Competitive interactions between Fucus spiralis L. and F. vesiculosus L. (Fucales,Phaeophyta)

Fucus spiralis forms a conspicuous belt in the upper intertidal on rocky shores of the NW Atlantic. The objective of this study was to determine whether competition among congeners plays a role in restricting the distribution of F. spiralis to the upper shore. A replacement series design was used to test the growth performance of F. spiralis and F. vesiculosus in monocultures and in mixtures set up in the mid shore level on an exposed rocky coast. For F. spiralis and F. vesiculosus, all individual and relative crowding coefficients were < 1 and > 1, respectively. Hence, F. vesiculosus succeeded relative to F. spiralis in all mixtures tested. This finding was confirmed by graphical analysis. Ratio diagrams showed that the outcome of long term interaction among the species is extinction of F. spiralis. The results lead to the conclusion that interaction among congeners is a major determinant of the restricted distribution of F. spiralis.     

Geographical variation in Fucus vesiculosus morphology in the Baltic and North Seas

Effects of stress and disturbance on morphology, reproductive effort, size and sex ratio were studied for Fucus vesiculosus populations from the Baltic Sea at Askö and the North Sea on the west coast of Sweden at Tjäm[otilde]. High morphological variation was found between Fucus populations, with significant differences in length and weight of individuals, thallus breadth, number of branches and receptacles and receptacle weight, not only between Baltic and North Sea populations but also between populations within the same area, differing in wave exposure. With increasing disturbance, individuals in both studied populations were smaller and less branched. Differences were observed in plant size, with longer, broader and more branched plants being found in Askö compared with Tjärnö. Fucus populations at Tjämö allocated more biomass to reproduction and had longer, heavier receptacles than at Askö. Although the observed morphological changes may be partly explained by differences in wave exposure and salinity between the two sites, it is not possible to rule out genetic differences between the Baltic and North Sea populations. However, it is unlikely that the variations observed within the populations and between populations from the same area are genetically determined.     

Intriguing asexual life in marginal populations of the brown seaweed Fucus vesiculosus

Reproduction of attached large brown algae is known to occur only by sexual zygotes. Using microsatellites we show evolution of asexual reproduction in the bladder wrack promoting population persistence in the brackish water Baltic Sea (< 6 psu). Here a dwarf morph of Fucus vesiculosus is dominated by a single clone but clonal reproduction is also present in the common form of the species. We describe a possible mechanism for vegetative reproduction of attached algae, and conclude that clonality plays an important role in persistence and dispersal of these marginal populations, in which sexual reproduction is impaired by low salinity.     

Variation in natural selection for growth and phlorotannins in the brown alga Fucus vesiculosus

Directional selection for plant traits associated with resistance to herbivory tends to eliminate genetic variation in such traits. On the other hand, balancing selection arising from trade-offs between resistance and growth or spatially variable selection acts against the elimination of genetic variation. We explore both the amount of genetic variation and variability of natural selection for growth and concentration of phenolic secondary compounds, phlorotannins, in the brown alga Fucus vesiculosus. We measured variation in selection at two growing depths and two levels of nutrient availability in algae that had faced two kinds of past growing environments. Genetic variation was low for growth but high for phlorotannins. The form and strength of selection for both focal traits depended on the past growing environment of the algae. We found strong directional selection for growth rate in algae previously subjected to higher ultraviolet radiation, but not in algae previously subjected to higher nutrient availability. Stabilizing selection for growth occurred especially in the deep growing environment. Selection for phlorotannins was generally weak, but in some past-environment-current-environment combinations we detected either directional selection against phlorotannins or stabilizing selection. Thus, phlorotannins are not selectively neutral but affect the fitness of F. vesiculosus. In particular, there may be a fitness cost of producing phlorotannins, but the realization of such a cost varies from one environment to another. Genetic correlations between selective environments were high for growth but nonexistent for phlorotannins, emphasizing the high phenotypic plasticity of phlorotannin production. The highly heterogeneous selection, including directional, stabilizing, and spatially variable selection as well as temporal change in selection due to responses to past environmental conditions, probably maintains a high amount of genetic variation in phlorotannins. Such variation provides the potential for rapid evolutionary response of phlorotannins under directional selection.     

Isolation and Analysis of the Cell Walls of Brown Algae: Fucus spiralis, F. ceranoides, F. vesiculosus, F. serratus, Bifurcaria bifurcata and Laminaria digitata

Mabeau, S. and Kloareg, B. 1987. Isolation and analysis of thecell walls of brown algae: Fucus spiralis, F. ceranoides, F.vesiculosus, F. serratus, Bifurcaria bifurcata and Laminariadigitata.—J. exp. Bot. 38: 1573–1580. Cell walls were isolated from six marine brown algae, Fucusspiralis, F. ceranoides, F. vesiculosus, F. serratus, Bifurcariabifurcata (Fucales, Phaeophyta) and Laminaria digitata (Laminariales,Phaeo-phyta). Yields of isolated cell walls ranged from 35–45%of thallus dry weight. Walls were composed mainly of alginatesand sulphatcd fucans, the proportions of which correlated withspecies zonation in the intertidal region. This result is consistentwith the hypothesis that the sulphated fucans are associatedwith the adaptation of macroalgae to the intertidal environment.Comparing the chemical composition of isolated cell walls withthat of whole plants, we conclude that alginic acid is mainlypart of the fibrillar wall while a significant proportion ofthe sulphated fucans probably belongs to the intercellular spacematrix. Since ascophyllan-like polysaccharides were more abundantin the fucan extracts from the isolated cell walls than fromthe whole plants, it is suggested that differences in the structureof fucans might be related to differences in their localizationthroughout the tissue. Key words: Cell walls, Phaeophyta, sulphated fucans     

Fouling mediates grazing: intertwining of resistances to multiple enemies in the brown alga Fucus vesiculosus

Macroalgae have to cope with multiple natural enemies, such as herbivores and epibionts. As these are harmful for the host, the host is expected to show resistance to them. Evolution of resistance is complicated by the interactions among the enemies and the genetic correlations among resistances to different enemies. Here, we explored genetic variation in resistance to epibiosis and herbivory in the brown alga Fucus vesiculosus, both under conditions where the enemies coexisted and where they were isolated. F. vesiculosus showed substantial genetic variation in the resistance to both epibiosis and grazing. Grazing pressure on the alga was generally lower in the presence than in the absence of epibiota. Furthermore, epibiosis modified the susceptibility of different algal genotypes to grazing. Resistances to epibiosis and grazing were independent when measured separately for both enemies but positively correlated when both these enemies coexisted. Thus, when the enemies coexisted, the fate of genotypes with respect to these enemies was intertwined. Genotypic correlation between phlorotannins, brown-algal phenolic secondary metabolites, and the amount of epibiota was negative, indicating that these compounds contribute to resistance to epibiosis. In addition, phlorotannins correlated also with the resistance to grazing, but this correlation disappeared when grazing occurred in the absence of epibiota. This indicates that the patterns of selection for the type of the resistance as well as for the resistance traits vary with the occurrence patterns of the enemies.     

Antioxidant and pro-oxidant activities of the brown algae, Laminaria digitata, Himanthalia elongata, Fucus vesiculosus, Fucus serratus and Ascophyllum nodosum

The ability of Laminaria digitata, Himanthalia elongata, Fucus vesiculosus, Fucus serratus and Ascophyllum nodosum to scavenge peroxyl radicals was investigated by kinetic studies in a model system. The thermal initiated oxidation of methyl linoleate was performed at 60°C in heptanol, with or without antioxidants. When they reached 1% of the substrate, seaweed extracts exhibited antioxidant activities by extending the induction period, but they did not suppress the rate of oxygen uptake as did vitamin E. A synergistic effect occurred when both algal extracts (1.5 g L^-1) and vitamin E (0.4 mmol L^-1) were present, and the effectiveness of the combined antioxidants during the whole induction period was vitamin E effectiveness. The synergistic effect of L. digitata, however, was subject to seasonal variations: samples collected in summer were effective synergists, whereas samples collected in winter displayed a marked negative synergism. The phospholipid fractions of F. vesiculosus, F. serratus and A. nodosum, including pigments, accounted for only 6% of the total lipid fraction, and did not exhibit a large synergistic effect. The main phospholipid was not phosphatidyl ethanolamine as usually related, but phosphatidyl inositol. Fucoxanthin had some antioxidant activity per se under our experimental conditions, but did not act as a synergist of vitamin E. The most potent synergists were recognized as chlorophyll a and related compounds by the application of liquid-liquid partition and chromatography for the identification of active components. This revised version was published online in June 2006 with corrections to the Cover Date.     

Dimethylsulphopropionate (DMSP) and proline from the surface of the brown alga Fucus vesiculosus inhibit bacterial attachment

It was demonstrated previously that polar and non-polar surface extracts of the brown alga Fucus vesiculosus collected during winter from the Kiel Bight (Germany) inhibited bacterial attachment at natural concentrations. The present study describes the bioassay-guided identification of the active metabolites from the polar fraction. Chromatographic separation on a size-exclusion liquid chromatography column and bioassays identified an active fraction that was further investigated using nuclear magnetic resonance spectroscopy and mass spectrometry. This fraction contained the metabolites dimethylsulphopropionate (DMSP), proline and alanine. DMSP and proline caused the anti-attachment activity. The metabolites were further quantified on the algal surface together with its associated boundary layer. DMSP and proline were detected in the range 0.12-1.08 ng cm(-2) and 0.09-0.59 ng cm(-2), respectively. These metabolites were tested in the concentration range from 0.1 to 1000 ng cm(-2) against the attachment of five bacterial strains isolated from algae and sediment co-occurring with F. vesiculosus. The surface concentrations for 50% inhibition of attachment of these strains were always <0.38 ng cm(-2) for DMSP and in four cases <0.1 ng cm(-2) for proline, while one strain required 1.66 ng cm(-2) of proline for 50% inhibition. Two further bacterial strains that had been directly isolated from F. vesiculosus were also tested, but proved to be the least sensitive. This study shows that DMSP and proline have an ecologically relevant role as surface inhibitors against bacterial attachment on F. vesiculosus.     

Influence of light and nitrogen on the phlorotannin content of the brown seaweeds Ascophyllum nodosum and Fucus vesiculosus

Phlorotannins, C-based defence compounds in brown seaweeds, show a high degree of spatial and temporal variation within seaweed species. One important model explaining this variation is the Carbon Nutrient Balance Model (CNBM), which states that the relative supply of carbon and limiting nutrients will determine the level of defence compounds in plants. Nitrogen is often considered to be the limiting nutrient for marine macroalgal growth and the CNBM thus predicts that when the carbon:nitrogen ratio is high, photosynthetically fixed carbon will be allocated to production of phlorotannins. In the present study, we evaluated the effects of light (i.e. carbon) and nitrogen on the phlorotannin content of two intertidal brown seaweeds, Ascophyllum nodosum and Fucus vesiculosus. This was done in an observational field study, as well as in a manipulative experiment where plants from habitats with different light regimes were subjected to different nitrogen and light treatments, and their phlorotannin content was measured after 14 days. The results showed that there was a negative relationship between tissue nitrogen and phlorotannin content in natural populations of F. vesiculosus, but not in A. nodosum. In the short term, the phlorotannin content in both algal species was not affected by changes in nitrogen availability. Exposure to sunlight had a positive effect on the phlorotannin content in natural populations of both algal species but, in the manipulative experiment, only F. vesiculosus showed a rapid response to changes in light intensities. Plants subjected to sunlight contained higher phlorotannin content than shaded plants. In conclusion, the results imply that nitrogen availability explains some of the natural variation in the phlorotannin content of F. vesiculosus, but the light environment has greater importance than nitrogen availability in predicting the phlorotannin content of each species.     

Physodes and the phenolic compounds of brown algae. Isolation and characterization of phloroglucinol polymers from Fucus vesiculosus (L.)

A series of vanillin-reactive compounds has been isolated from extracts of the brown alga Fucus vesiculosus. The first three members of this series have been examined by mass spectrometry, nuclear magnetic resonance spectrometry, and chemical methods, and were shown to be phloroglucinol, its phenyl-linked dimer, and a timer also composed of phloroglucinol units. Evidence is presented for the presence of tetrameric, pentameric, and hexameric phloroglucinol derivatives. Polymeric vanillin-reactive compounds were isolated and separated into two fractions. Oxidative hydrolysis of the major one produced phloroglucinol and related oligomers in low yields. The classical vanillin reactivity of physodes may thus be attributed to phloroglucinol-containing derivatives, as originally proposed by Crato (Crato, E.: Ber. Dtsch. Bot, Ges. 10, 295-302 (1892)).     

Hybridization and introgression in a mixed population of the intertidal seaweeds Fucus evanescens and F. serratus

The introduced Fucus evanescens (hermaphroditic) and the native F. serratus (dioecious) have been in secondary contact along the Danish coast of the Kattegat Sea for 60–100 years and dioecious hybrids have been observed at Blushøj for several years. Hybridization in Fucus is unusual because it appears to always involve a hermaphroditic and a dioecious parental pair. We determined the degree and spatial patterns of introgression for 286 individuals using 10 microsatellite loci and cpDNA. Hybrids accounted for nearly 13% of the population, yet parental species were well differentiated (F_ST = 0.633). The presence of F. evanescens chloroplasts in 100% of F₁ hybrids revealed asymmetrical hybridization. Fucus evanescens cpDNA was observed in 50% of introgressed and 5.4% of pure F. serratus, but no F. serratus cpDNA was found in F. evanescens. In contrast, nuclear DNA introgression was symmetrical with an equal amount (≈1.5%) of genes introgressed into each parental species. Survivorship and viability data suggest selection against hybrids in the hybrid zone.     

Cadmium tolerance and adsorption by the marine brown alga Fucus vesiculosus from the Irish Sea and the Bothnian Sea

Cadmium (Cd) uptake capacities and Cd tolerance of the marine alga Fucus vesiculosus from the Irish Sea (salinity 35 psu) and from the Bothnian Sea (northern Baltic, 5 psu) were quantified. These data were complemented by measurements of changes in maximal photosynthetic rate (P(max)), dark respiration rate and variable fluorescence vs. maximal fluorescence (F(v):F(m)). At concentrations between 0.01 and 1 mmol Cd l(-1), F. vesiculosus from the Bothnian Sea adsorbed significantly more (about 98%) Cd compared with F. vesiculosus from the Irish Sea. The photosynthetic measurements showed that the Bothnian Sea F. vesiculosus were more sensitive to Cd exposure than the Irish Sea algae. The algae from the Irish Sea showed negative photosynthetic effects only at 1 mmol Cd l(-1), which was expressed as a decreased P(max) (-12.3%) and F(v):F(m) (-4.6%). On the contrary, the algae from the Bothnian Sea were negatively affected already at Cd concentrations as low at 0.1 mmol Cd l(-1). They exhibited increased dark respiration (+11.1%) and decreased F(v):F(m) (-13.9%). The results show that F. vesiculosus from the Bothnian Sea may be an efficient sorption substrate for Cd removal from Cd contaminated seawater and this algae type may also have applications for wastewater treatment.     

Changes in the composition of photosynthetic pigments and cellular structure of the brown algae Fucus vesiculosus L. and F. serratus L. from the Barents Sea during a prolonged period of darkness

The effect of a prolonged absence of light on the photosynthetic pigment content and cell ultrastructure in Fucus vesiculosus L. and F. serratus L. from the Barents Sea was investigated during a 30-day experiment in summer and under natural conditions in winter (at the end of the polar night). With decreasing irradiance, the accumulation of photosynthetic pigments, mainly violaxanthin, occurred. In the absence of light, the contents and ratio of pigments remained unchanged. At the end of the experiment, the following changes occurred in the cellular ultrastructure: the disappearance of the stored substances and the change in the shape and compression of the physodes. Higher proportions of mitochondria in the cell and of cristae in the mitochondria are indicative of the enhancement of the respiration processes. No degradational changes were found in the tissues and cells of fucoids during prolonged periods of darkness. This suggests the possibility of their transition from photoautotrophy to endogenous nutrition.     

Variation in phlorotannin content within two species of brown macroalgae (Desmarestia anceps and D. menziesii) from the Western Antarctic Peninsula

Phlorotannins are metabolites found only in the Phaeophyceae (brown algae) and have a variety of metabolic roles, including both primary (e.g. cell wall construction) and secondary (e.g. herbivore defence and UV protection). This study aimed to establish the level of variation of phlorotannins in two species of dominant Antarctic macroalgae (Desmarestia anceps and D. menziesii). Thirteen samples were taken from specific locations throughout the thallus of multiple individuals at two depths at three locations near Anvers Island, Antarctica. Overall average concentrations were 0.117±0.003 g g^–1dwt in D. anceps and 0.052±0.002 g g^–1dwt in D. menziesii. Concentrations varied greatly at the 13 sampling locations in each individual, but not in any consistent manner. Significant variation occurred between locations in both species and between depths in D. anceps.     

Isolation of humic acid from the brown algaeAscophyllum nodosum,Fucus vesiculosus,Laminaria saccharina and the marine angiospermZostera marina

Recent discovery of humic acid (HA) in the free-living, brown algaPilayella littoralis has prompted a search for HA in other live plants. Marine algaeCodium fragile andMonostoma oxyspermum (greens),Chondrus crispus,Palmaria palmata andPolysiphonia lanosa (reds),Ascophyllum nodosum, Fucus vesiculosus andLaminaria saccharina (browns) andZostera marina (marine angiosperm) were investigated for their HA content. Only the brown algae and the marine angiosperm contained HA, which was extracted by a standard procedure augmented with necessary removal of alginic acid (where applicable). The isolated products were identified as HA by comparison of their analytical data, uv-visible, FTIR,¹H NMR spectra and morphologies with those of authentic HA isolated from municipal compost.Authors for correspondence     

Lipid Metabolism in the Brown Marine Algae Fucus vesiculosus and Ascophyllum nodosum

Lipid metabolism and environmental effects on this process havebeen studied in the marine brown algae Fucus vesiculosus andAscophyllum nodosum. These algae showed very similar patternsof lipid metabolism during 24 h incubations. Labelling from[1-¹⁴C]acetate showed the major labelled lipids to be the ß-alanineether lipid and the neutral lipid fraction in both algae. Ofthe glycolipids, only sulphoquinovosyldiacylglycerol was welllabelled and the phosphoglycerides were all poorly labelled.The major labelled fatty acids were palmitate and oleate, againin both algae, although Fucus vesiculosus also showed significantlabelling of stearate and behenate. Although the amount of fattyacid labelling increased with time, the proportion of labelin palmitate and oleate remained approximately constant. Verylong chain fatty acids (arachidic, behenic) were increasinglylabelled with time. Lowered incubation temperatures decreased labelling of the saturatedfatty acids. Cu²⁺ increased the proportion of oleate labelledin both algae, and of linoleate in Fucus vesiculosus. This cationdecreased the percentage labelling of stearate and myristatein Ascophyllum nodosum. Lipid metabolism in Ascophyllum nodosumwas more sensitive to raised Cu²⁺ levels than in Fucus vesiculosus Key words: Acyl lipid metabolism, Fucus vesiculosus, temperature effects, Ascophyllum nodosum, copper pollution     

Rhodobacteraceae on the marine brown alga Fucus spiralis are abundant and show physiological adaptation to an epiphytic lifestyle

Macroalgae harbour specific microbial communities on their surface that have functions related to host health and defence. In this study, the bacterial biofilm of the marine brown alga Fucus spiralis was investigated using 16S rRNA gene amplicon-based analysis and isolation of bacteria. Rhodobacteraceae (Alphaproteobacteria) were the predominant family constituting 23% of the epibacterial community. At the genus level, Sulfitobacter, Loktanella, Octadecabacter and a previously undescribed cluster were most abundant, and together they comprised 89% of the Rhodobacteraceae. Supported by a specific PCR approach, 23 different Rhodobacteraceae-affiliated strains were isolated from the surface of F. spiralis, which belonged to 12 established and three new genera. For seven strains, closely related sequences were detected in the 16S rRNA gene dataset. Growth experiments with substrates known to be produced by Fucus spp. showed that all of them were consumed by at least three strains, and vitamin B₁₂ was produced by 70% of the isolates. Since growth of F. spiralis depends on B₁₂ supplementation, bacteria may provide the alga with this vitamin. Most strains produced siderophores, which can enhance algal growth under iron-deficient conditions. Inhibiting properties against other bacteria were only observed when F. spiralis material was present in the medium. Thus, the physiological properties of the isolates indicated adaption to an epiphytic lifestyle.