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Seaweeds along KwaZulu-Natal Coast of South Africa--4: Elemental uptake by edible seaweed Caulerpa racemosa (sea grapes) and the arsenic speciation.

The elemental uptake by edible seaweed Caulerpa racemosa (Sea grapes), a marine macroalgae (chlorophyta, green alga) grown richly along KwaZulu-Natal coastline. The total concentrations of seven elements, namely Mn, Fe, As, B, Ti, Zn and Hg in Caulerpa racemosa were monitored for a one-year cycle (June 2002 to May 2003) at four selected sampling sites spread over 150 km wide from North to South. The C. racemosa possess high arsenic, boron and titanium accumulating ability, but low iron uptake. A typical C. racemosa sample at Treasure Beach in the vicinity of Durban Metropolis in autumn contained Mn (5.2+/-ppm), Fe (0.21+/-0.01 ppm), As (8.5+/-0. ppm), B (1090+/-ppb), Ti (159+/-ppb), Zn (3.8+/-0.1 ppb), and Hg (189+/-ppb). The general trend found at all sites was high elemental concentrations in winter and a decrease in concentrations from winter to spring and summer. C. racemosa recorded highest mercury levels (>205 ppb) during the summer season at the Zinkwasi site. The arsenic speciation in four abundant seaweeds from the beaches of Indian Ocean in the KwaZulu-Natal coast is investigated. The speciation of arsenic in two rhodophyta seaweeds, Plocamium corallorhiza and Gelidium abbottiorum, and two chlorophyta seaweeds, Ulva lactuca and Caulerpa racemosa at four sampling sites during the summer of 2003, is elucidated. Caulerpa racemosa had highest total arsenic (in ppb) reaching (8850+/-200) at Zinkwasi, and the concentrations of the other arsenic species analysed are As (III) (194+/-10), As(V) (568+/-27), methylarsonic acid (494+/-22) and dimethylarsinic acid (373+/-12). In an indirect estimation, C. racemosa had 81+/-2% aresenosugars.

Arsenic↗

Suppression of 125I-uptake in mouse thyroid by seaweed feeding: possible preventative effect of dietary seaweed on internal radiation injury of the thyroid by radioactive iodine.

We conducted an animal experiment to determine how dietary seaweeds rich in iodine and dietary fibers suppress radioactive iodine uptake by the thyroid, using mice and four kinds of experimental diets, three with 1% or 2% powdered fronds of the kelp Laminaria religiosa and 2% powdered laver Porphyra yezoensis, and one with cellulose. Iodine content of a hot-water extract of the kelp was 0.530 +/- 0.001%, and its dietary fiber (DF) values were 52.8 +/- 1.2%. Iodine in an extract of the laver was 0.008 +/- 0.001%, and its DF values were 41.4% +/- 0.7%. A statistically significant reduction of 125I uptake by the thyroid, 3 hours after intragastric administration of the radionuclide at a dosage of 18.5 kBq or 185 kBq in 0.3 ml aqueous solution per mouse, was observed in mice previously fed the experimental diets containing 1% and 2% kelp during periods varying from 24 hours to 7 days. The degree of the suppression was observed to depend on the amount of iodine in the diet or in the injected sample, no matter whether organic or inorganic, judging from the results of an additional experiment. Thus, we conclude that previously fed iodine-rich material, especially dietary seaweeds rich in iodine and other minerals, vitamins, and beta-carotene, such as kelps or laver supplemented with inorganic iodine, may be effective in prevention of internal radiation injury of the thyroid.

Administration, Oral↗

Mesoherbivores reduce net growth and induce chemical resistance in natural seaweed populations.

Herbivory on marine macroalgae (seaweeds) in temperate areas is often dominated by relatively small gastropods and crustaceans (mesoherbivores). The effects of these herbivores on the performance of adult seaweeds have so far been almost exclusively investigated under artificial laboratory conditions. Furthermore, several recent laboratory studies with mesoherbivores indicate that inducible chemical resistance may be as common in seaweeds as in vascular plants. However, in order to further explore and test the possible ecological significance of induced chemical resistance in temperate seaweeds, data are needed that address this issue in natural populations. We investigated the effect of grazing by littorinid herbivorous snails (Littorina spp.) on the individual net growth of the brown seaweed Ascophyllum nodosum in natural field populations. Furthermore, the capacity for induced resistance in the seaweeds was assessed by removing herbivores and assaying for relaxation of defences. We found that ambient densities of gastropod herbivores significantly reduced net growth by 45% in natural field populations of A. nodosum. Seaweeds previously exposed to grazing in the field were less consumed by gastropod herbivores in feeding bioassays. Furthermore, the concentration of phlorotannins (polyphenolics), which have been shown to deter gastropod herbivores, was higher in the seaweeds that were exposed to gastropod herbivores in the field. This field study corroborates earlier laboratory experiments and demonstrates that it is important to make sure that the lack of experimental field data on marine mesoherbivory does not lead to rash conclusions about the lack of significant effects of these herbivores on seaweed performance. The results strongly suggest that gastropods exert a significant selection pressure on the evolution of defensive traits in the seaweeds, and that brown seaweeds can respond to attacks by natural densities of these herbivores through increased chemical resistance to further grazing.

Animals↗

Biotransformation and accumulation of arsenic in soil amended with seaweed.

For many coastal regions of the world, it has been common practice to apply seaweed to the land as a soil improver and fertilizer. Seaweed is rich in arsenosugars and has a tissue concentration of arsenic up to 100 micro/g g(-1). These arsenic species are relatively nontoxic to humans; however, in the environment they may accumulate in the soil and decompose to more toxic arsenic species. The aim of this study was to determine the fate and biotransformation of these arsenosugars in soil using HPLC-ICP-MS analysis. Data from coastal soils currently manured with seaweeds were used to investigate if arsenic was accumulating in these soils. Long-term application of seaweed increased arsenic concentrations in these soils up to 10-fold (0.35 mg of As kg(-1) for nonagronomic peat, 4.3 mg of As kg(-1) for seaweed-amended peat). The biotransformation of arsenic was studied in microcosm experiments in which a sandy (machair) soil, traditionally manured with seaweed, was amended with Laminaria digitata and Fucus vesiculosus. In both seaweed species, the arsenic occurs in the form of arsenosugars (85%). The application of 50 g of seaweed to 1 kg of soil leads to an increase of arsenic in the soils, and the dominating species found in the soil pore water were dimethylarsinic acid (DMA(V)) and the inorganic species arsenate (As(V)) and arsenite (As(III)) after the initial appearance of arsenosugars. A proposed decomposition pathway of arsenosugars is discussed in which the arsenosugars are transformed to DMA(V) and further to inorganic arsenic without appreciable amounts of methylarsonic acid (MA(V)). Commercially available seaweed-based fertilizers contain arsenic concentration between 10 and 50 mg kg(-1). The arsenic species in these fertilizers depends on the manufacturing procedure. Some contain mainly arsenosugars while others contain mainly DMA(V) and inorganic arsenic. With the application rates suggested by the manufacturers, the application of these fertilizers is 2 orders of magnitude lower than the maximum permissible sewage sludge load for arsenic (varies from 0.025 kg ha(-1) yr(-1) in Styria, Austria, to 0.7 kg ha(-1) yr(-1) in the U.K.), while a direct seaweed application would exceed the maximum arsenic load by at least a factor of 2.

Arsenates↗

Suppressive effects of the extracts of Japanese edible seaweeds on mutagen-induced umu C gene expression in Salmonella typhimurium (TA 1535/pSK 1002) and tumor promotor-dependent ornithine decarboxylase induction in BALB/c 3T3 fibroblast cells.

Some of epidemiological data indicated that ubiquitous consumption of seaweeds in Japan may be a possible protective factor against some types of tumor. To analyse this problem, the authors studied the antimutagenic and antitumor promotion activities in methanol-soluble extracts of typical edible seaweeds which showed suppressive effects on 3-amino-1,4-dimethyl-5H-pyrido[4,3-b]indol (Trp-P-1)-induced umu C gene expression in SOS response of Salmonella typhimurium (TA 1535/pSK 1002) and 12-O-tetradecanoylphorbol-13-acetate (TPA)-dependent ornithine decarboxylase induction in BALB/c 3T3 fibroblast cells. Although eight varieties of edible seaweeds including chlorophyta, Phaenophyta and Rhodophyta showed significant antimutagenic and antipromotion activities, they expressed the activities different from each other. Among these seaweeds, Enteromorpha prolifera ('Sujiaonori' in Japanese) and Porphyra tenera ('Asakusanori') showed relatively strong suppressive activities in both antimutagenic and antipromotion assays compared with other seaweeds. These seaweeds contained considerable amounts of beta-carotene as a possible active principle with anticarcinogenic activity. This compound was partially associated with the antimutagenic activity in the seaweed extract, but did not contribute to the antipromotion activity of seaweed extract under our experimental conditions. These results strongly suggest that Japanese edible seaweeds have possible antimutagenic and antipromotion activities probably associated with antitumor activity.

3T3 Cells↗

Effect of fiber-rich brown seaweeds on protein bioavailability of casein in growing rats.

The effect of three subtropical fiber-rich brown seaweeds, Sargassum hemiphyllum (Turn.) C. Ag., Sargassum henslowianum C. Ag. and Sargassum patens Ag., on protein bioavailability of casein were evaluated by growing rats. The effect of the fiber-rich seaweeds was determined by comparing the net protein ratio (NPR), true protein digestibility (TD), nitrogen balance (NB), biological value (BV), net protein utilization (NPU), and fecal and urinary nitrogen loss of seaweed-based diet groups with those of the cellulose-based control group. There were no significant differences on NPR and urinary N loss in all diet groups. However, the values of TD (range 85.6-89.6%), BV (range 96.9-97.5%), NPU (range 83.5-86.8%) and NB (range 145-150 mg/rat/day) in all seaweed-based diet groups were significantly (p<0.05, one-way ANOVA, Tukey-HSD) lower than those of the control group. On the contrary, the fecal weight (range 5.59-6.30 g/rat/day) and fecal nitrogen (N) loss (range 31.1-41.2 mg/rat/day) of rats fed seaweed-based diets were significantly (p<0.05, one-way ANOVA, Tukey-HSD) higher than that of the control one. However, the differences on the body weight gain and weight of internal organs including liver, kidney and spleen between the seaweed-based diets and control groups were insignificant except that the weight of caecum of rats fed seaweed-based diets was significantly (p<0.05, one-way ANOVA, Tukey-HSD) higher than that of control group. The possible mechanism of fiber-rich brown seaweeds in affecting the protein bioavailability of casein and their related nutritional implications were discussed.

Animals↗

B cell stimulating activity of seaweed extracts.

The activity of seaweed extracts on murine and human lymphocytes was studied in vitro. The extracts of some kind of seaweed, such as Hizikia fusiformis and Meristotheca papulosa, stimulated normal mouse spleen cells to proliferate. The responder cells are B cells, because the response was depleted by the treatment of spleen cells with anti-immunoglobulin (Ig) antibody and complement and being passed through a nylon wool column. This response is not due to lipopolysaccharide (LPS) contamination, because seaweed extracts can stimulate spleen cells of C3H/HeJ mice which are LPS low responders. Seaweed extracts also enhanced Ig production by B cells and tumor necrosis factor (TNF) production by macrophages. Furthermore, seaweed extracts stimulated human lymphocytes to proliferate. All these B cell stimulating activities of seaweed extracts associated with glycoproteins whose molecular weights resided in 100 kD. These results suggest that seaweed extracts have stimulating activity on B cells and macrophages and this ability could be clinically for the modulation of immune responses.

Animals↗

Effect of dietary Laminaria angustata (brown seaweed) on azoxymethane-induced intestinal carcinogenesis in male F344 rats.

The effect of dietary Laminaria angustata (brown seaweed) on azoxymethane (AOM)-induced intestinal carcinogenesis was studied in male F344 rats. Five-week old rats were fed semipurified diets containing 0 and 10% seaweed. When the rats were 7 weeks old, all except the vehicle-treated groups received weekly subcutaneous injections of AOM in normal saline for two weeks (20 mg/kg body wt/week). All animals were fed the experimental diets until the termination of the experiment, which was 28 weeks after the last AOM injection. The incidence (percent of animals with tumors) and multiplicity (tumors/animal) of small intestinal tumors did not differ significantly between the control and seaweed groups. The incidence and multiplicity of colon adenomas along with the size of colon tumors were increased in rats fed the seaweed diet compared with those fed the control diet. Dietary seaweed had no major effect on the concentration of fecal bile acids; however, the concentration of fecal cholesterol and total neutral sterols was decreased in the seaweed group. These results suggest that dietary seaweed increases the risk for colon tumors.

Adenocarcinoma↗

Are the reductions in nematode attack on plants treated with seaweed extracts the result of stimulation of the formaldehyde cycle?

Soil application to the roots of tomato plants (Lycopersicon esculentum) of a commercially-available alkaline extract of the brown alga, Ascophyllum nodosum, resulted in a significant reduction in the number of second-stage juveniles of both Meloidogynejavanica and M. incognita invading the roots, compared to those of plants treated with water alone. Egg recovery from the seaweed extract treated plants was also significantly lower. The three major betaines found in the seaweed extract (gamma-aminobutyric acid betaine, delta-aminovaleric acid betaine and glycinebetaine) also led to significant reductions in both the nematode invasion profile and egg recovery when applied at concentrations equivalent to those present in the extract. This led to the conclusion that the betaines present in the seaweed extract play a major role in bringing about the observed effects. Treatment of Arabidopsis thaliana plants with seaweed extract also resulted in a significant decrease in the number of females of M. javanica which developed in the roots. Significant reductions in egg recovery were also achieved from plants treated with the seaweed extract and similar effects were produced with the betaines found in the seaweed extract. As the experiments were conducted under monoxenic conditions, it can be concluded that the results obtained with the application of either the seaweed extract or betaines are not dependent on microorganisms associated with the rhizosphere.

Animals↗

Benthic macrofauna changes in areas of Venice lagoon populated by seagrasses or seaweeds.

Two areas of the Venice lagoon populated by seagrasses (three stations covered by Cymodocea nodosa (Ucria) Asherson, Zostera marina Linnaeus, Zostera noltii Hornemann) or seaweeds (two stations: one covered by Ulva rigida C. Agardh and another at present without seaweed biomass) were monitored by means of six surveys over a year in order to study macrofaunal composition and seasonal changes. The seagrass stations showed a mean species richness (28-30 S m(-2)), individual abundance (1854-4018 N m(-2)) and biomass (22.3-37.7 g m(-2) ash-free-dry-weight, AFDW) ca. 3-8 times higher than those populated by seaweeds (10-15 S m(-2), 494-1395 N m(-2) and 5.6-13.7 g m(-2) AFDW). Differences among seagrass or seaweed stations were much lower. The Ulva-dominated station showed a macrofauna completely different both from the other stations and the communities recorded ca. 30 years ago, before the prolific growth of Ulva. In this station, frequent biomass decompositions and anoxic crises created critical conditions for life favouring organisms with reduced life cycles, younger individuals and the epifaunal species instead of the infaunal ones. In particular, Ulva grazers and scrapers such as Gammarus aequicauda Stock and Gibbula adriatica Philippi were found to be by far the most abundant species, whereas the taxa characteristic of the associations found in the past, in the presence of seagrasses or seaweeds and typical of low eutrophicated environments, appear strongly reduced. Marked differences in the macrophyte dominance and in the bio-physico-chemical variables which characterise the main environmental conditions of the Venice lagoon support the different distribution and composition of macrofaunal communities. Seaweed stations appear mainly governed by the seasonal cycles of these un-rooted macrophytes which, by alternating periods of production and decomposition, are responsible for the drastic reduction of macrofauna biodiversity and biomass. Conversely, seagrass stations exhibit a better oxidisation of the environment and show conditions more favourable for macrofauna colonisation, especially in the presence of macrophytes which are characterised by very well developed below-ground systems such as Cymodocea nodosa.

Animals↗

Geographic and genetic variation in feeding preference for chemically defended seaweeds.

Genetic variation in tolerance for seaweed chemical defenses among populations of marine herbivores is poorly described, either because few marine studies have addressed the issue or because the feeding preferences of populations of marine herbivores rarely differentiate. We address this using the polyphagous amphipod Ampithoe longimana (Crustacea), which commonly consumes, and lives on, terpene-rich brown seaweeds in the tropical genus Dictyota. Previous work demonstrated that in areas where the amphipod and seaweeds overlap geographically, these chemically deterrent seaweeds provide the amphipod with spatial refuges from predation by omnivorous fishes that avoid consuming Dictyota. However, the amphipod's distribution extends northward of the seaweed's distribution, making this benefit of association unavailable to more northerly populations of amphipods. On average, populations sympatric with Dictyota have stronger feeding preference for Dictyota species and greater fitness when raised on Dictyota than do populations that are outside Dictyota's geographic endpoint. These results are consistent with the hypothesis that in areas where the amphipods co-exist with Dictyota, selection favors amphipods that tolerate Dictyota's chemical defenses and thereby access its enemy-free space. Amphipods allopatric with Dictyota are unable to tolerate Dictyota's chemical defenses, either because of selection, drift, or some mix of both forces. A trade-off between preference for Dictyota and for the chemically distinct seaweed Hypnea musciformis is suggested by the finding that populations with high preference for Dictyota tended to have lower preference for Hypnea. However, this population-level pattern appears to be the result of independent evolution and not a genetic trade-off: an analysis of full-sib families within a single population detected heritable variation in preferences for Dictyota and suggested this for Hypnea (although the latter was not statistically significant), yet no family-level covariation was detected. Phylogeographic analysis of mitocondrial DNA and nuclear sequences indicates a strong historical break between populations sympatric with Dictyota and populations more than 500 km beyond Dictyota's geographic endpoint. The historical separation of these most northerly populations could have allowed a random accumulation of alleles to effectively degrade tolerance for Dictyota, although selection may also be responsible for the degradation.

Amphipoda↗

Nutritional analysis of Vietnamese seaweeds for food and medicine.

Vietnamese edible marine macro-algae (seaweed) are of interest because of their value in nutrition and medicine. Vietnamese living in the coastal have traditionally utilized seaweeds species as food supplement and herbal medicine. They consumed seaweed as food in various forms: raw as salad and vegetable, pickle with sauce or with vinegar, relish or sweetened jellies and also cooked for vegetable soup. As herbal medicine, seaweed is usually used for traditional comestics, treatments for cough, asthma, hemorrhoid, boils, goitres, stomach ailments, urinary diseases, reduce the incidence of tumors, ulcers and headaches. Although Vietnam has an abundance of algae floral with total number of species is estimated to be nearly 1000 spp. in which there are 638 species of marine algae identified. However, there have been no intense study regarding to changes in marine algal chemistry. The fifteen species of edible seaweeds studied in this paper include green, brown and red algae. The proximate compositions as ash, protein, lipid, carbohydrate, fatty acids, vitamins, pigments, macro and micro-elements were compared among different species examined and the genetic relationships among them by analyzing the species-specific differences in nucleotide sequences of ITS-1 region of the ribosomal DNA was identified.

Dietary Supplements↗

An HPLC method for the quantification of sterols in edible seaweeds.

This study presents an HPLC method for the quantification of sterols in edible seaweeds. Sterols were identified by HPLC/mass spectrometry (HPLC-MS) in positive APCI mode. The samples were saponified by refluxing with 1 m ethanolic KOH, and the non-saponifiable fraction was extracted with hexane. Sterols were quantified by HPLC with UV detection (HPLC-UV), on a 15 x 0.4 cm Kromasil 100 C(18) 5 micro m column (mobile phase 30:70 v/v methanol:acetonitrile; fl ow rate 1.2 mL/min; column temperature 30 degrees C; detection wavelength 205 nm). Method repeatability for fucosterol was good (coefficient of variation 2.4%). Sterol contents were determined in canned or dried brown seaweeds (Himanthalia elongata, Undaria pinnatifida, Laminaria ochroleuca) and red seaweeds (Palmaria sp., Porphyra sp.). The predominant sterol was fucosterol in brown seaweeds (83-97% of total sterol content; 662-2320 micro g/g dry weight), and desmosterol in red seaweeds (87-93% of total sterol content; 187-337 micro g/g dry weight).

Chromatography, High Pressure Liquid↗

Extraction and detection of arsenicals in seaweed via accelerated solvent extraction with ion chromatographic separation and ICP-MS detection.

An accelerated solvent extraction (ASE) device was evaluated as a semi-automated means of extracting arsenicals from ribbon kelp. The effect of the experimentally controllable ASE parameters (pressure, temperature, static time, and solvent composition) on the extraction efficiencies of arsenicals from seaweed was investigated. The extraction efficiencies for ribbon kelp (approximately 72.6%) using the ASE were fairly independent (< 7%) of pressure, static time and particle size after 3 ASE extraction cycles. The optimum extraction conditions for the ribbon kelp were obtained by using a 3 mL ASE cell, 30/70 (w/w) MeOH/H2O, 500 psi (1 psi = 7 KPa), ambient temperature, 1 min heat step, 1 min static step, 90% vol. flush, and a 120 s purge. Using these conditions, two other seaweed products produced extraction efficiencies of 25.6% and 50.5%. The inorganic species present in the extract represented 62.5% and 27.8% of the extracted arsenic. The speciation results indicated that both seaweed products contained 4 different arsenosugars, DMA (dimethylarsinic acid), and As(V). One seaweed product also contained As(III). Both of these seaweed products contained an arsenosugar whose molecular weight was determined to be 408 and its structure was tentatively identified using ion chromatography-electrospray ionization-mass spectrometry/mass spectrometry (IC-ESI-MS/MS).

Arsenicals↗

Total arsenic, inorganic arsenic, lead and cadmium contents in edible seaweed sold in Spain.

Total arsenic, inorganic arsenic, lead and cadmium contents were determined in 112 samples of seaweed preparations sold in Spain (seaweed packed in plastic or cardboard box, seaweed in the form of tablets and concentrates, foods containing seaweed, and canned seaweed). The concentration ranges found, expressed in mg/kg, dry weight, were: total As (0.031-149), inorganic As (<0.014-117), Pb (<0.050-12.1) and Cd (<0.003-3.55). For all the contaminants there were failures to comply with legislated values. In particular, all the samples of Hizikia fusiforme exceeded the inorganic As limit established in some countries, and a considerable number of species exceeded the Cd limit set by international regulations. With respect to food safety, consumption of 3 g/day of the samples analysed could represent up to 15% of the respective Tolerable Daily Intakes (TDI) established by the WHO. The situation is especially alarming for intake of inorganic As from H. fusiforme, which can be three times the TDI established.

Arsenic↗

Statistical evaluation of ecosystem properties influencing the uptake of As, Cd, Co, Cu, Hg, Mn, Ni, Pb and Zn in seaweed (Eucus vesiculosus) and common mussel (Mytilus edulis).

In order to evaluate the influence of geographically varying marine ecosystem properties on the uptake of trace elements in bioindicators, samples were taken of seaweed (Fucus vesiculosus) and common mussel (Mytilus edulis) along the North Sea and Baltic Sea coast. Seasonal variations of the bioindicator status were minimized by sampling within 1 month. Ecosystem properties considered were the geographical position, the salinity and the concentrations of the macroelements Ca, Fe, K, Mg, Na, P and S in the bioindicators. Trace elements studied were As, Cd, Co, Cu, Hg, Mn, Ni, Pb and Zn. Factor analysis of the concentration patterns in the bioindicators and of salinity as a function of location confirmed the influence of the geographically varying salinity on the biological uptake of macroelements and trace elements. This influence of salinity was higher in the case of seaweed than in the case of mussel. Comparison of the geographical courses of the macroelement and trace-element concentrations by cluster analysis revealed corresponding courses for As and Hg in both bioindicators. All other elements showed different courses in seaweed and mussel. Subsequent cluster analysis comparing locations with respect to the macroelement or trace-element concentration patterns in the bioindicators, indicated a clear separation of North and Baltic Sea locations. However, the trace-element concentration patterns provided a regionally less distinctive ecosystem arrangement than those of the macroelement ones. The results of the cluster analysis were verified by discriminant analysis forming groups of locations with respect to geographical position and salinity. Results of discriminant analysis demonstrated, both for seaweed and for mussel as bioindicators, that the location groups formed according to the macroelement concentration patterns corresponded well with the geographical regions in the order of salinity. On the other hand, location groups based on the trace-element concentration patterns again showed a modified less distinctive ecosystem arrangement than the location groups based on macroelement concentration patterns. This confirms modified conditions for the uptake of trace elements in seaweed or mussel in comparison to the uptake of macroelements.

Animals↗