CHEMISTRY, PROPERTIES, AND PHYLOGENETIC IMPLICATIONS OF THE METHYLATED CARRAGEENANS FROM RED ALGAE OF THE GENUS ARESCHOUGIA (ARESCHOUGIACEAE, GIGARTINALES, RHODOPHYTA)

The three Australian-endemic species comprising the genus Areschougia have been examined to determine the structure of their nonfibrillar wall components. The polysaccharide extracted from the most widely distributed species, A. congesta (Turner) J. Agardh, was shown by compositional analyses, Fourier transform infrared (FTIR) spectroscopy, linkage analysis, and ¹³C-NMR spectroscopy to be a carrageenan composed predominantly of the repeating disaccharides 6'- O -meth- ylcarrabiose 2,4'-disulfate, carrabiose 2,4'-disulfate (the repeating unit of ι-carrageenan), 4',6'- O -(1-carboxyethylidene)carrabiose 2-sulfate, and 6'- O -methylcarrabiose 2-sulfate. The carrageenan also contained small amounts of 4-linked Gal p residues, some bearing methyl ether substitution at O-3 and some possibly bearing sulfate ester and/or glycosyl substitutions at O-3. The A. congesta carrageenan had unique rheological properties, its gels having some similarities to those of commercial ι-carrageenan but with the viscosity of commercial λ-carrageenan. Polysaccharides from A. ligulata Harvey ex J. Agardh and A. stuartii Harvey were shown by constituent sugar and FTIR analyses to be sulfated galactans rich in mono- O -methylgalactose. The carrageenan structures of Areschougia spp. were consistent with those of the genera Rhabdonia , Erythroclonium , and Austroclonium , the other genera constituting the family Areschougiaceae.     

Bioconversion of red seaweed galactans: a focus on bacterial agarases and carrageenases

Agars and carrageenans are 1,3-α-1,4-β-galactans from the cell walls of red algae, substituted by zero (agarose), one (κ-), two (ι-), or three (λ-carrageenan) sulfate groups per disaccharidic monomer. Agars, κ-, and ι-carrageenans auto-associate into crystalline fibers and are well known for their gelling properties, used in a variety of laboratory and industrial applications. These sulfated galactans constitute a crucial carbon source for a number of marine bacteria. These microorganisms secrete glycoside hydrolases specific for these polyanionic, insoluble polysaccharides, agarases and carrageenases. This article reviews the microorganisms involved in the degradation of agars and carrageenans, in their environmental and taxonomic diversity. We also present an overview on the biochemistry of the different families of galactanases. The structure–function relationships of the family GH16 β-agarases and κ-caraggeenases and of the family GH82 ι-carrageenases are discussed in more details. In particular, we examine how the active site topologies of these glycoside hydrolases influence their mode of action in heterogeneous phase. Finally, we discuss the next challenges in the basic and applied field of the galactans of red algae and of their related degrading microorganisms.     

Carrageenans from Cystocarpic and Sterile Plants of Chondrus Pinnulatus (Gigartinaceae, Rhodophyta) Collected from the Russian Pacific Coast

The chemical structure, gel properties and biological activity of the carrageenans isolated from cystocarpic and sterile plants of Chondrus pinnulatus were investigated. The total carrageenan content of the sterile plant was observed to be twice that of the cystocarpic plants. According to data obtained by ¹³C-NMR and FT IR, the gelling polysaccharides from cystocarpic and sterile plants of C. pinulatus have similar structures and were identified as κ/ι-carrageenans. The difference between these polysaccharides was in the ratio of the κ- and ι-segments, with a predominant content of κ-segments in cystocarpic plants (80%). Moreover, KCl-insoluble fractions possibly contain hetero-disperse μ/ν precursor: amounts of this in the polysaccharide from sterile plants were more than that extracted from the cystocarpic plants. The KCl-soluble fractions (non gelling) were λ-carrageenans with another carrageenan type that had a low amount of 3,6-anhydrogalactose. Carrageenans from cystocarpic stages showed good gelling properties, whereas those from sterile plants formed a very weak gel. Structural differences and molecular weight of carrageenans obviously determine the biological activity of the polysaccharides. Non gelling-carrageenans from both types of ιt C. pinnulatus plants showed high macrophage-phosphatase activity and κ/ι-carrageenan from cystocarpic plant possessed a potent anti-coagulant activity, which was extremely strong in a low concentration of 100 μg ml⁻¹.     

Chemical structure and gel properties of carrageenans from algae belonging to the Gigartinaceae and Tichocarpaceae, collected from the Russian Pacific coast

The chemical and gel characteristics of carrageenans isolated from the most abundant algal species growing on the Russian Pacific coast – Chondrus pinnulatus, C. armatus and Iridaea cornucopiae belonging to the Gigartinaceae and Tichocarpus crinitus from the Tichocarpaceae were investigated. The polysaccharides were identified by FTIR and NMR spectroscopy as predominantly κ-carrageenans with traces of ι-type (Gigartinaceae) and κ / β-type repeating structures (Tichocarpaceae) together with a small quantity of λ-carrageenan (10%). The chemical structure and the hydrodynamic properties play a determinant role on the rheology of these carrageenans. κ-Carrageenans from the Gigartinaceae displayed good gelling properties. The highest gel strength was obtained from C. pinnulatus (1232.7 Pa) at a 2.5% polymer concentration, while carrageenans from the Tichocarpaceae formed very weak gels (77.4 Pa) at the same concentration. Optimum gel characteristics were found with 1.0–2.0% KCI concentrations for kappa- carrageenans from Gigartinaceae and 0.75% for T. crinitus. The flow curves of λ-carrageenans solutions from the Gigartinaceae were similar, all between 20 and 65 °C, and characteristic of conformational disordered ‘random coil’ polysaccharides. Carrageenans from T. crinitus displayed the properties of ’random coil' only at high temperatures. This revised version was published online in June 2006 with corrections to the Cover Date.     

Carrageenans, sulphated polysaccharides of red seaweeds, differentially affect Arabidopsis thaliana resistance to Trichoplusia ni (cabbage looper)

Carrageenans are a collective family of linear, sulphated galactans found in a number of commercially important species of marine red alga. These polysaccharides are known to elicit defense responses in plant and animals and possess anti-viral properties. We investigated the effect of foliar application of ι-, κ- and λ-carrageenans (representing various levels of sulphation) on Arabidopsis thaliana in resistance to the generalist insect Trichoplusia ni (cabbage looper) which is known to cause serious economic losses in crop plants. Plants treated with ι- and κ-carrageenan showed reduced leaf damage, whereas those treated with λ- carrageenan were similar to that of the control. In a no-choice test, larval weight was reduced by more than 20% in ι- and κ- carrageenan treatments, but unaffected by λ-carrageenan. In multiple choice tests, carrageenan treated plants attracted fewer T. ni larvae by the fourth day following infestation as compared to the control. The application of carrageenans did not affect oviposition behaviour of T. ni. Growth of T. ni feeding on an artificial diet amended with carrageenans was not different from that fed with untreated control diet. ι-carrageenan induced the expression of defense genes; PR1, PDF1.2, and TI1, but κ- and λ-carrageenans did not. Besides PR1, PDF1.2, and TI1, the indole glucosinolate biosynthesis genes CYP79B2, CYP83B1 and glucosinolate hydrolysing QTL, ESM1 were up-regulated by ι-carrageenan treatment at 48 h post infestation. Gas chromatography-mass spectrometry analysis of carrageenan treated leaves showed increased concentrations of both isothiocyanates and nitriles. Taken together, these results show that carrageenans have differential effects on Arabidopsis resistance to T. ni and that the degree of sulphation of the polysaccharide chain may well mediate this effect.     

CARRAGEENANS BIOSYNTHESIZED BY CARPOSPOROPHYTES OF RED SEAWEEDS GIGARTINA SKOTTSBERGII (GIGARTINACEAE) AND GYMNOGONGRUS TORULOSUS (PHYLLOPHORACEAE)1

Carrageenans biosynthesized by gametophytic and tetrasporic plants of seaweeds belonging to the Gigartinaceae and Phyllophoraceae are different: gametophytes produce carrageenans of the kappa family, whereas lambda‐carrageenans are extracted from tetrasporophytes. For Gigartina skottsbergii Setchell and Gardner and Gymnogongrus torulosus Hooker et Harvey, mature cystocarps were isolated and carrageenans were extracted. Structural determination by methylation analysis, Fourier transform infrared spectroscopy, and ¹³C‐NMR spectroscopy showed that they were kappa/iota‐carrageenans. For the extract obtained from cystocarps of Gigartina skottsbergii with water at room temperature, the ratio kappa:iota was 1:0.30 and at 90° C was 1:0.43; significant amounts of precursors were also present. The extract obtained from cystocarps of Gymnogongrus torulosus at 90° C showed prevalence of iota‐carrageenans (ratio kappa:iota 1:1.21). These extracts are similar to the polysaccharides produced by gametophytes of these seaweeds. For Gigartina skottsbergii, it was possible to separate the pericarpic tissue from the carposporophyte. Thus, they were extracted separately, and the carrageenans isolated were studied as described before, obtaining similar conclusions. These results clearly show that whereas the carposporophytes are located inside the cystocarp, they produce carrageenans of the kappa family despite of being diploid cells.     

BETA/KAPPA-CARRAGEENANS AS EVIDENCE FOR CONTINUED SEPARATION OF THE FAMILIES DICRANEMATACEAE AND SARCODIACEAE (GIGARTINALES,RHODOPHYTA)1

The Dicranemataceae consists of five species in four genera of macroscopic red algae endemic to the southern half of Australia plus a single species from southern Japan. Investigations of the nonfibrillar wall components of five of the six species show that all are composed mainly of hybrid (or mixed) beta (β)/kappa(κ)-type carrageenans. Detailed studies of Tylotus obtusatus (Sonder) J. Agardh show that it produces the largest dry-weight percentage of β-carrageenan yet recorded. Monosaccharide composition, total sulfate content, sulfation pattern revealed by infrared and ¹³C-nuclear magnetic resonance spectroscopy and a positive specific optical rotation ([α]_D+ 54°) are indicative of a low-sulfate-containing carrageenan with gelling properties similar to those of agar and furcellaran. β-carrageenan is recorded in only five other red algal species belonging to relatively unrelated families, and we conclude that its uniform occurrence in the highly specialized family Dicranemataceae has phylogenetic significance. Chemical and anatomical examination of the genus Sarcodia, which produce lambda-type carrageenan in both its gametophytic and tetrasporophytic phases, suggests that, despite the recent proposal to incorporate the Dicranemataceae into the Sarcodiaceae, the two families should continue to be separated.     

Carrageenophytes of occidental Portuguese coast: 1-spectroscopic analysis in eight carrageenophytes from Buarcos bay

Infrared and Raman spectroscopic analysis of the carrageenan (alkaline extraction) in eight species (representing seven genera and four families) of Gigartinales, in different reproductive phases from Buarcos bay (Figueira da Foz, Portugal), were studied. Female gametophytes and non-fertile thalli samples of Chondrus crispus, Mastocarpus stellatus, Chondracanthus teedei var. lusitanicus, Gigartina pistillata and Chondracanthus acicularis present a kappa-carrageenan profile or varying degrees of a kappa-iota hybrid. The presence of kappa-iota hybrid carrageenan in C. teedei var. lusitanicus was confirmed by 13C NMR. The carrageenans extracted from Gymnogongrus crenulatus and Ahnfeltiopsis devoniensis are constituted mainly by iota-carrageenan but seasonal variations in the nature of carrageenans are present. lambda-Family carrageenans were found in tetrasporophytes of C. crispus, M. stellatus, C. teedei var. lusitanicus, C. acicularis and G. pistillata. Calliblepharis jubata presents carrageenans of iota-type in all reproductive stages.     

Chemotaxonomy of New Zealand red algae in the family Gigartinaceae (Rhodophyta) based on galactan structures from the tetrasporophyte life-stage

The identification of the polysaccharides from tetrasporophytic plants of nine endemic New Zealand species belonging to the Gigartinaceae, ‘Gigartina’ ancistroclada, ‘G.’ grandifida, Gigartina dilatata, G. divaricata, G. macrocarpa, G. marginifera, G. pachymenioides, G. sp. ‘Lindauer 164’ and Sarcothalia livida using infra-red spectroscopy in conjunction with constituent sugar and glycosyl linkage/substitution analysis is reported. All nine species contain galactans with structures consistent with λ-type carrageenans. Differences in the structures of the galactans in these and a further six previously studied species indicate chemotaxonomically distinct groupings that correspond to Sarcothalia, ‘Sarcothalia’ and Gigartina genera plus some outliers. These distinct, chemotaxonomic groupings are aligned to those determined by rbcL sequence analysis reported in the literature.     

Comparative study of carrageenans from reproductive and sterile forms of <Emphasis Type="Italic">Tichocarpus crinitus</Emphasis> (Gmel.) Rupr (Rhodophyta,Tichocarpaceae)

A comparative study of the structure and properties of the sulfated polysaccharides (carrageenans) isolated from the vegetative and reproductive forms of the red alga Tichocarpus crinitus was performed. The polysaccharides were separated into the gelling (KCl-insoluble) and non-gelling (KCl-soluble) fractions by precipitation with 4% KCl. The total content of polysaccharides extracted from the reproductive form of the alga was 1.8-fold more than that extracted from the vegetative form, and in the first case, the gelling polysaccharides mostly accumulated. The gelling polysaccharides from the vegetative form have the highest molecular weight (354 kD). According to the results of FT-IR and 13C-NMR spectroscopy, the gelling polysaccharide fractions from both forms are kappa/beta carrageenans. The differences concern the content of the kappa- and beta-disaccharide units and the presence of a small content of the sulfated disaccharide segments (precursors of the kappa-carrageenans) in the polysaccharide from the reproductive form of the alga. The non-gelling polysaccharide fractions from both forms of the plant are mixtures of sulfated galactans with a low content of 3,6-anhydrogalactose.     

樟科植物花粉形态研究

樟科植物为我国南方常绿阔叶林重要组成成分,我国约有20属420余种。过去对于樟科花粉形态的研究仅限于在光学显微镜下进行观察记载,以台湾学者王仁礼的研究最为全面,共记载了台湾产14属40余种的花粉形态。本文作者以国产樟科为主,应用扫描电镜及光学显微镜对该科花粉形态进行了更为全面系统的研究,共观察记载了22属150余种,对各属的花粉形态特征均进行了描述;首次报道了樟科花粉有小穿孔的存在,并依据花粉的形状、外壁纹饰及小穿孔存在与否将樟科花粉划分为7个类型,即:1．厚壳桂型(Cryptocarya,Neocimamomum);2．檬果樟型(Caryodaphnopsis);3．木姜子型(Litsea,Lindera,Neolitsea);4．月桂型(Laurus);5．鳄梨型(Persea,Machilus,Syndiclis,Nothaphoebe, Beilschemiedia,Phoebe,Alseodaphne,Octea);6．檫木型(Sassafras,Cinnamomum,Umbellularia,Actinodaphne,Dehaasia);7．无根藤型(Cassytha)。文中还讨论了樟科花粉的多样性、有关属之间的亲缘关系和分类问题以及同邻近的肉豆蔻科、莲叶桐科的关系。     

我国罂粟科(Papaveraceae)植物的花粉形态

本文用光学显微镜对我国16属60种5变种罂粟科植物花粉形态进行了系统地观察研究。其中对15属27种进行了扫描观察。本科花粉形态为多类型。3沟为原始的、基本的花粉类型。根据萌发孔的形状、数目和特征,将罂粟科的花粉分为2沟,3沟,4-6沟、散沟、散孔和无萌发孔6个花粉类型。根据孢粉学资料,本文对绿绒蒿属(Meconopsis Vig.)花粉粒和其它器官的进化趋势,罂粟科在进化上的亲缘关系和本科花粉萌发孔的演化趋向等问题进行了讨论。     

Polysaccharides enzymatic degradation in heterogeneous phase: example of agarases and carrageenases

Agars and carrageenans are sulphated galactans which assemble in the red algal cell wall as a dense network of semi-crystalline fibers. These polysaccharides are degraded in heterogeneous phase by bacterial enzymes, namely agarases and carageenases. Crystallographic as well as enzymologic investigations of the sulphated galactans/galactanases systems highlight that the properties of these catalysts are well adapted to the degradation of solid polyanionic substrates. Indeed, as for cellulases or amylases, they are able to depolymerize their respective substrates according to a processive mode of action. However, at the molecular level, they are distinguished by the ionic nature of the interactions involved which do not allow the direct transposition of the processivity models developed for the degradation of neutral polysaccharides.     

中国蕨科的数量分类研究

本文用聚类分析的方法,对中国蕨科23个种代表9个属做了数量分类研究,通过23OTUs×37个性状的原始数值矩阵标准化后,用欧氏距离系数计算相似性,采用WPGMA法进行聚类分析,以探索各属之间的系统位置和亲缘关系。其结果与我国植物学家经典分类处理基本上一致,并对粉背蕨属(Aleuritopteris)也做了修正,认为粉背蕨系(Ser.Farinosae)和银粉背蕨系(Sef.Argenteae)也可做为两个新属处理比较合理。另外对距离系数和结合线在类群划分申的作用进行了讨论和评价。     

Use of FTIR,FT-Raman and 13C-NMR spectroscopy for identification of some seaweed phycocolloids

Many seaweeds produce phycocolloids, stored in the cell wall. Members of the Rhodophyceae produce polysaccharides the main components of which are galactose (galactans)-agar and carrageenan. In addition, alginic acid is extracted from members of the Phaeophyceae. This is a binary polyuronide made up of mannuronic acid and guluronic acid. The wide uses of these phycocolloids are based on their gelling, viscosifying and emulsifying properties, which generate an increasing commercial and scientific interest. In this work, the FTIR and FT-RAMAN spectra of carrageenan and agar, obtained by alkaline extraction from different seaweeds (e.g. Mastocarpus stellatus, Chondrus crispus, Calliblepharis jubata, Chondracanthus acicularis, Chondracanthus teedei and Gracilaria gracilis), were recorded in order to identify the type of phycocolloid produced. The spectra of commercial carrageenan, alginic acid and agar samples (SIGMA and TAAB laboratories) were used as references. Special emphasis was given to the 500-1500 cm(-1) region, which presents several vibrational modes, sensitive to the type of polysaccharide and to the type of glycosidic linkage. The FT-Raman spectra present a higher resolution than FTIR spectra, this allowing the identification of a larger number of characteristic bands. In some cases, phycocolloids can be identified by FT-Raman spectroscopy alone.     

中国五加科花粉形态研究

本文对我国五加科(Araliaceae)植物21个属、34种、2变种的花粉形态进行了详细的观察研究, 每一个属均选择1至2个代表种进行电镜扫描。从花粉学方面进一步探讨了各属间的亲缘关系, 分类位置、以及分属的界限等问题。     

我国瑞香科(Thymelaeaceae)植物的花粉形态

本文对我国8属22种3变种的瑞香科植物花粉形态进行了观察。本科花粉具散孔,花粉形态属于单类型。根据纹饰特点,本文将瑞香科花粉分为具瘤状纹饰和具网状一巴豆型图案两个类型。另外,还对巴豆型图案和本科在被子植物中的系统位置等问题进行了讨论。     

海南岛尖峰岭灵芝科的种类组成及其生态分布

报道海南岛尖峰岭的灵芝科真菌４９种,分属于２个属,３个亚属,２个组。尖峰岭地区的灵芝在不同的热带植被类型中,其灵芝种类的组成不同。就种类和数量而言施雨林和热山地雨林多,热带半落叶季雨林次之,山顶苔藓矮林少。     

中国马鞭草科植物的花粉形态

本文对我国18属91种3变种马鞭草科(Verbenaceae)植物的花粉形态进行了详细地观察研究。其中对15属27种进行了扫描电镜观察。本科为花粉多类型的科之一。根据萌发孔的数目、特征和纹饰特点,本文将马鞭草科分为9个花粉类型:1.具疣3沟花粉类型;2.具刺3沟花粉类型;3.具细网8沟花粉类型;4.具模糊颗粒-细网3沟花粉类型;5.具网3沟花粉类型;6.具散沟(6-8条)花粉类型;7.具3孔花粉类型;3.具细网3沟花粉类型,4.具模糊颗粒一细网3沟花粉类型,5.具网3沟花粉类型,6.具散沟(6-8条)花粉类型,7.具3孔花粉类型,8.具3孔沟花粉类型,9.具3(4)孔沟花粉类型。     

中国蟹蛛科（蜘蛛目）新种记述

记述我国蟹蛛科花蟹蛛属６新种和蟹蛛属２新种,共计８新种,金林花蟹蛛Ｘｙｓｔｉｃｕｓｊｉｎｌｉｎ,玛纳斯花蟹蛛Ｘ．ｍａｎａｓ,林芝花蟹蛛Ｘ．ｎｙｉｎｇｃｈｉｅｎｓｉｓ,拟斑花蟹蛛Ｘ．ｐａｒａｐｕｎｃｔａｔｕｓ,似旋花蟹蛛Ｘ．ｔｏｒｓｉｖｏｉｄｅｓ,吴氏花蟹蛛Ｘ．ｗｕａｅ,广西蟹蛛Ｔｈｏｍｉｓｕｓｇｕａｎｇｘｉｃｕｓ和胡氏蟹蛛Ｔ．ｈｕｉ。