalactana-proteínas (AGPs). Inúmeros relatos demonstram que as pectinas, bem como as AG e AGPs, podem atuar como moduladores do sistema imunológico, sendo, por isso, consideradas modificadores da resposta biológica. A imunomodulação pode estar relacionada tanto com a atividade de macrófagos quanto com as vias do sistema complemento. Em geral, os polissacarídeos provocam um estímulo da atividade fagocitária; no aumento da produção de espécies reativas de oxigênio e da secreção de citocinas pró-inflamatórias. Em relação ao sistema complemento, os polissacarídeos podem modular tanto a via clássica como a via alternativa. A presente revisão tem como objetivo principal descrever os aspectos estruturais de pectinas e suas atividades biológicas relacionadas à modulação do sistema imune. Utilizando literatura específica, estão descritas informações de 29 espécies de plantas medicinais, que apresentam como constituintes pectinas, arabinogalactanas e/ou AGPs, correlacionando suas propriedades terapêuticas com as atividades biológicas associadas ao sistema imune. Na maioria dos casos descritos na literatura, é difícil determinar como as características estruturais específicas podem estar envolvidas na modulação da atividade de macrófagos. Porém, em relação à modulação da atividade do sistema complemento fica sugerido que a presença de estruturas tipo AG-II contribuiria mais significativamente para esta atividade. Entretanto, os possíveis mecanismos de modulação de pectinas, AGs e AGPs sobre a atividade de macrófagos e/ou sobre o sistema complemento ainda não estão totalmente esclarecidos, mesmo assim, estes polímeros podem ser considerados potenciais candidatos para estudos que visam ao desenvolvimento de novos agentes terapêuticos com propriedades moduladoras benéficas para o sistema imunológico.

ABSTRACT Medicinal plants have many therapeutic properties that are related to the presence of biologically active compounds. Pectins, a group of acid polysaccharides that have relevant medicinal and nutraceutical properties, are an example of such biological compounds. Pectins contain a main chain with galacturonic acid units that are α-(1→4) linked; they can be classified into homogalacturonans and type I and type II rhamnogalacturonans (RG-I and RG-II). Other polysaccharides containing arabinose, galactose, or both have been isolated in association with pectin-type polysaccharides are known as arabinogalactans (AGs, type I and type II). Arabinogalactan-proteins (AGPs) comprise AG-II associated with proteins. Several studies have reported that pectins, as well as AG and AGPs, can act as modulators of the immune system and can therefore be considered biological response modifiers. The immunomodulation is related to the activity of macrophages as on the complement system pathways. In general, polysaccharides cause stimulation of phagocytic activity, increase production of reactive oxygen species and secretion of pro-inflammatory cytokines. Polysaccharides can modulate the classical and alternative complement pathways. The aim of this review has to describe the structural aspects of pectins and their biological activities related to the modulation of the immune system. Using literature, we reported data of 29 medicinal plant species, which present as constituents pectins, arabinogalactan and/or AGPs, correlating their therapeutic properties with biological activities associated to the immune system. In most cases described in the literature, it is difficult to determine how the specific structural characteristics can be involved in modulation of macrophage activity. However, with respect to the modulation of the activity of the complement system is proposed that the presence of AG-II-type structures would contribute most significantly to this activity. The possible mechanisms of modulation of pectins, AGs and AGPs on macrophage activity and/or the complement system are not yet fully clear, even if, these polymers can be considered potential candidates for studies aimed at the development of new therapeutic agents with modulatory properties beneficial to the immune system.

Effect of the native polysaccharide of cashew-nut tree gum exudate on murine peritoneal macrophage modulatory activities.

The native polysaccharide of cashew-nut tree gum exudate (CNTG) and its arabinogalactan-protein component (CNTG-AGP) were tested by using immuno-stimulant and anti-inflammatory in vitro assays of murine peritoneal macrophage activities. In the assay for immuno-stimulant activity (without previous treatment with lipopolysaccharide; LPS), CNTG increased the production of interleukin (IL)-10 and both CNTG and CNTG-AGP decreased the concentrations of IL6. When the macrophages were incubated in the presence of LPS and CNTG a decrease in the levels of nitric oxide (NO(·)) and IFN-γ was observed. The results could explain the popular use of CNTG as an anti-inflammatory. In addition, CNTG is the main component of the cashew-nut tree gum exudate, which has been considered a versatile polymer with potential pharmaceutical and food industry applications. These data may contribute to the study of the immunomodulation activity of plant polysaccharides, as well as encourage future experiments in the field of cashew-nut tree gum exudate applications.