Polysaccharides of Calcareous Algae and Their Effect on the Calcification Process

The composition and structure of polysaccharides from several groups of calcareous algae (including calcareous cyanobacteria), which differ in the calcification mode (extracellular, cell wall, or intracellular), are reviewed. Two families of marine algae, Corallinaceae (Rhodophyta) and Coccolithophoraceae (Prymnesiophyta = Haptophyta), are considered in detail; they exhibit the cell wall and intracellular calcification modes, respectively, and synthesize unusual polysaccharides that seem to directly participate in the calcification process.     

Nutrient-limited productivity of calcareous versus fleshy macroalgae in a eutrophic,carbonate-rich tropical marine environment

The results of a study of nutrient enrichment with nitrogen (N) and phosphorus (P) on productivity and calcification of fleshy and calcareous algae are reported in this study. Plants were collected from a nearshore eutrophic site in the Florida Keys (USA) and experimentally pulsed during the night with combinations of N and P. After several days of pulsing (7–10 days), net productivity, calcification, and alkaline phosphatase activity (APA), were measured. Productivity of fleshy algae were frequently enhanced by N, P, and N+P, during both summer and winter. Phosphorus limited the productivity of Hydroclathrus clathratus during winter and Ulva spp. during summer, whereas nitrogen limited the productivity of Laurencia intricata during both seasons. During summer, Dictyota cervicornis productivity was not enhanced by any nutrient enrichment. Nitrogen limited the productivity of the three calcareous species Penicillus capitatus, Penicillus dumetosus and Halimeda opuntia during winter and that of H. opuntia during summer. Neither N nor P enrichment increased calcification of calcareous species, and P enrichment greatly inhibited calcification of P. dumetosus during winter. Nutrient enrichment enhanced the productivity of the fleshy species to a greater extent than that of calcareous algae. The seawater DIN:SRP molar ratio was low at our eutrophic study site (molar ratio average of 3:1 during winter and 9:1 during summer) compared to more oligotrophic sites in the Florida Keys, suggesting that in carbonate-rich environments, eutrophication shifts nutrient regulation of productivity from P to N. APA activities of fleshy macroalage were higher than calcareous algae, and rates of all macro algae were 2- to 7-fold higher in summer compared to winter. Productivity was also about 3-fold higher in fleshy compared to calcareous species and about 2-fold higher in summer compared to winter. These results suggest that nutrient enrichment enhances productivity of fleshy algae to a greater extent than that of calcareous algae. Thus, overgrowth of calcareous algae by more opportunistic fleshy forms could reduce carbonate accretion in tropical coastlines experiencing increased eutrophication.     

Seagrass meadows mixed with calcareous algae have higher plant productivity and sedimentary blue carbon storage

Seagrass meadows capture and store large amounts of carbon in the sediment beneath, thereby serving as efficient sinks of atmospheric CO₂. Carbon sequestration levels may however differ greatly among meadows depending on, among other factors, the plant community composition. Tropical seagrass meadows are often intermixed with macroalgae, many of which are calcareous, which may compete with seagrass for nutrients, light, and space. While the photosynthetic CO₂ uptake by both seagrasses and calcareous algae may increase the overall calcification in the system (by increasing the calcium carbonate saturation state, Ω), the calcification process of calcareous algae may lead to a release of CO₂, thereby affecting both productivity and calcification, and eventually also the meadows’ carbon storage. This study estimated how plant productivity, CaCO₃ production, and sediment carbon levels were affected by plant community composition (seagrass and calcareous algae) in a tropical seagrass‐dominated embayment (Zanzibar, Tanzania). Overall, the patterns of variability in productivity differed between the plant types, with net areal biomass productivity being highest in meadows containing both seagrass and calcareous algae. Low and moderate densities of calcareous algae enhanced seagrass biomass growth, while the presence of seagrass reduced the productivity of calcareous algae but increased their CaCO₃ content. Sedimentary carbon levels were highest when seagrasses were mixed with low or moderate cover of calcareous algae. The findings show that plant community composition can be an important driver for ecosystem productivity and blue carbon sequestration.     

钙质红藻化石的主要类群特征及演化

钙质红藻是指可以发生生物钙化作用在其细胞壁上沉淀碳酸钙的红藻。钙质红藻可以保存为化石,是红藻古生物研究中的重要类群,具有重要的生态意义,但以往的研究对钙质红藻类群的系统分类及地史分布缺乏清晰认识。本文详细综述了钙质红藻化石的系统分类,归属于红藻门(Rhodophyta)红藻纲(Rhodophyceae)的4个目7个科,分别为珊瑚藻亚纲(Corallinophycidae)珊瑚藻目(Corallinales)的珊瑚藻科(Corallinaceae)、石叶藻科(Lithophyllaceae)、宽珊藻科(Mastophoraceae)和管孔藻科(Solenoporaceae),混石藻目(Hapalidiales)的混石藻科(Hapalidiaceae),孢石藻目(Sporolithales)的孢石藻科(Sporolithaceae)以及真红藻亚纲(Florideophycidae)耳壳藻目(Peyssonneliales)的耳壳藻科(Peyssonneliaceae)。最早的钙质红藻为管孔藻科,出现于中奥陶世,于中新世灭绝。珊瑚藻科最早出现于晚志留世并于白垩纪辐射演化至今,其他科均于白垩纪...     

Rates of Turnover and Net Accretion of Calcium and the Role of Calcium Binding Polysaccharides During Calcification in the Calcareous Alga Halimeda opuntia (L.)

Calcification in the calcareous alga Halimeda opuntia involves Calcium exchange with the seawater. Calcium influx and-outflux is nearly the same and approximately 300 μg Ca/g alga/min. Calcium net accretion is 3.6 ± 1.1 μg Ca/g alga/min. Calcium outflux alga-seawater is higher in darkness than in light. Calcium exchange in live algae is 3–6 times faster than in dead algae. Calcium is lost from the seawater at two distinctly different rates and is taken up by watersoluble polysaccharides in the algal mucilage and by the skeleton. These two Calcium pools represent 0.1 and 99.9% of the algal Calcium respectively. Between 0.8–2.0% of the skeletal Calcium is exchangeable. The analysis of the ⁴⁵Ca specific activity curves suggests that there are more than 2 Calcium pools in the alga. The polysaccharides constitute a small but rapidly exchanging Calcium pool which communicates with the seawater and at least 2 other pools. The Calcium binding polysaccharides are involved in the Calcium metabolism of the alga and may play a role in algal calcification.     

MULTIMINERAL CALCAREOUS DEPOSITS IN THE MARINE ALGA ACETABULARIA ACETABULUM (CHLOROPHYTA; DASYCLADACEAE)

The ultrastructure and mineralogy of cultured and wild specimens of the green algae Acetabularia acetabulum were studied. Sites of calcification were clarified using TEM and SEM. Minerology was determined via electron diffraction, energy-dispersive x-ray microanalysis, and Fourier transform infrared spectral analyses of the calcareous deposits. These deposits were found covering the surface and within the cell wall of the cultured juveniles and adult wild specimens of A. acetabulum. Deposits within and directly over the wall were amorphous calcium carbonate (ACC) in granular form. In-wall and outer cell wall ACC appear to develop independently from each other. The formation of the in-wall ACC may be mediated by the wall structure and/or chemistry, whereas mucin may be the factor mediating formation of outer cell wall ACC. In contrast to the stable in-wall ACC, outer cell wall ACC could be transitory, transforming into aragonite. In cultured cells, a small amount of monohydrocalcite crystals and calcite were found. It is possible that these were transformation products of ACC. In wild specimens, whewellite (calcium oxalate monohydrate) was present inside and outside of the cap wall. It is unclear if whewellite is present within the cytoplasm. Local and temporal differences in the microenvironments for the multiphase mineralization are suggested.     

西伯利亚地台早寒武世钙藻的发育特征

西伯利亚地台的新元古代到早寒武世间的钙藻化石特别丰富,为研究钙藻化石的发育历史提供了模式。在几次演化事件背景下记录了演化的两次转折,其中一个是藻类的普遍钙化,还有一个是Botomian晚期到Toyonian早期之间在西伯利亚地台缺失钙藻化石记录,而在其它地区包括西伯利亚地台的边缘(Altay—Sayan地区),钙藻的丰度却达到了最大。     

Chemical and enzymatic fractionation of cell walls from Fucales: insights into the structure of the extracellular matrix of brown algae

Background and Aims

Brown algae are photosynthetic multicellular marine organisms evolutionarily distant from land plants, with a distinctive cell wall. They feature carbohydrates shared with plants (cellulose), animals (fucose-containing sulfated polysaccharides, FCSPs) or bacteria (alginates). How these components are organized into a three-dimensional extracellular matrix (ECM) still remains unclear. Recent molecular analysis of the corresponding biosynthetic routes points toward a complex evolutionary history that shaped the ECM structure in brown algae.

Methods

Exhaustive sequential extractions and composition analyses of cell wall material from various brown algae of the order Fucales were performed. Dedicated enzymatic degradations were used to release and identify cell wall partners. This approach was complemented by systematic chromatographic analysis to study polymer interlinks further. An additional structural assessment of the sulfated fucan extracted from Himanthalia elongata was made.

Key Results

The data indicate that FCSPs are tightly associated with proteins and cellulose within the walls. Alginates are associated with most phenolic compounds. The sulfated fucans from H. elongata were shown to have a regular α-(1→3) backbone structure, while an alternating α-(1→3), (1→4) structure has been described in some brown algae from the order Fucales.

Conclusions

The data provide a global snapshot of the cell wall architecture in brown algae, and contribute to the understanding of the structure–function relationships of the main cell wall components. Enzymatic cross-linking of alginates by phenols may regulate the strengthening of the wall, and sulfated polysaccharides may play a key role in the adaptation to osmotic stress. The emergence and evolution of ECM components is further discussed in relation to the evolution of multicellularity in brown algae.     

Skeletal ultrastructure of the calcified red alga Galaxaura oblongata,Hainan Island,China

Aragonite calcification in the red alga Galaxaura oblongata (Ellis et Solander) Lamouroux (Chaetangiaceae, Nemaliales) from southern China, occurs at three sites in the cortex: (1) intercellular spaces (ICS); (2) the cell wall; and (3) the inner surface of the cortex. The ICS is the principal site of calcification and contains aragonite mainly in the form of equidimensional granules ∼1–2 μm in size, although needle-like crystals up to 3 μm in length are also present locally. Granules range from densely to loosely packed. A dense and even layer peripheral to the ICS is interpreted to represent calcification of the cell wall. Coarser radial clumps of elongate crystals, forming a thin discontinuous layer on the inner surface of the cortex, resemble a cement fringe facing into the medulla, but form during the life of the alga. Calcification in these specimens shows similarities, and differences, to both coralline red algae, in which calcification is limited to the cell wall, and halimedacean green algae in which it is limited to the ICS. Some members of the Palaeozoic family Moniliporellaceae show similarities in their skeletal organization to Galaxaura.     

Characterization of Laurencia arbuscula spore mucilage and cell walls with stains and FITC-labelled lectins

In most red algae, spores are liberated without a cell wall, within a sheath of mucilage that is responsible for its primary attachment. Utilizing fluorescent-labelled lectins, we identified carbohydrate residues and their location in the mucilage and cell walls of spores of Laurencia arbuscula. Cell wall formation and mucilage composition were studied with Calcofluor, Toluidine Blue (AT-O), Alcian Blue (AB) and periodic acid-Schiff (PAS). In the mucilage, we identified α-d-mannose, α-d-glucose, N-acetyl-glucosamine, N-acetyl-galactosamine and β-d-galactose. All sugar residues were found in the cell wall, in the spore body rather than in the rhizoid, which suggests that the residues may be related to initial substrate adhesion. A cell wall is produced soon after the spore's attachment, beginning with a deposition of cellulose around the spore, as indicated by Calcofluor. A polarization of the cell wall triggers the process of germination. The cell-wall matrix was positive to AB and metachromatic to AT-O, indicating acidic polysaccharides, while neutral polysaccharides were positive to PAS.     

Monoclonal Antibodies Directed to Fucoidan Preparations from Brown Algae

Cell walls of the brown algae contain a diverse range of polysaccharides with useful bioactivities. The precise structures of the sulfated fucan/fucoidan group of polysaccharides and their roles in generating cell wall architectures and cell properties are not known in detail. Four rat monoclonal antibodies, BAM1 to BAM4, directed to sulfated fucan preparations, have been generated and used to dissect the heterogeneity of brown algal cell wall polysaccharides. BAM1 and BAM4, respectively, bind to a non-sulfated epitope and a sulfated epitope present in the sulfated fucan preparations. BAM2 and BAM3 identified additional distinct epitopes present in the fucoidan preparations. All four epitopes, not yet fully characterised, occur widely within the major brown algal taxonomic groups and show divergent distribution patterns in tissues. The analysis of cell wall extractions and fluorescence imaging reveal differences in the occurrence of the BAM1 to BAM4 epitopes in various tissues of Fucus vesiculosus. In Ectocarpus subulatus, a species closely related to the brown algal model Ectocarpus siliculosus, the BAM4 sulfated epitope was modulated in relation to salinity levels. This new set of monoclonal antibodies will be useful for the dissection of the highly complex and yet poorly resolved sulfated polysaccharides in the brown algae in relation to their ecological and economic significance.     

Isolation of a Unique Marine Bacterium Capable of Growth on a Wide Variety of Polysaccharides from Macroalgae

A unique marine bacterium has been isolated which can be cultured on a variety of polysaccharides and cell wall preparations from red and brown algae.     

Dasyclads, cyclocrinitids and receptaculitids: comparative morphology and paleoecology

The cyclocrinitids are an extinct tribe of dasycladacean green algae. They were anatomically very similar to certain Recent dasyclads, even at early growth stages. The morphology and preservation of cyclocrinitids strongly suggest that they had a siphonous cellular organization with extracellular, aragonitic calcification; these features are characteristic of living dasyclads. The light surficial calcification of cyclocrinitids and other dasyclads had important paleoecological effects. It restricted them to low-energy waters, as it provided relatively little structural support. It also confined them to warm, tropical waters; they are good paleoequatorial indicators. The decline of these algae during the late Ordovician and early Silurian may therefore reflect the simultaneous cooling and glaciation. Receptaculitids are entirely unrelated organisms. Their meroms have several distinctive features; they are not homologous to the lateral branches of cyclocrinitids or dasyclads. Receptaculitid calcification was extensive and their thalli were apparently quite sturdy; they often occurred in reefs. Receptaculitids also lived in high-latitude, cold-water environments. Thus, they were ecologically unlike any calcareous green algae, and cannot be used as paleoequatorial indicators. Receptaculitids remain problematical, although the arrangement of meroms suggests plant affinities. □ Calcareous algae, Problematica, Dasycladales, Cyclocriniteae, Receptaculitales, morphology, classification, paleoecology, paleogeography .     

Spore Adhesion and Cell Wall Formation in Gelidium Floridanum (Rhodophyta, Gelidiales)

The attachment of spores to a substratum is essential for their germination and, therefore, to the completion of the life cycle of the red algae. In most red algae, spores are liberated without a cell wall, within a sheath of mucilage which is responsible for their primary attachment. Utilizing fluorescent-labeled lectins, we identified carbohydrate residues and their locations in the mucilage and cell walls of spores of Gelidium floridanum. Cell wall formation and mucilage composition were studied with calcofluor, toluidine blue (AT-O), alcian blue (AB) and periodic acid-Schiff (PAS). In the mucilage we identified α-D mannose, α-D glucose, β-D-galactose, N-acetyl-glucosamine and N-acetyl-galactosamine. The first two sugar residues were not found in the cell wall of the germ tube but they were present on the rhizoid’s cell wall indicating their importance to substrate adhesion. A cell wall is produced soon after the spore’s attachment, beginning with a polar deposition of cellulose and its gradual spread around the spore as indicated by calcofluor. The cell wall matrix was positive to AB and metachromatic to AT-O, indicating acidic polysaccharides, while cellulose microfibrills were positive to PAS. A polar disorganization of the cell wall triggers the process of germination. As spores are the natural form of propagation of Gelidium, the understanding of the mechanisms of spore attachment may contribute to the cultivation of this valuable seaweed.     

Structure and functional properties of ulvan, a polysaccharide from green seaweeds

With today's interest in novel renewable chemicals and polymers, the underexploited marine green algae belonging to species of Ulva and Entermorpha stimulated interest as sources of polysaccharides with innovative structure and functional properties. These algae are common on all seashores and can produce in time an important amount of biomass in nutrient-enriched waters. The major water-soluble polysaccharide, ulvan, extracted from the cell wall represents about 8-29% of the algae dry weight. The original physicochemical, rheological, and biological properties recently unraveled for this complex sulfated aldobiouronan open the way for novel potential applications.     

Stable chloroplast transformation of the unicellular red alga Porphyridium species

Red algae are extremely attractive for biotechnology because they synthesize accessory photosynthetic pigments (phycobilins and carotenoids), unsaturated fatty acids, and unique cell wall sulfated polysaccharides. We report a high-efficiency chloroplast transformation system for the unicellular red microalga Porphyridium sp. This is the first genetic transformation system for Rhodophytes and is based on use of a mutant form of the gene encoding acetohydroxyacid synthase [AHAS(W492S)] as a dominant selectable marker. AHAS is the target enzyme of the herbicide sulfometuron methyl, which effectively inhibits growth of bacteria, fungi, plants, and algae. Biolistic transformation of synchronized Porphyridium sp. cells with the mutant AHAS(W492S) gene that confers herbicide resistance gave a high frequency of sulfomethuron methyl-resistant colonies. The mutant AHAS gene integrated into the chloroplast genome by homologous recombination. This system paves the way for expression of foreign genes in red algae and has important biotechnological implications.     

Characterization of the cell-wall polysaccharides of Arabidopsis thaliana leaves.

The cell-wall polysaccharides of Arabidopsis thaliana leaves have been isolated, purified, and characterized. The primary cell walls of all higher plants that have been studied contain cellulose, the three pectic polysaccharides homogalacturonan, rhamnogalacturonan I and rhamnogalacturonan II, the two hemicelluloses xyloglucan and glucuronoarabinoxylan, and structural glycoproteins. The cell walls of Arabidopsis leaves contain each of these components and no others that we could detect, and these cell walls are remarkable in that they are particularly rich in phosphate buffer-soluble polysaccharides (34% of the wall). The pectic polysaccharides of the purified cell walls consist of rhamnogalacturonan I (11%), rhamnogalacturonon II (8%), and homogalacturonan (23%). Xyloglucan (XG) accounts for 20% of the wall, and the oligosaccharide fragments generated from XG by endoglucanase consist of the typical subunits of other higher plant XGs. Glucuronoarabinoxylan (4%), cellulose (14%) and protein (14%) account for the remainder of the wall. Except for the phosphate buffer-soluble pectic polysaccharides, the polysaccharides of Arabidopsis leaf cell walls occur in proportions similar to those of other plants. The structure of the Arabidopsis cell-wall polysaccharides are typical of those of many other plants.     

A dye binding assay for the quantification of soluble and cell-bound acidic polysaccharides produced by red algae

Polysaccharides produced in technical scale from red algae serve in a variety of industrial applications. Ruthenium red (RR) is a cytochemical stain which has been used to visualize these mostly acidic polysaccharides in light microscopy. The binding of RR to algal polysaccharides is highly dependent on the ionic strength of the surrounding medium. The dye is firmly bound at low ionic strength, but is released at increasing salt concentrations. These properties were exploited to develop a new method to quantify the amount of cell wall material synthesized by algal cells under different physiological conditions. Soluble or extracted polysaccharides are quantified by a dot-blot procedure with subsequent image analysis, while cell-bound polysaccharides are determined in situ by a dye-binding assay followed by photometry. In comparison to the current methods of quantification, the new assay is quick, reliable, and avoids extensive use of hazardous chemicals.     

Marine macroalgae as foods for fishes: an evaluation of potential food quality

Synopsis A revitalized view of feeding by herbivorous marine fishes is sought through two questions. First, What characteristics of major taxa of algae identify them as predictably high or low quality foods? Second, are marine algae valuable foods for fishes which do not mechanically disrupt cell walls and do not harbor specialized enzymes or microbes capable of lysing cell walls? Energy, ash and nutrient content of 16 species of marine algae were employed to assess food quality of fleshy red, green, brown and calcareous red algae. On the basis of ash, calories, total protein and total lipid content, fleshy algae should be superior to calcareous algae as foods for fishes; in addition, green algae should be superior to brown algae and brown algae superior to red algae. When the probable digestibility of storage and extracellular carbohydrates is considered, green and red algae are predicted superior to brown algae as food. Two species of damselfishes (Pomacentridae) from the Gulf of California,Eupomacentrus rectifraenum andMicrospathodon dorsalis, eat red and green algae and ignore brown and calcareous algae. They feed, therefore, in a fashion consistent with predictions based only on algal chemistry. These fishes absorb at least 20–24% of the biomass, 57–67% of the protein, 46–56% of the lipid and 37–44% of the carbohydrate contained in algae eaten in the wild. Since these damselfishes do not masticate their food, it appears that herbivorous fishes can digest major fractions of algal nutrients without mechanical destruction of algal cells.