SETTLEMENT AND SURVIVAL OF POLYSIPHONIA LANOSA (CERAMIALES) SPORES ON ASCOPHYLLUM NODOSUM AND FUCUS VESICULOSUS (FUCALES)1

The settlement patterns of spores of Polysiphonia lanosa (L.) Tandy on Ascophyllum nodosum (L.) Le Jolis and Fucus vesiculosus L. were studied using a flow tank. Settlement sites were defined as ‘sheltered’ or ‘exposed.’ Surface area calculations revealed non-random settlement on A. nodosum, with higher than expected spore frequencies on the thallus and lateral pits and lower than expected frequencies on the vesicles. Settlement of F. vesiculosus was random and significantly lower than on A. nodosum. On the shore, survival of sporelings from September (post-sporulation) to May (pre-sporulation) was highly non-random on both basiphytes. On A. nodosum, lateral pits ('sheltered') showed the highest survival frequency. Here the proportion of surviving sporelings increased over the study period, whereas the proportion on open thallus area ('exposed') decreased. On F. vesiculosus also preferential survival occurred on ‘sheltered’ sites such as vesicle/thallus interfaces and wounds. Between September and May, all P. lanosa sporelings were lost from ‘exposed’ areas (thallus surface and vesicles). Overall, frequencies of surviving sporelings were much greater on A. nodosum than on F. vesiculosus. These results are discussed with reference to basiphyte morphology, epiphyte removal mechanisms and the survival stratagy of P. lanosa.     

PHENOTYPIC DIFFERENTIATION AT SOUTHERN LIMIT BORDERS: THE CASE STUDY OF TWO FUCOID MACROALGAL SPECIES WITH DIFFERENT LIFE‐HISTORY TRAITS1

Marginal populations are often geographically isolated, smaller, and more fragmented than central populations and may frequently have to face suboptimal local environmental conditions. Persistence of these populations frequently involves the development of adaptive traits at phenotypic and genetic levels. We compared population structure and demographic variables in two fucoid macroalgal species contrasting in patterns of genetic diversity and phenotypic plasticity at their southern distribution limit with a more central location. Models were Ascophyllum nodosum (L.) Le Jol. (whose extreme longevity and generation overlap may buffer genetic loss by drift) and Fucus serratus L. (with low genetic diversity at southern margins). At edge locations, both species exhibited trends in life‐history traits compatible with population persistence but by using different mechanisms. Marginal populations of A. nodosum had higher reproductive output in spite of similar mortality rates at all life stages, making edge populations denser and with smaller individuals. In F. serratus, rather than demographic changes, marginal populations differed in habitat, occurring restricted to a narrower vertical habitat range. We conclude that persistence of both A. nodosum and F. serratus at the southern‐edge locations depends on different strategies. Marginal population persistence in A. nodosum relies on a differentiation in life‐history traits, whereas F. serratus, putatively poorer in evolvability potential, is restricted to a narrower vertical range at border locations. These results contribute to the general understanding of mechanisms that lead to population persistence at distributional limits and to predict population resilience under a scenario of environmental change.     

Production of fucoid brown algae <Emphasis Type="Italic">Ascophyllum nodosum</Emphasis> and <Emphasis Type="Italic">Fucus vesiculosus</Emphasis> in the White Sea

Year-round studies of photosynthesis and respiration in the fucoid brown algae Ascophyllum nodosum and Fucus vesiculosus in the White Sea were performed. The annual specific production of the macrophytes was determined to be 1314 and 1642 cal/g of wet weight for A. nodosum and F. vesiculosus, respectively. The total annual production of fucoids (4.88 × 10¹¹ kcal) comprised about 0.8% of the phytoplankton production in the White Sea.     

Isolation and cross‐species amplification of microsatellite loci from the fucoid seaweeds Fucus vesiculosus,F. serratus and Ascophyllum nodosum (Heterokontophyta,Fucaceae)

The Fucaceae is a family of brown seaweeds that dominate and frequently co‐occur on North Atlantic rocky shores. We developed nine polymorphic microsatellite markers for the fucoid seaweeds Fucus vesiculosus, F. serratus and Ascophyllum nodosum using a combined, enriched library. Six of these loci were polymorphic in at least two species, showing from two to eight alleles with heterozygosities ranging from 0.41 to 0.85. Loci were also tested on F. spiralis, revealing five polymorphic microsatellite loci in this species.     

Extracts of the Brown Seaweed <Emphasis Type="Italic">Ascophyllum nodosum</Emphasis> Induce Gibberellic Acid (GA<Subscript>3</Subscript>)-independent Amylase Activity in Barley

Extracts of the brown seaweed Ascophyllum nodosum have been used as a biostimulant to promote growth and productivity in a number of agricultural production systems. Although the extracts have been shown to improve seedling emergence and vigor in a variety of plants, including barley, the mechanism(s) of this growth-promoting effect is(are) largely unknown. In our study, A. nodosum extract induced amylase activity in barley seed-halves; a significant difference in amylase activity was observed in seeds without an embryo. The addition of activated charcoal to the treatment media negated the bioactivity of the extracts suggesting the organic nature of bioactive compounds in A. nodosum extracts. The extracts induced amylase activity in a gibberellic acid (GA)-deficient barley mutant (grd2). LC-MS-MS analysis failed to detect the presence of GA₃ in the extracts. ABA supplementation of the medium caused a significant reduction of amylase activity in GA-treated seeds compared with those treated with the A. nodosum extract. Taken together, our results suggest that the organic components of A. nodosum extract induce amylase activity independent of GA₃ and might act in concert with GA-dependent amylase production leading to enhanced germination and seedling vigor in barley. Being derived from a renewable resource, the bioactive compounds from A. nodosum could be used to improve crop productivity in sustainable agricultural systems.     

Effect of epiphytism on reproductive and vegetative lateral formation in the brown, intertidal seaweed Ascophyllum nodosum (Phaeophyceae)

In the brown alga Ascophyllum nodosum (L) Le Jolis, a common species on sheltered Northern temperate rocky shores, gametes are produced in receptacles that emerge from small depressions (lateral pits) along the branched frond. These lateral pits are also the preferred settling site for the obligate epiphyte Polysiphonia lanosa (L) Tandy. Therefore, epiphytism can be expected to interfere with host reproductive output. The present study investigated the potential impact of the epiphyte on A. nodosum in two series of laboratory experiments that measured: (i) the direct shading of the host plant underneath an epiphyte canopy; and (ii) the development of receptacles in clean and epiphytised A. nodosum segments (excised from individual fronds) over a 6 month period. These experiments showed that light reaching emerged fronds underneath a dense epiphyte cover was reduced by 40%, and this was independent of the degree of desiccation the epiphyte experienced. Concurrently, in the growth study with epiphytised A. nodosum segments (segments with one clean and one epiphytised lateral pit) total receptacle biomass per epiphytised fragment was significantly reduced compared with clean segments (0.52 g and 1.25 g per gram of frond segment, respectively), although this effect was only significant in A. nodosum from sheltered shores. However, expressed as biomass per lateral pit, receptacle biomass in the remaining clean lateral pits in epiphytised segments was significantly increased in segments from both shores, demonstrating that A. nodosum can at least partially compensate for the loss of production resulting from epiphytism.     

Biomass and height of Ascophyllum nodosum after two decades of continuous commercial harvesting in eastern Canada

With the increasing demand for seaweed resources worldwide, management must ensure that the harvest of wild seaweed stocks is sustainable. We evaluate the impact of over 25 years of commercial harvesting of Ascophyllum nodosum in eastern Canada by comparing the biomass and height of the seaweed in the late 1990s to the late 2010s over a broad spatial scale spanning the provinces of Nova Scotia and New Brunswick. There has been no significant decrease in the biomass of A. nodosum in either province, and biomass has increased in some regions of New Brunswick during that period. The average height of A. nodosum has decreased by 7.8 cm in Nova Scotia while it increased by 13.8 cm in New Brunswick. Biomass of A. nodosum in unharvested sectors was 7% higher than that in harvested sectors while height of A. nodosum in unharvested sectors in New Brunswick is similar to the values observed in harvested sectors. Over the same period, water temperature has increased in both provinces and, in recent years, has at times exceeded the optimal growing temperature for A. nodosum within bays in Nova Scotia. We conclude that the current management and harvest of A. nodosum in eastern Canada are sustainable and maintain the biomass and height of A. nodosum beds but that control sites are necessary to offer adequate comparisons as environmental conditions are changing.

     

Temporal dynamics of inducible anti‐herbivory defenses in the brown seaweed Ascophyllum nodosum (Phaeophyceae)1

Anti‐herbivory defenses support persistence of seaweeds. Little is known, however, about temporal dynamics in the induction of grazer‐deterrent seaweed traits. In two induction experiments, consumption rates of the periwinkle Littorina obtusata (L.) on the brown seaweed Ascophyllum nodosum (L.) Le Jolis were measured in 3‐d intervals. Changes in palatability of directly grazed A. nodosum were tested every 3 d with feeding assays using fresh and reconstituted seaweed pieces. Likewise, assays with fresh A. nodosum assessed changes in seaweed palatability in response to water‐borne cues from nearby grazed conspecifics. Consumption rates of L. obtusata varied significantly during the 27‐d induction phase of each experiment. Direct grazing by L. obtusata lowered palatability of fresh and reconstituted A. nodosum pieces to conspecific grazers after 15 d as well as after 6 and 12 d, respectively. After 12, 18, and 24 d, fresh A. nodosum located downstream of L. obtusata‐grazed conspecifics was significantly less palatable than A. nodosum located downstream of ungrazed conspecifics. Changes in L. obtusata consumption rates and A. nodosum palatability during both induction experiments suggest temporal variation of grazer‐deterrent responses, which may complicate experimental detection of inducible anti‐herbivory defenses.     

Changes in composition of rockweed (Ascophyllum nodosum) beds due to possible recent increase in sea temperature in Eastern Canada

Ascophyllum nodosum (rockweed) is the main economic resource of the seaweed industry in the Atlantic Provinces of Canada. The annual harvest steadily increased since 1995, reaching a historic peak of 37,000 tonnes in 2007. Due to a high demand for fertilizers and animal feed supplements derived from rockweed, this trend seems likely to continue. The current management plan for the sustainable harvest of the A. nodosum resource is considered conservative. The resource has been managed with a precautionary approach since 1995 to protect the integrity of the habitat. Acadian Seaplants Limited (ASL) has been granted approximately 90% of the government-issued licenses to harvest A. nodosum resources in the Maritimes. The Canadian approach is based on an annual harvest from a given bed and not strip-and-return after several years. Since 1995, ASL has proactively undertaken extensive annual surveys and research on biomass productivity of this renewable resource to establish acceptable annual exploitation rates. Historically, the rockweed beds of southwestern Nova Scotia (NS) have been almost 99% pure A. nodosum, with a minor component of Fucus vesiculosus. However, since 2004, a steady increase in F. vesiculosus, with a peak of 4.6% of the total biomass in 2008, was recorded. This coincided with one of the mildest winters on record for the Maritimes. This increase in temperature seemed to be also responsible for an unusual recruitment of the blue mussel Mytilus edulis in rockweed beds in some areas of southern New Brunswick (NB) in 2006, causing the detachment of up to 30% of the seaweed biomass in some harvesting sectors. Another phenomenon observed in southwestern NS during 2003 and 2004 was extensive ice damage on rockweed beds produced by an early melting of the ice, with losses of up to 90% of the rockweed biomass in some areas.     

The effects of priming treatments using seaweed suspensions on the water sensitivity of barley (Hordeum vulgare L.) caryopses

Two seaweed suspensions, one prepared from Ascophyllum nodosum (Linnaeus) Le Jolis (ANS), another from Laminaria hyperborea (Gunn.) Foslie (LHS), were evaluated for their effects on the water sensitivity of barley (Hordeum vulgare L.) seeds. Priming in either seaweed suspension did not reduce viability. Priming in ANS was beneficial to seeds germinated in elevated water levels. The suspension of A. nodosum reduced the water sensitivity of barley seeds better than either water or polyethylene glycol (PEG) priming treatments. ANS also reduced the microbial population on the seeds by 86%. The hygroscopic properties and the antibiotic effect of the suspension of A. nodosum resulted in greater oxygen availability to the embryo, enabling more seeds to germinate under oxygen deficient conditions.     

Fucose-containing polysaccharides in the brown algae Ascophyllum nodosum and Fucus vesiculosus

The fucose-containing, sulfated polysaccharides from Ascophyllum nodosum and Fucus vesiculosus were isolated by extraction with water adjusted to pH 2. Pure fractions were carefully separated by fractional precipitation with ethanol from aqueous solutions containing magnesium or calcium chloride. Progress in the fractionation efforts and purity of the fractions isolated were established by free-boundary and cellulose acetate clectrophoresis. Ascophyllan, two “complexes”, and a galactofucan were isolated from A. nodosum. An ascophyllan-like fraction, and a “complex” were isolated from F. vesiculosus. Mild, acid hydrolysis (0.02m hydrochloric acid, 1 h, 80°) converted each of the “complexes” into an electrophoretically faster-moving and a slower-moving component. The “complex” from F. vesiculosus comprised a greater proportion of the extract than did the two “complexes” from A. nodosum. In addition, the Fucus “complex” was richer in fucose*. However, the data suggest that neither species contains a pure fucan sulfate in the native state.     

Low Divergence in rDNA ITS Sequences Among Five Species of Fucus (Phaeophyceae) Suggests a Very Recent Radiation

Sequences from the two ribosomal DNA internal transcribed spacers (ITS1 and ITS2) were compared among five species of Fucus. Based on the present taxon sampling, parsimony analysis showed that Fucus serratus is the sister-group of the remaining Fucus species when Ascophyllum nodosum was used as an outgroup. The topology of the tree was (Fucus serratus (F. lutarius (F. vesiculosus (F. spiralis+F. ceranoides)))). The extremely low variation observed suggests a very recent radiation of the genus which supports the view widely accepted that the Fucales are among the most evolutionarily advanced of the brown algae. We further note that sequence differences between Fucus and Ascophyllum were 28%: this does not rule out the utility of ITS sequences within the Fucaceae. The very low number of informative positions allows to demonstrate empirically that distance matrix methods group on the basis of symplesiomorphies. Received: 17 February 1997 / Accepted: 17 April 1997     

An evaluation of the mortality of the brown seaweed <Emphasis Type="Italic">Ascophyllum nodosum</Emphasis> (L.) Le Jol. produced by cutter rake harvests in southern New Brunswick,Canada

Ascophyllum nodosum (rockweed) landings in the Atlantic Maritime provinces of Canada totalled 36,500 wet tonnes in 2009. Due to the relative slow growth, stochastic recruitment, and the importance of habitat protection, strict harvest regulations are in place in the region to maintain the integrity of this resource. Special harvesting rakes have been designed to cut the plants and not to dislodge the clumps and their holdfasts from the substratum. However, for various reasons, close to 6% of the biomass harvested annually contains holdfast material. This proportion is closely monitored by the province as it is assumed to represent clump mortality. However, due to the complex structure of A. nodosum clumps, this relationship with mortality is not simple. A study was carried out to evaluate the real impact of this detachment on the A. nodosum population of southern New Brunswick in 2004. The structure of harvested A. nodosum clumps with associated holdfast material was analyzed and compared to non-harvested clumps from the same harvest area. Results showed that when a rake strips a clump, it only detaches 17.4% of the holdfast surface, leaving 36.8% of the plant biomass and 80.3% of the shoot density intact. An analysis of storm-cast material from the same study area showed a similar effect in the clump structure, although the incidence of holdfast in the detached biomass could be as high as 30%. Due to the high biomass detached each year by coastal storms in New Brunswick, their impact on the A. nodosum resource is 21 times higher than the annual harvest.     

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.     

Pollution effects on littoral algal communities in the inner Oslofjord,with special reference toAscophyllum nodosum

Summary 1. Littoral algal vegetation was studied at three localities in the inner Oslofjord; one in the Drøbak area, and two in the more polluted inner basin.2. Regrowth and succession on cleared strips were followed for 16 months.3. The phaeophyceanAscophyllum nodosum was a common species in the inner part of the fjord at the turn of the century, but has disappeared with increase in pollution.4. Well-grown specimens ofAscophyllum nodosum were transplanted from the Drøbak area to the inner basin. No essential differences in vegetative growth and maturation of generative structures were observed.5. Growth and development ofFucus distichus ssp.edentatus were followed from the state of germlings to fertile plants.6. Regrowth proceeded more slowly in the Drøbak locality than in the localities in the inner basin, where a dense carpet ofEnteromorpha spp. etc. was formed.7. It is assumed that the increased competition for substrate and the shading effect of theEnteromorpha carpet reduces the chances ofAscophyllum nodosum germlings of becoming established in the inner basin.

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Verunreinigungseffekte auf litorale Algengemeinschaften im inneren Oslofjord unter besonderer Berücksichtigung vonAscophyllum nodosum

Kurzfassung In dem verunreinigten inneren Oslofjord tritt die BraunalgeAscophyllum nodosum (L.)Le Jol. nicht mehr auf. Mit der vorliegenden Untersuchung sollte geklärt werden, ob das verunreinigte Fjordwasser einen unmittelbar hemmenden Effekt auf die Entwicklung dieser Alge ausübt oder ob ein indirekter Einfluß vorliegt, indem bestehende biotische Faktoren eine ungünstige Konkurrenzsituation für diese Art bedingen. Zur Klärung dieser Frage wurden 3 Versuchsareale von Algenvegetation reingeschabt; anschließend wurden Wiederbesiedelung und Artensukzessionen über einen Zeitraum von 16 Monaten verfolgt. Das eine Versuchsgebiet lag bei Dröbak (30 km von Oslo entfernt), woFucus vesiculosus undAscophyllum nodosum in der litoralen Vegetation dominieren. Die beiden anderen Versuchsgebiete befanden sich in dem verunreinigten inneren Teil des Oslofjords, wo einjährige Grünalgen vorherrschen. Exemplare vonAscophyllum nodosum wurden von Dröbak in die im inneren Fjord gelegenen Versuchsgebiete transplantiert, um Wachstum und Entwicklung der Algen mit der Population bei Dröbak vergleichen zu können. Es wurde festgestellt, daß keine wesentlichen Unterschiede hinsichtlich des vegetativen Wachstums und der Reifung von Rezeptakeln auftraten. Eine kritische Phase stellt die Entwicklung der Keimpflanzen auf dem Substrat dar. Als Folge der Eutrophierung bildet sich innerhalb weniger Wochen ein dichter Bewuchs vonEnteromorpha spp. Es zeigte sich, daß die Konkurrenz um das Substrat und die reduzierte Lichtzufuhr durch denEnteromorpha-Bewuchs sich als entscheidend für das Fehlen vonAscophyllum nodosum erweisen.

     

Improved methods of analysis for betaines in <Emphasis Type="Italic">Ascophyllum nodosum</Emphasis> and its commercial seaweed extracts

Beneficial effects of seaweeds and their extracts on crop performance have been attributed to a variety of compounds, including the betaines which are quaternary ammonium betaines. Methods of analysis of betaines published thus far suffer from low sensitivity, lack of baseline separation of individual betaines and from interference from other sample constituents. A rapid cleanup protocol and a sensitive LC-MS/MS method of analysis were developed to afford baseline separation of four betaines in the brown alga Ascophyllum nodosum and its commercial seaweed extract. Using this method, the presence of glycine betaine, δ-aminovaleric acid betaine, γ-aminobutyric acid betaine and laminine in A. nodosum, and commercial extracts derived from A. nodosum, were confirmed and quantified. The major betaine present was γ-aminobutyric acid betaine accounting for 0.008–0.014% of the dry weight of the seaweed and 0.014–0.027% of the dry weight of the commercial extracts. Seasonal variation in betaine content was observed. Differences in the total betaine content were observed between A. nodosum of the yellow (0.011–0.017% dry weight) and the olive green (0.017–0.021% dry weight) coloured morphologies.     

COLONIZATION AND GROWTH OF ASCOPHYLLUM NODOSUM (PHAEOPHYTA) IN MAINE1

Colonization and growth of Ascophyllum nodosum (L.) Le Jol. were investigated in two Maine estuaries from 1972 to 1978. Following denudation of intertidal rock, substrata were initially colonized by Fucus vesiculosus L.; eventually, Ascophyllum supplanted Fucus, and became dominant in terms of percentage cover. Ascophyllum settled first and most densely in the low intertidal zone, but its fastest growth occurred in the mid-intertidal zone. Some, but not all, Ascophyllum germlings produced a vesicle within one year of colonization. The mean annual growth of A. nodosum was variable among sites, zones and years.     

Rapid bioassays to evaluate the plant growth promoting activity of <Emphasis Type="Italic">Ascophyllum nodosum</Emphasis> (L.) Le Jol. using a model plant, <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> (L.) Heynh

Ascophyllum nodosum extract products are used commercially in the form of liquid concentrate and soluble powder. These formulations are manufactured from seaweeds that are harvested from natural habitats with inherent environmental variability. The seaweeds by themselves are at different stages of their development life-cycle. Owing to these differences, there could be variability in chemical composition that could in turn affect product consistency and performance. Here, we have tested the applicability of using Arabidopsis thaliana as a model to study the activity of two different extracts from A. nodosum. Three different bioassays: Arabidopsis root-tip elongation bioassay, Arabidopsis liquid growth bioassay and greenhouse growth bioassay were evaluated as growth assays. Our results indicate that both extracts promoted root and shoot growth in comparison to controls. Further, using Arabidopsis plants with a DR5:GUS reporter gene construct, we provide evidence that components of the commercial A. nodosum extracts modulates the concentration and localisation of auxins which could account, at least in part, for the enhanced plant growth. The results suggest that A. thaliana could be used effectively as a rapid means to test the bioactivity of seaweed extracts and fractions.     

Species‐specific mediation of temperature and community interactions by multiple foundation species

Foundation species can provide habitat that modify abiotic and biotic processes that contribute to ecosystem function. While many studies have focused on the processes and consequences of a focal foundation species, understanding the ecological equivalence of co‐occurring foundation species is important to identify key species responsible for ecosystem function. Here, we investigated the relative contributions of co‐occurring foundation species on abiotic (temperature) and biotic responses of invertebrate species (recruitment, persistence, growth and survival). In a series of experimental field studies, we manipulated foundation species to measure invertebrate recruitment, persistence, and predation. A laboratory experiment measured foundation species effects on herbivore growth. Results demonstrated that macroalgal (Fucus vesiculosus ecad and Ascophyllum nodosum ecad scorpioides) intermediate foundation species provide habitat, food, and alleviate abiotic stress for dominant littorinid herbivores that surpass that provided by the primary species (Spartina alterniflora). These foundation effects were species‐specific with F. vesiculosus ecad important for early life‐history stages (enhanced recruitment and early growth of littorinid snails) and A. nodosum ecad important later on as a refuge from predators (Carcinus meanas) and stressful temperature. Understanding of the different effects of co‐occurring foundation species on population and community processes is necessary for predicting community response to natural disturbance, species invasion, and ecosystem‐based management actions.