Interactions of Patella and macroalgae with settling Semibalanus balanoides (L.)

Biological interactions affecting densities of settling and newly-settled Semibalanus balanoides (L.) have been investigated by manipulative field experiments on the Isle of Man.The effects of sweeping by fucoid clumps of different species and Patella browsing have been compared on moderately-exposed shores. Patella allowed barnacle settlement by preventing growth of competing green algae, but reduced post-settlement densities. Small clumps of Fucus spiralis L., F. vesiculosus L, and F. serratus L. all reduced settlement considerably more so than limpets. F. serratus had the greatest sweeping effect.Interactions between macroalgae and Semibalanus balanoides have been investigated at all levels on sheltered shores and low down on more exposed shores. In the Fucus spiralis and F. vesiculosus zones, post-settlement numbers were higher than in adjacent areas where the canopy was removed. Barnacles did not settle readily in the Ascophyllum zone in either experimental or control areas. Settlement occurred in the upper part of the Fucus serratus zone in experimental areas where the canopy was removed but not in control areas. No settlement occurred in either treatment or control areas lower in the F. serratus zone. At all levels on the shore fucoid canopies seemed to reduce cyprid settlement, but the effect was greatest amongst F. serratus where there was total prevention. High on the shore the effect of enhanced post-settlement survival under the canopy outweighs reduction of cyprid settlement thus there are greater numbers in the controls. Competition with red algal turfs was shown to set the lower limit of the barnacle zone on a vertical pier face.     

Factors controlling the upper limits of fucoid algae on the shore

Year-round observations on the condition of intertidal seaweeds growing in situ on the shore, show that the upper limits of the zones characterized by Pelvetia canaliculata (L.) Done et Thur., Fucus spiralis L. and Ascophyllum nodosum (L.) Le Jol. were periodically pruned back by environmental conditions. The uppermost plants of each species showed clear signs of tissue damage 21 to 28 days after a time when drying conditions coincided with neap tides which exposed the plant to aerial conditions for long periods. High air temperatures aggravated the damage, but neither frost nor prolonged rain had any obvious adverse effects. On spring tides the plants were wetted every day and no damage resulted regardless of the weather.These species clearly all reach up to their physiological limits on the shore investigated, but presumably Fucus vesiculosus L. and F. serratus L. do not, for they were never observed to show signs of tissue damage attributable to exposure to air. Transplant experiments did, however, prove that F. serratus cannot survive in the F. spiralis zone and nor can F. spiralis persist in the Pelvetia canaliculata zone.Laboratory experiments also demonstrated that the ability to tolerate desiccation and then to resume photosynthesis and growth when re-submerged was greatest in P. canaliculata, the species found highest on the shore, and was progressively less in species inhabiting successively lower levels.     

The habitat and feeding preferences of Littorina obtusata (L.) and L. mariae sacchi et rastelli

The vertical zonation of the flat periwinkles Littorina obtusata (L.) and L. mariae Sacchi et Rastelli is different, but overlaps. At a sheltered location on the Isle of Cumbrae, Scotland, L. obtusata, although most abundant on Fucus vesiculosus L. and Ascophyllum nodosum (L.) Le Jol. on the midshore, ranged over the entire shore from the Pelvetia zone down to the upper limits of Laminaria. Littorina mariae exhibited a much more restricted distribution being associated predominantly with Fucus serratus L. on the lower shore. In laboratory trials with adult plants, flat periwinkles found fucoid algae both more attractive and more palatable than other seaweeds. Littorina obtusata also preferred the reproductive receptacles of fucoid algae to their vegetative tissue. Tiny germlings of a diversity of seaweeds were readily consumed by L. obtusata. In choice experiments germling Ulva lactuca L. was preferred to adult U. lactuca, whereas adult Fucus serratus was preferred to germling F. serratus. Contrasting foraging behaviours between different species of flat periwinkle are discussed in relation to differences in their radular dentition.     

PROTEINS IMMUNOLOGICALLY RELATED TO DEHYDRINS IN FUCOID ALGAE

Adult fucoid algae on Atlantic shores have well-characterized, species-specific tolerances to the varying levels of desiccation that occur from the low to high intertidal zones; however, less is known about embryonic tolerances and their mechanistic basis. We investigated this by 1) exposing embryos of Fucus evanescens C. Agardh, F. spiralis L., and F. vesiculosus L. from the Maine shore to osmotic desiccation in hypersaline seawater and 2) examining whether these embryos contain species-specific dehydrins, proteins first identified in higher plants that are hypothesized to confer tolerance to dehydration. Embryonic survival when cultured in hypersaline seawater >100 practical salinity units (psu) correlated with the position of these species in the intertidal zone (F. spiralis > F. vesiculosus > F. evanescens), but all 1-day-old embryos of these species tolerated treatment with 100 psu or lower seawater. Proteins (17–105 kDa) immunologically related to dehydrins were detected on western blots with dehydrin antibodies raised against a synthetic peptide representing the conserved motif of dehydrins in higher plants. These proteins were constitutive and unstable when subjected to prolonged (>15 min) temperatures above 55° C, unlike most higher plant dehydrins, which are inducible and remain soluble at 75°–100° C. The presence of these proteins was species- and stage-specific. Sperm of F. vesiculosus had a characteristic protein of 76 kDa, whereas eggs and embryos (6 h to 3 days old) had a 92-kDa protein. By 1 week of age, expression of the 92-kDa protein decreased, and the 35-kDa protein of adults was present. Embryos of A. nodosum L. and Pelvetia compressa J. Agardh DeToni contained an 85-kDa protein rather than the 92-kDa protein of Fucus embryos (F. distichus L., F. evanescens, F. spiralis, and F. vesiculosus). The 92-kDa protein became more abundant in embryos exposed to hyperosmotic seawater at 50 psu (F. evanescens and F. vesiculosus) or 150 psu (F. spiralis); however, dehydrin-like proteins of some molecular masses decreased in abundance simultaneously. Further characterization of these proteins is required to establish whether they protect embryos against intertidal desiccation.     

An ecological study of fucus spiralis L.

An ecological study of Fucus spiralis L. has been made at Jaffrey Point, Newcastle, New Hampshire and the adjacent Great Bay Estuary System from 1972–1975. The distribution, growth, reproductive periodicity, attrition and longevity of the plants are described in relation to a variety of environmental factors. F. spiralis shows a broad but discontinuous estuarine distribution within the Great Bay Estuary System. The presence or absence of appropriate substrata is considered to be a factor determining its discontinuous distribution, since it is usually associated with metasedimentary or metavolcanic rock outcrops. The maximum growth and reproduction of F. spiralis are during the summer. The summer growth rates ranged from 1.9 to 2.8 cm/month while the average growth rate throughout the year was 1.2 cm/month.F. spiralis populations showed two periods of major attrition, namely, during the winter and summer. The average longevity of F. spiralis plants is ≈ 2 yr. The F. spiralis zone lies in the uppermost intertidal area between +2.12 and +2.31 m above M.L.W. There is a micro-stratification of biomass, stature, and reproductive activity within this zone. Average plant weight, length, and fertility tend to decrease with increasing elevation.     

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.     

Seasonal growth and reproduction of estuarine fucoid algae in New England

Estuarine populations of Ascophyllum nodosum (L.) Le Jolis and Fucus vesiculosus L. var. spiralis Farlow from the Piscataqua River of the Great Bay Estuary System (New Hampshire-Maine) showed their maximum growth during the spring and fall of two consecutive years. The maximum growth rates (in terms of elongation) for Ascophyllum and Fucus were 3.5 and 3.6 cm/month, respectively, during 1973, and 3.2 and 2.3 cm/month in 1974. The average length of whole Fucus fronds showed a trend towards longer plants during the early spring and late summer-fall, while the average weight was significantly greater in the spring and early summer. Ascophyllum showed a distinct reproductive periodicity, with a maximum in March or April of both years; after the spring period of maximum reproduction, the receptacles dehisced simultaneously and the plants were vegetative during the summer. The reproductive periodicity of Fucus was not so distinct as that of Ascophyllum; even so, its maximum reproduction was in April of both years. The level of reproduction for Ascophyllum populations was three times greater in 1974 than in 1973. The reproduction and growth of the two fucoid algae are related to seasonal variations in temperature and light.     

The Effect of Elevated UV Radiation on Fucus spp. (Fucales,Phaeophyta) Zygote and Embryo Development

Abstract: This study has shown that in Fucus serratus and Fucus distichus, young zygotes and embryos are highly susceptible to elevated levels of both UVA (UVAR) and UVB radiation (UVBR). Zygotes treated with UVAR are able to polarise and germinate, but are very slow to divide; if they do, they often have skewed division planes or deformed rhizoids. Those treated with UVAR and UVBR remain spherical, they do not polarise, germinate to form rhizoids or undergo cell division. We suggest that the UVR may be affecting the cytoskeleton. Conversely, zygotes and embryos of Fucus spiralis are able to withstand these same UVR levels and, at the light microscope level, appear to develop normally. When the brown algal phenolic compound phloroglucinol was placed in a filter covering the developing embryos, normal development was seen under all treatments. Phenolic compounds protect the developing fucoids from UVR. In comparison with the other two species, Fucus spiralis grows high up on the shore and is exposed for much longer periods of time and, presumably, to higher levels of natural UVR. The failure of the juvenile stages of F. serratus and F. distichus to withstand UVR stress may have implications for the continued survival of these species in the intertidal, and may prove detrimental to the population as a whole if UVR levels increase.     

Annual variation in growth rate of perennial littoral fucoid algae from the west coast of Norway

The growth of apical segments of six perennial fucoid littoral algae (Pelvetia canaliculata (L.) Dec. et Thur., Ascophyllum nodosum (L.) Le Jol., Fucus spiralis L., F. vesiculosus L., F. serratus L. and F. distichus L. subsp. edentatus (Pyl.) Powell) was measured at approximately monthly intervals during 1 year.In winter, elongation was limited by temperature and light. Maximum growth in May–June coincided with minimum N concentration in the seawater, but with high N concentration in the apical tissue. Throughout summer, both growth and tissue N concentration decreased, indicating that the growth was nutrient limited.A close linear correlation is indicated between elongation and weight increase in apices of A. nodosum.     

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.     

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     

Short-term effects of temperature upon the growth of intertidal fucales

The growth rate of five species of intertidal Fucales (Pelvetia canaliculata (L.) Dec. et Thur., Fucus spiralis L., Fucus vesiculosus L., Fucus serratus L., Ascophyllum nodosum (L.) Le Jolis) was measured at temperatures from 2.5 to 35 °C. An increase in temperature immediately causes a high growth rate, and during the first hour it increases linearly with temperature; at 35 °C it is 20 times the control at 7 °C. This acceleration of growth is based mainly on stored photosynthate. After the first few hours the growth rate decreases rapidly, particularly at the highest temperatures. After 2–3 weeks a temperature optimum below 17.5 °C is indicated. High temperatures, 30–35 °C, were lethal to all species, with a survival time corresponding to their vertical zonation in the natural habitat.     

The effect of zinc on the increase in length of five species of intertidal fucales

The growth rate of Pelvetia canaliculata (L.) Dec. et Thur., Fucus spiralis L., F. vesiculosus L., F. serratus L., and Ascophyllum nodosum (L.) Le Jolis was measured in six different concentrations of zinc (0.025–14 mg/l) during a period of ten days. No significant growth response was observed at 0.1 mg/l of zinc. A significant reduction of growth during the first two to three days of exposure to 1.4 mg/l or more was followed by no or small further decline. Apices of A. nodosum from localities with different background load of zinc (0.1–0.5 mg/l and <0.01 mg/l) showed no difference in response to 2.6 mg/l of zinc.     

The effect of lead,cadmium, and mercury on the increase in length of five intertidal fucales

The increase in length of Pelvetia canaliculata (L.) Dec. et Thur., Fucus spiralis L., F. vesiculosus L., F. serratus L., and Ascophyllym nodosum (L.) Le Jolis was measured in various concentrations of lead, cadmium, and mercury during a period of 9–10 days. Concentration ranges of the three metals were 45 2600 μg 1, 1.5 1040 μg 1 and 0.9 1250 μg 1, respectively.Significant reductions of growth rate compared with the controls were observed at ? 810 μg 1 of lead, ? 450 μg/l of cadmium, and ? 10 μg/l of mercury, and regressions of growth reduction on log concentration were indicated.Growth was significantly enhanced in Pelvetia canaliculata and Ascophyllum nodosum when exposed to cadmium, and in this case there was a significant regression of growth iincrease on log concentration. The growth of Pelvetia canaliculata was also enhanced at all concentrations of lead.     

Longitudinal profiles of carbon dioxide fixation capacities in marine macroalgae

Fucus serratus L., Fucus spiralis L., and Fucus vesiculosus L. (Fucales, Phaeophyceae) as well as Laminaria digitata (Huds.) Lamour., Laminaria hyperborea (Gunn.) Fosl., and Laminaria saccharina (L.) Lamour. (Laminariales, Phaeophyceae) have been investigated for the distribution of enzymic CO₂ fixation capacities via phosphoenolpyruvate carboxykinase (EC 4.1.1.32) (PEP-CK) and via ribulose-1,5-bisphosphate carboxylase (EC 4.1.1.39) (RubP-C) in different regions of the thalli. The maximum of PEP-CK activity is found to be confined to the growing regions of the algae, while the activity of RubP-C achieves its highest values in the entirely differentiated parts of the fronds. These findings are confirmed by the results of photosynthetic and light-independent (dark) carbon assimilation as determined by in vivo ¹⁴CO₂ fixation. The physiological significance of these differential patterns of carboxylation patterns is discussed with respect to the ontogenetic stage and the chemical constitution of the different thallus parts.     

GENETIC ISOLATION BETWEEN THREE CLOSELY RELATED TAXA: FUCUS VESICULOSUS,F. SPIRALIS,AND F. CERANOIDES (PHAOPHYCEAE)1

All traditional markers, both phenotypic and phylogenetic, have failed to discriminate between the taxa composing the Fucus vesiculosus L., F. spiralis L., and F. ceranoides L. species complex, particularly in Brittany (France), so we used five microsatellite markers to compare the allelic frequencies of populations of the three taxa in this region. The aim of this study was to assess whether the different populations were grouped according to their geographical location, their habitat (open coast versus estuary), or their a priori taxonomic assignment. Species‐specific alleles were identified at one locus, demonstrating the utility of microsatellite markers for recognizing the three taxa in Brittany. Moreover, our results clearly support the separation of F. vesiculosus, F. spiralis, and F. ceranoides into distinct species, independently of geography. We also identified genetic differentiation between estuarine and coastal populations of F. vesiculosus.     

Growth interactions between littoral diatoms and juvenile marine algae

Interactions have been studied between juvenile plants of green, brown, and red marine algae and 31 diatom clones isolated from a variety of marine eulittoral habitats. The interactions seemed to be of an individual nature for both juvenile plants and diatoms. Germlings of Ulva lactuca L. mostly showed enhanced growth, often with significant increases in population sizes of the accompanying diatoms. Fucus spiralis L. germlings were mainly unaffected by growth in the presence of the diatom clones, but growth of the diatoms was often stimulated. Ascophyllum nodosum (L.) Le Jol. germlings were little affected, whilst the accompanying diatoms were less noticeably affected than with Fucus spiralis. Germlings of F. vesiculosus L. often showed growth inhibitions in the presence of diatoms, with many diatom populations showing enhanced growth. Similarly, the discoid encrusting sporelings of Chondrus crispus Stackh. showed growth inhibitions where there were measurable interactions, although the accompanying diatoms usually failed to show growth stimulations. The discoid sporelings of Gigartina stellata (Stackh. in With.) Batt. showed high mortalities, usually with marked increases in population sizes of accompanying diatoms.     

Characterization of microsatellite loci in the marine seaweed Ascophyllum nodosum (Phaeophyceae; Fucales)

Ascophyllum nodosum L. dominates rocky intertidal shores throughout the temperate North Atlantic. Six microsatellite loci were developed for A. nodosum using enriched libraries. The number of alleles ranged from 9 to 24 and heterozygosities from 0.2213 to 0.7785. Ascophyllum is monotypic. There was no cross‐reactivity observed with Fucus serratus, F. vesiculosus or F. evanescens.     

D-VITAMINS AND THEIR PRECURSORS AS GROWTH REGULATORS IN AXENICALLY CULTIVATED MARINE MACROALGAE1

Light petroleum extracts from thalli of Fucus vesiculosus (L.) and Laminaria digitata (Huds.) Lamour. stimulate growth of axenically cultured Fucus spiralis L. and Enteromorpha compressa (L.) Grev. in amounts per litre culture medium corresponding to one to 10 mg wet weight of the brown algae. This indicates the presence of a lipophilic, very active growth regulator in the blades. Extracts from the meristematic zone of Laminaria are much less active. Vitamins D₃, D₂ and 1,25-(OH)₂-D₃ at 1·10^?8– 1·10^?7 M increase the fresh weight of Enteromorpha compressa and Fucus spiralis up to 100% or more. The dry weight of Nemalion helminthoides (Veil, in With.) Batt. is also increased under the same conditions. Ergosterol and cholesterol are also growth-stimulating but higher concentrations (e.g. 1·10^?6 M) are necessary. The effects of the D vitamins manifest themselves without any auxins being added to the medium.     

Adventitious Bud Formation from Bulb-scale Explants of Lilium speciosum Thunb. in vitro Effects of aminoethoxyvinyl-glycine, 1-aminocyclopropane-1-carboxylic acid,and ethylene

Several representatives of marine brown macroalgae (Phaeophyceae) including Fucus serratus L., Fucus spiralis L. and Fucus vesiculosus L. as well as Laminaria digitata (Huds.) Lamour., Laminaria hyperborea (Gunn.) Foslie and Laminaria saccharina (L.) Lamour. were investigated with particular regard to features of biosynthesis of the storage product mannitol. The respective catalytic system involved in the last step of mannitol formation, mannitol-1-phosphate dehydrogenase, appears to be a cytoplasmic enzyme as may be judged from the degree of correlation with the chloroplast key enzyme ribulose-1,5-bisphosphate carboxylase in different tissues of Laminaria digitata and Laminaria saccharina. Activity of mannitol-1-phosphate dehydrogenase in vitro is not affected by mannitol-l-phosphate or free mannitol, suggesting that mannitol biosynthesis in vivo) is mainly controlled by the environment and/or developmental stage. Certain inorganic ions such as NO₃^- (including K⁺) exert a strong influence on the activity of mannitol1-phosphate dehydrogenase thus suggesting that the intracellular pools of stored NO₃^- and mannitol are confined to spatially separated cellular compartments.