Seasonal changes in situ grazing of the mesoherbivores Idotea baltica and Gammarus oceanicus on the brown algae Fucus vesiculosus and Pylaiella littoralis in the central Gulf of Finland, Baltic Sea

The in situ grazing experiments were performed in the shallow water rocky habitat of the northern Baltic Sea during ice-free season 2002. In the experiments the effects of algal species and choice on the grazing of the mesoherbivores Idotea baltica (Pallas) and Gammarus oceanicus Segerstråle were tested. Salinity, temperature, concentration of nutrients in water and macroalgae and net production of macroalgae were considered as random effects in the analysis. The invertebrate feeding rate was mainly a function of the net photosynthetic activity of Pylaiella littoralis (L.) Kjellman and Fucus vesiculosus L. Feeding rate increased significantly with decreasing algal photosynthetic activity. When the two algal species were incubated together invertebrates fed primarily on P. littoralis. Low selectivity towards P. littoralis coincided with its high photosynthetic activity. The presence of F. vesiculosus did not modify the invertebrate feeding on P. littoralis. The results indicated that (1) the grazing on F. vesiculosus depended on the availability of P. littoralis, (2) the photosynthetic activity of algae explained the best the variation in grazing rate and (3) the grazers are not likely to control the early outbreak of filamentous algae in the northern Baltic Sea by avoiding young and photosynthetically active algae. The likely mechanism behind the relationship is that the increased photosynthetic activity of macroalgae coincides with their higher resistance to herbivory.     

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.     

Genetic and environmental variation in performance of a marine isopod: effects of eutrophication

Environmental variation in food resources modifies performance of herbivores, in addition to genetic variation and maternal effects. In marine benthic habitats, eutrophication may modify herbivores diets by changing host species composition or nutritional quality of algae for herbivores. We studied experimentally the effects of diet breadth and nutrient availability for the host algae on fitness components of the herbivorous isopod Idotea baltica. We fed the adult isopods with the brown algae Fucus vesiculosus and Pilayella littoralis and juveniles with the green alga Cladophora glomerata. By using half-sib families, we were able to separate the genetic, environmental and maternal effects on intermolt duration and size of the juveniles. The mothers confined to the diet consisting of both Fucus and Pilayella grew better and produced larger egg mass than those having consumed Fucus alone. Nutrient enhancement of algae did not influence the performance of the adult herbivores. However, the juveniles achieved twice the weight as well as shorter intermolt duration when consuming nutrient-treated C. glomerata. Mothers nutrition, either nutrient enrichment of her food algae or diet breadth, did not affect juvenile survival or growth as such, but we found evidence that the broader diet consumed by the mother mediated offspring performance by further accelerating growth rate of the offspring that fed on nutrient-treated alga. Intermolt duration was a highly heritable trait, but size showed very low heritability. Instead, maternal effects on size were substantial, suggesting that differences among mothers in their egg-provisioning ability strongly affect weight gain of their offspring. A high amount of additive genetic variance in intermolt duration implies potential for quick evolutionary responses in the growth schedule in the face of changes in the selective environment. We conclude that eutrophication, in addition to improving growth and reproduction of I. baltica by enhancing food quality and by providing opportunity for broader, more profitable diets, may act as a selective agent on life-history traits. Eutrophication of coastal waters is thus likely to reflect in herbivore density, population dynamics and eventually, grazing pressure of littoral macroalgae.     

Interaction between isopod grazing and wave action: a structuring force in macroalgal communities in the southern Baltic Sea

The macroalgal belt in the southern Baltic Sea may be partly structured by the interaction of physical and biological factors. A field study, spanning the 1990s, describes a rapid decline of the Fucus spp. stands along the wave-exposed Swedish southeast coast. During this period, a relative dominance of Fucus vesiculosus L. shifted to a relative dominance of Fucus serratus L. The decline of F. vesiculosus coincided with observations of large numbers of the grazing isopods Idotea baltica (Pallas) and Idotea granulosa Rathke, or with field observations of frequent grazing marks on Fucus fronds. I. baltica, but not I. granulosa, tended to aggregate in the declining Fucus spp. stands, indicating a strong preference for Fucus spp. In a mesocosm experiment I. baltica, when given a choice, grazed both Fucus species at weak water motion. At strong water motion grazing was concentrated on F. vesiculosus. It is hypothesized that one of the reasons I. baltica preferred F. vesiculosus to F. serratus at strong water motion may have been differences in habitat quality, like width of thallus, influencing the ability to cling to the plant. Smaller thallus, as in F. vesiculosus, thus is the preferred habitat for grazing of I. blatica. We postulate that the existence of F. serratus in the area may be favoured by strong wave action and moderate but not strong grazing by I. baltica, relaxing the interspecific competition from F. vesiculosus.     

Responses of growth and phlorotannins in <Emphasis Type="Italic">Fucus vesiculosus</Emphasis> to nutrient enrichment and herbivory

Discharge from anthropogenic sources may modify both macroalgal growth patterns and resource allocation to carbon based secondary compounds, thereby affecting their susceptibility for herbivory. We tested the effect of eutrophication in terms of nutrient enhancement on growth and phlorotannin concentration of Fucus vesiculosus by conducting manipulative experiments in the field and mesocosms. In the field experiment we utilised fish farms as nutrient sources and in the mesocosm-experiment we manipulated ambient nutrient levels and occurrence of the herbivorous isopod Idotea baltica. Vicinity of a fish farm affected neither growth nor the phlorotannin concentration of Fucus but increased the amount of epiphytes growing on Fucus. Other organisms such as epiphytic filamentous algae and periphyton, which are more capable of quickly utilizing excess nutrients, may restrain the direct effects of nutrient enhancement on Fucus. In a manipulative mesocosm experiment, neither nutrient enrichment nor occurrence of herbivores affected phlorotannin concentration implying lack of induced defences, at least in terms of increasing phlorotannin concentration. Feeding of thallus decreased the growth rate of algae, but the number of reproductive organs, receptacles, was not affected by herbivory. The negative effect of herbivory on the amount of apical tips tended to be stronger under nutrient enriched conditions. We conclude that eutrophication processes, in terms of nutrient enrichment, does not have strong direct effect on growth or phlorotannin production of F. vesiculosus. However, there may be important indirect consequences. First, herbivory may be targeted more to apical parts of the thallus under eutrophicated conditions. Second, the result that Fucus growing close to nutrient sources were smaller than those in control areas may reflect differences in mortality schedules of algae between eutrophicated and control areas.     

Photosynthesis and UV-B Tolerance of the Marine Alga Fucus vesiculosus at Different Sea Water Salinities

The marine algal species in the Baltic Sea are few due to the low sea water salinity. One of the few species that can be found is Fucus vesiculosus. Even this species is affected by the low salinity and becomes smaller in size in the Baltic. In present work the photosynthesis of F. vesiculosus in the northern Baltic (Bothnian Sea) was compared to the photosynthesis of F. vesiculosus in the Atlantic. Oxygen evolution was measured before and after exposure to 2.3 W of UV-B (280–320 nm) radiation for 5 h, as well as after 48 h recovery in low light. The plants were kept in their own sea water salinity as well as in a changed salinity, this to examine possible correlations between salinity and photosynthesis. The results show a significant higher initial maximal photosynthesis (P _max) for Atlantic plants (10.3 nmol O₂ g⁻¹ FW s⁻¹) compared to Baltic plants (4.0 nmol O₂ g⁻¹ FW s⁻¹). The Baltic plants were found more sensitive to UV-B with a 40–50% decrease of P _max as well as a lower degree of recovery (60–70% compared to 75–95% for the Atlantic plants). The higher salinity (35 psu) had a positive effect on the Baltic F. vesiculosus with increased P _max as well as increased tolerance to UV-B. The lower salinity (5 psu) had a negative effect on the Atlantic plants with a decreased P _max as well as a lower tolerance to UV-B. Pigment content was found higher in Atlantic F. vesiculosus. The pigment content decreased then the Atlantic plants were transferred to 5 psu. The concentration of Chl a as well as the total content of violaxanthin, diadinoxanthin and zeaxanthin in Baltic plants increased when transferred to 35 psu. The Atlantic F. vesiculosus can not survive the low salinity in the northern Baltic (died within 8 weeks). It is likely that a long time acclimation or adaptation to low salinity has taken place for F. vesiculosus in northern Baltic. If this is an ecotypic or genotypic development it is too early to say.     

Seasonal interactive effects of pCO2 and irradiance on the ecophysiology of brown macroalga Fucus vesiculosus L.

ABSTRACT

Stochastic upwelling of seawater in the Baltic Sea from the deep, anoxic bottoms may bring low-pH water rich in CO₂ close to the surface. Such events may become more frequent with climate change and ongoing ocean acidification (OA). Photoautotrophs, such as macroalgae, which are important foundation species, have been proposed to benefit from increased carbon availability due to reduced energetic cost in carbon acquisition. However, the exact effects of CO₂ fertilization may depend on the ambient light environment, as photosynthesis rates depend on available irradiance. In this experimental study, interacting effects of CO₂ addition and irradiance on the habitat-forming macroalga Fucus vesiculosus were investigated during two seasons – winter and summer – in the northern Baltic Sea. Growth rates remained unaffected by CO₂ or irradiance during both seasons, suggesting that direct effects of elevated CO₂ on mature F. vesiculosus are small. Increases in CO₂ affected algal elemental ratios by increasing carbon and decreasing nitrogen content, with resulting changes in the C:N ratio, but only in winter. In summer, chlorophyll a content increased under low irradiance. Increases in CO₂ caused a decline in light-harvesting efficiency (decrease in F_v/F_m and α) under high irradiance in summer, and conversely increased α under low irradiance. High irradiance caused increases in the maximum relative electron transport rate (rETR_max) in summer, but not in winter. Differences between winter and summer indicate that F. vesiculosus responses to CO₂ and irradiance are season-specific. Increases in carbon content during winter could indicate slightly positive effects of CO₂ addition in the long run if the extra carbon gained may be capitalized in growth. The results of this study suggest that increases in CO₂, either through upwelling or OA, may have positive effects on F. vesiculosus, but these effects are probably small.     

The antibacterial effect of a polyhydroxylated fucophlorethol from the marine brown alga,Fucus vesiculosus

An antibacterial compound was isolated from the brown alga Fucus vesiculosus. Purification consisted of extraction of plant material with 0.05% trifluoroacetic acid, concentration on a C₁₈ cartridge, and reverse-phase high performance liquid chromatography on a C₁₈ semi-preparative column. The isolated compound exhibited antibacterial activity against both the Gram-positive and the Gram-negative bacteria tested. Killing studies conducted indicated that the activity was bactericidal. The compound showed no haemolytic effect against human red blood cells. Results obtained by electrospray ionization mass spectrometry indicated that the antibacterial activity was caused by a polyhydroxylated fucophlorethol.     

Present and past depth distribution of bladderwrack (Fucus vesiculosus) in the Baltic Sea

This study aimed to (1) assess the present depth distribution of Fucus vesiculosus in the Baltic Sea and evaluate differences between districts and (2) assess long-term and recent changes in depth distribution and evaluate reasons for such changes. This was done through compilation and analysis of existing data (3356 obs.). Depth limits were shallowest in the Kattegat, the Danish Belts and the Øresund (∼1.5 m on average), located at the entrance of the Baltic Sea and markedly deeper in the central and inner parts of the Baltic (up to ∼4.5 m on average). This increase in depth limits to some extent matched the decline in salinity and may in part be explained by reduced competition when species diversity decreases successively along the Baltic salinity gradient. In the central and inner Baltic Sea, Secchi depths explained part of the variation (16%) in depth limits and the majority (85%) of the variation in maximum attainable depth limits whereas at the entrance of the Baltic Secchi depths explained a negligible part of the variation (∼1%). In most districts, depth limits moved upwards during the 20th century. In many cases this happened during or shortly after the 1960s/1970s, and was most likely due to eutrophication.     

Isolation of protoplasts from Fucus serratus and F. vesiculosus(Fucales, Phaeophyceae): factors affecting protoplast yield

Protoplasts were isolated enzymatically from meristematic tissues of the brown algae, Fucus serratus, using a combination of 2% cellulase R-10 Onozuka, 0.5% macerozyme and 1% crude extract of gland gut of Aplysia vaccaria. The main factors affecting protoplast yield were identified. Protoplasts were produced in large quantities from apical region of thallus and from plantlets compared to mature explants. Yields were greatly improved by the addition of sodium citrate and bovine serum albumin in the enzymatic solution and could reach 5.8 × 10⁶ protoplasts per gram of fresh wt. The applicability of these optimal parameters to other species Fucus vesiculosus was shown.     

Effects of filamentous algae and deposited matter on the survival of Fucus vesiculosus L. germlings in the Baltic Sea

As a result of increased nutrient levels in the Baltic Sea during thepast 50 years, mass developments of filamentous algae have become a commonfeature along the Swedish east coast and deposition of organic matter has alsoincreased. To test whether these two factors have any effects on the early lifestages of Fucus vesiculosus a number of laboratory andfield studies were conducted. The amount of epilithic and epiphytic filamentousalgae on F. vesiculosus and the amount of deposited matterin the littoral zone were quantified during the two reproductive periods ofF. vesiculosus, early summer (May–June) and lateautumn (September–October). Both filamentous algae (Cladophoraglomerata) and deposited matter (introduced either before or aftersettlement of fertilized eggs) were shown to significantly decrease the numberof surviving germlings. The survival of germlings seeded on stones withfilamentous algae, or seeded on culture dishes concurrently with the lowestconcentration of deposited matter (0.1 g dm^–2),was 5% or less. In the field, the amount of filamentous algae was significantlyhigher during F. vesiculosus summer reproduction, whereasthe amount of deposited matter collected in traps was significantly higherduring the period of autumn reproduction. The greatest biomass of filamentousalgae was observed at sheltered sites. Based on the negative effects offilamentous algae and deposited matter on Fucusrecruitmentand the observation of local and seasonal differences in abundance offilamentous algae and deposition, we suggest that the prerequisites for thesurvival of either summer or autumn-reproducing populations of F.vesiculosus in the Baltic Sea may differ locally.     

Long-term decline and recent recovery of Fucus populations along the rocky shores of southeast Sweden, Baltic Sea

The Fucus populations on rocky shores along 300 km of the coastal waters of southeast Sweden in the Baltic proper have been studied since 1984 at a number of fixed sites as part of monitoring programmes. This paper describes changes in distribution and abundance of F. vesiculosus and F. serratus during the period 1984–2001. Sheltered sites showed a consistent temporal and spatial pattern of Fucus spp. distribution over a coastline of 300 kilometres. The depth penetration and abundance of Fucus spp. increased during the 1980s. Around 1990 the development reversed as a consequence of grazing and in 1997 many sites were almost devoid of Fucus spp. Since 1998 both abundance and depth penetration have increased again, possibly as a result of local measures against eutrophication. Exposed sites, on the other hand, lost their Fucus populations at the beginning of the 1990s, and they have not recovered. Extended field studies lead us to deduce that the fixed sites referred to above were representative of the Fucus populations in the area investigated. Major declines, both at sheltered and exposed sites, are attributed to grazing by the isopod Idotea baltica. The populations of I. baltica may have been favoured by the continuing eutrophication of the Baltic, a series of mild winters in the 1990s, and a contemporary decline in some potential predators.     

Reckless males, rational females: dynamic trade-off between food and shelter in the marine isopod Idotea balthica

Habitat choice of herbivores is expected to be a resolution of a trade-off between food and shelter. The resolution of this trade-off may, however, be dynamic within a species because distinct phenotypes may value these factors differently and the value may vary temporally. We studied this hypothesis in the marine herbivore Idotea balthica (Isopoda), by simultaneously manipulating both food and shelter, and investigated whether the resolution of the trade-off differed between sexes, colour morphs and day and night (i.e. high and low predation risk). Isopods chose between exposing and concealing backgrounds in which the quantity or quality of food varied. When choosing between the backgrounds in the absence of food, females preferred the concealment more than males did. However, in a trade-off situation the isopods traded shelter for food, and females more so than males. Thus, males' lower preference for the shelter was not counterbalanced by a stronger preference for food. The microhabitat use also differed between night and day showing adaptation to diurnally fluctuating predation risk. We suggest that microhabitat utilization of females is more strongly tied to variation in risk and resources than that of males, for whom other factors, such as seeking mates, may be more important.     

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.     

Phytoplankton vertical distributions and composition in Baltic Sea cyanobacterial blooms

We studied the vertical structure of the phytoplankton community in two toxic cyanobacterial blooms in the offshore Baltic Sea. In 1994, vertically separated potentially toxic, diazotrophic and mixotrophic species (belonging to Cyanophyceae, Dinophyceae and Prymnesiophyceae) dominated. In 1997, picocyanobacteria, mainly in colonies, made up 40–50% of the total phytoplankton carbon biomass in the top 20 m both day and night. Colony-forming species of picocyanobacteria seem to be occasionally important and hitherto underestimated in the Baltic Sea.We found species-specific depth distribution patterns. Nodularia spumigena and Anabaena spp. were observed mainly above 10 m depth, while Aphanizomenon sp. was mostly found deeper, especially at night. Dinophysis norvegica was only abundant near the seasonal pycnocline and showed very limited diurnal migration. Other flagellates, including small Cryptophyceae and 10 identified Chrysochromulina species, occurred down to 40 m depth. Their vertical migration may help to retrieve nutrients from below the summer pycnocline.We conclude that considerable differences in dominating functional groups may occur between years/bloom stages, and that the vertical distribution pattern of many species is recurring at similar environmental conditions, suggesting species-specific niche-separation.     

Seasonal variability in the grazing potential of the invasive amphipod Gammarus tigrinus and the native amphipod Gammarus salinus (Amphipoda: Crustacea) in the northern Baltic Sea

Mesograzers are known to reduce the biomass of their host plant and modify the structure of the whole macrophyte community in many ecosystems. Thus, the introduction of an efficient mesograzer may destabilize macrophyte community and also affect the native grazers. We estimated how large proportion of macrophyte production are consumed by the alien gammarid G. tigrinus and the native gammarid G. salinus in the species poor ecosystem of the northern Baltic Sea. We analysed whether G. tigrinus consumes different diet as the native G. salinus and whether the effect of G. tigrinus on the survival of the native G. salinus is macrophyte species specific. Grazing experiments showed that there was a clear difference in the grazing rates of gammarids among the studied macrophyte species in summer and autumn but not in spring. The grazing rates were significantly higher in the prevailing macrophyte Pilayella littoralis as compared to other macrophytes. The grazing was inversely related to the diurnal net photosynthetic values of macrophytes. The gammarid amphipods potentially removed only a minor part of plant primary production except for summer and autumn when grazing of a few perennial species exceeded macrophyte production. Macrophyte species and presence of G. salinus had no effect on the survival of G. tigrinus. The presence of G. tigrinus, however, reduced the survival of the native gammarids within P. littoralis in summer. To conclude it is likely that both native and alien gammarid amphipods do not exert significant pressure on the macroalgal communities in the northern Baltic Sea. Competitive interactions between G. tigrinus and G. salinus within the prevailing macrophyte P. littoralis is the likely explanation of the decline of the native gammarid amphipods after the establishment of G. tigrinus in the northern Baltic Sea.     

Ecology of the hybrid Marram Grass x Calammophila baltica in Britain

Variation in growth habit between British populations of Ammophila arenaria, and its intergeneric hybrid, x Calammophila baltica was investigated. Competition between the two species was examined with a de Wit type competition experiment. The morphology of the species was found to be significantly different in terms of flowering, leaf dimensions, and tillering and branching patterns. It was found that they competed for the same space, except when nutrients were added and there was space for rhizome extension. These results imply that when planting for sand dune conservation, both species should be utilized in a mixed sward, with fertilizer added in a mulch.     

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     

Influence of abiotic factors on the structure of the population of the brown alga <Emphasis Type="Italic">Fucus vesiculosus</Emphasis> in East Murman (Barents Sea)

The peculiarities of the structure of the population of the brown alga Fucus vesiculosus in East Murman at different combinations of abiotic factors were studied. Salinity below 10‰ and high surf activity reduced the average life span of the plants (to 1 year) and caused a significant predominance of female specimens (60–90% of the fertile specimens). The proportion of female specimens increased with age. The greatest average age (3 branchings) and the highest survival of F. vesiculosus occurred in the wave-protected parts of bays. The distribution of F. vesiculosus was related to water movement intensity, salinity, and substrate type. The size-weight characteristics of F. vesiculosus thalli were decreased during maximum salinity fluctuations. The optimal conditions for this species were those existing on a weakly protected shore with short-term fluctuations of salinity (24–25‰).