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.     

Chemical settlement inhibition versus post-settlement mortality as an explanation for differential fouling of two congeneric seaweeds

It has been proposed that seaweed secondary metabolites, e.g. brown algal phlorotannins, may have an ecologically important function as a chemical defence against epiphytes, by acting against colonisation of epiphytic organisms. We tested whether the low epiphytic abundance on the invasive brown seaweed Fucus evanescens, compared to the congeneric F. vesiculosus, is due to a more effective chemical defence against epiphyte colonisation. A field survey of the distribution of the common fouling organism Balanus improvisus (Cirripedia) showed that the abundance was consistently lower on F. evanescens than on F. vesiculosus. However, contrary to expectations, results from experimental studies indicated that F. vesiculosus has a more effective anti-settlement defence than F. evanescens. In settlement experiments with intact fronds of the two Fucus species, both species deterred settlement by barnacle larvae, but settlement was lower on F. vesiculosus both in choice and no-choice experiments. Phlorotannins from F. vesiculosus also had a stronger negative effect on larval settlement and were active at a lower concentration than those from F. evanescens. The results show that Fucus phlorotannins have the potential to inhibit settlement of invertebrate larvae, but that settlement inhibition cannot explain the lower abundance of the barnacle Balanus improvisus on F. evanescens compared to F. vesiculosus. Assessment of barnacle survival in the laboratory and in the field showed that this pattern could instead be attributed to a higher mortality of newly settled barnacles. Observation suggests that the increased mortality was due to detachment of young barnacles from the seaweed surface. This shows that the antifouling mechanism of F. evanescens acts on post-settlement stages of B. improvisus.     

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.     

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.     

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.     

The Influence of Salinity on the Rate of Dark Respiration and Structure of the Cells of Brown Algae Thalli from the Barents Sea Littoral

We studied the changes in the rate of dark respiration (DR) and structure of the cells in Ascophyllum nodosum and Fucus vesiculosus thalli during the incubation at 40, 34, 20, 10, and 2 salinity for 14 days. The changes in salinity affected the rate of DR and the structure of the thallus apical cells: the organelles swollen and were destroyed later. The effect of hyposalinity on the algae was more pronounced as compared to hypersalinity. The stress intensity directly increased with the degree of desalination. Further adaptation of the algae to low salinity enhanced DR and, hence, was an energy-dependent process. Despite higher DR rates (during the stress and adaptation) in F. vesiculosus as compared to A. nodosum, the seaweeds had similar pattern of adaptation to the variation of salinity. Different primary response of the seaweeds to 20 salinity was an important exception; apparently, the salinity around 20 is the limit of these species distribution in desalination zones.     

Polar extracts of the brown alga Fucus vesiculosus (L.) reduce assimilation efficiency but do not deter the herbivorous isopod Idotea baltica (Pallas)

Plant secondary metabolites provide resistance against herbivory if they are capable of decreasing herbivore performance or deterring feeding. Here we test the effects of phlorotannin-containing polar extracts from the brown alga Fucus vesiculosus on the assimilation efficiency and feeding preferences of the herbivorous isopod Idotea baltica. We test the effects separately for each sex of the isopod to test the hypothesis that the digestive efficiencies of the sexes may have diverged. Phlorotannin-containing polar extracts were incorporated into an agar-based artificial food in varying amounts. We also varied the quality of the food, using high-quality food containing animal proteins and low-quality food containing only algal material. The total assimilation efficiency as well as the assimilation efficiencies of carbon and nitrogen decreased with an increasing concentration of phlorotannin-containing extracts. The negative effects on carbon and nitrogen assimilation were higher when extracts were incorporated in the high-quality food. Females were consistently more efficient than males in all assimilation measures; there were no interactions between sex and food quality or phlorotannin concentration, suggesting that the digestive efficiency of the sexes had diverged but that the two sexes responded similarly to phlorotannin enriched artificial food. Although phlorotannin-containing extracts were harmful to performance, they did not deter feeding in preference tests. On the contrary: the isopods preferred the food containing more phlorotannins, suggesting that phlorotannins may act as cues in host recognition. Thus phlorotannins were inefficient deterrents, and I. baltica seems to be adapted to utilizing phlorotannin-containing foods despite their detrimental effects on assimilation. We suggest that F. vesiculosus provides a predation refuge and temporal constancy in availability for I. baltica, both of which select for the ability to utilize it as a food. Phlorotannin-containing polar extracts from F. vesiculosus are still harmful digestibility reducers, and may thereby prevent complete specialization and generate selection for utilization of a wider host range.     

Competitive ranks of three<Emphasis Type="Italic"> Fucus</Emphasis> spp. (Phaeophyta) in laboratory experiments—testing of Keddy's competitive hierarchy model

Keddy's competitive hierarchy model describes species distribution patterns along gradients under equilibrium conditions and can potentially serve as an explanation for zonation patterns of intertidal seaweeds on rocky shores. One of the assumptions of the model is a competitive hierarchy with the top competitor occupying the benign end of the gradient. Another assumption is the consistency of competitive ranks of species in all environmental conditions included in the shared parts of species' fundamental niches. In laboratory replacement series experiments, the competitive ranks of pairs of Fucus species that occupy adjacent zones in the field were analysed and compared to ranks found in previous field experiments. Unattached thalli of Fucus serratus versus F. vesiculosus or F. vesiculosus versus F. spiralis, respectively, were held in aerated beakers to establish the competitive ranking of the three congeners. Each replacement series was conducted at three total densities. F. vesiculosus was clearly competitively dominant over F. serratus. In competition with F. spiralis, F. vesiculosus was only dominant at its lowest absolute input frequencies, but at higher frequencies dominance was reversed. At high densities, the total ranking was F. spiralis > F. vesiculosus > F. serratus, which is the opposite order to that which would be expected from Keddy's model. Although all three species thrived well under the laboratory conditions, the results did not reflect in situ competitive dominances, which may be an effect of nutrient competition in the laboratory. Keddy's assumption that competitive ranks are consistent over the whole range of fundamental niches cannot be supported for Fucus spp. Communicated by K. Lüning     

THE USE OF NEAR INFRARED REFLECTANCE SPECTROMETRY FOR CHARACTERIZATION OF BROWN ALGAL TISSUE1

Measuring qualitative traits of plant tissue is important to understand how plants respond to environmental change and biotic interactions. Near infrared reflectance spectrometry (NIRS) is a cost‐, time‐, and sample‐effective method of measuring chemical components in organic samples commonly used in the agricultural and pharmaceutical industries. To assess the applicability of NIRS to measure the ecologically important tissue traits of carbon, nitrogen, and phlorotannins (secondary metabolites) in brown algae, we developed NIRS calibration models for these constituents in dried Sargassum flavicans (F. K. Mertens) C. Agardh tissue. We then tested if the developed NIRS models could detect changes in the tissue composition of S. flavicans induced by experimental manipulation of temperature and nutrient availability. To develop the NIRS models, we used partial least squares regression to determine the statistical relationship between trait values determined in laboratory assays and the NIRS spectral data of S. flavicans calibration samples. Models with high predictive power were developed for all three constituents that successfully detected changes in carbon, nitrogen, and phlorotannin content in the experimentally manipulated S. flavicans tissue. Phlorotannin content in S. flavicans was inversely related to nitrogen availability, and nitrogen, temperature, and tissue age interacted to significantly affect phlorotannin content, demonstrating the importance of studies that investigate these three variables simultaneously. Given the speed of analysis, accuracy, small tissue requirements, and ability to measure multiple traits simultaneously without consuming the sample tissue, NIRS is a valuable alternative to traditional methods for determining algal tissue traits, especially in studies where tissue is limited.     

Testing for the induction of anti-herbivory defences in four Portuguese macroalgae by direct and water-borne cues of grazing amphipods

Herbivory is a key factor in regulating plant biomass, thereby driving ecosystem performance. Algae have developed multiple adaptations to cope with grazers, including morphological and chemical defences. In a series of experiments we investigated whether several species of macroalgae possess anti-herbivore defences and whether these could be regulated to demand, i.e. grazing events. The potential of direct grazing on defence induction was assessed for two brown (Dictyopteris membranacea, Fucus vesiculosus) and two red seaweeds (Gelidium sesquipedale, Sphaerococcus coronopifolius) from São Rafael and Ria Formosa, Portugal. Bioassays conducted with live algal pieces and agar-based food containing lipophilic algal extracts were used to detect changes in palatability after exposure to amphipod attacks (=treatment phase). Fucus vesiculosus was the only species significantly reducing palatability in response to direct amphipod-attacks. This pattern was observed in live F. vesiculosus pieces and agar-based food containing a lipophilic extract, suggesting that lipophilic compounds produced during the treatment phase were responsible for the repulsion of grazers. Water-borne cues of grazed F. vesiculosus as well as non-grazing amphipods also reduced palatability of neighbouring conspecifics. However, this effect was only observed in live tissues of F. vesiculosus. This study is the first to show that amphipods, like isopods, are capable to induce anti-herbivory defences in F. vesiculosus and that a seasonally variable effectiveness of chemical defences might serve as a dynamic control in alga–herbivore interactions.     

Integrated outdoor culture of two estuarine macroalgae as biofilters for dissolved nutrients from <Emphasis Type="Italic">Sparus auratus</Emphasis> waste waters

An integrated outdoor cultivation of two macroalgal species: Ulva rotundata (Chlorophyta) and Gracilariopsis longissima (Rhodophyta) was designed. The macroalgae were cultured in effluents from an intensive marine culture (growout phase) of gilthead seabream Sparus aurata. The biomass evolution of the algal tanks followed a logistic curve, where the approach to the maximum stocking density of seaweeds was governed by thalli self-shading, as nutrient limitation in the cultivation tank was unlikely. The maximum stocking density of the system was approximately 27.8 g U. rotundata L⁻¹ (16.7 Kg m⁻²) and 11.9 g G. longissima L⁻¹ (7.12 Kg m⁻²). Yield was more than 3 times higher in U. rotundata than in G. longissima. Overall, U. rotundata removed a greater percentage of phosphate (8.9%) and total dissolved inorganic nitrogen (54%) flowing into the algal tanks than G. longissima. The latter species biofiltered approximately 3.2% of phosphate and 17% of the total dissolved inorganic nitrogen input. However, mean nutrient uptake rates on wet weight basis were usually higher in G. longissima than in U. rotundata. The production of total oxidised nitrogen in the algal tanks, considered as being the nitrification rate occurring on the algal fronds by nitrifying bacteria, was less than half of the ammonium uptake by the macroalgae, suggesting that seaweeds competed efficiently for ammonia against the nitrifyers. The biofiltration during a diel cycle showed that mean phosphate biofiltration was lower than 4.5% in the two species whereas ammonium was biofiltered efficiently (up to 67%), especially in U. rotundata. The metal and heavy metal content in the algal tissue at the end of the monitoring period suggested no metal contamination of tissues so that both macroalgal species could be used in the food industry. The study reveals the value of ecological engineering techniques in reducing the dissolved nutrient content in effluents from the fish farm, with the prospect of a better management practises, based on integrated mariculture designs, being developed by the local farmers.     

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

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

Biostimulant activity of brown seaweed species from Strangford Lough: compositional analyses of polysaccharides and bioassay of extracts using mung bean (Vigno mungo L.) and pak choi (Brassica rapa chinensis L.)

The aims of this study were to characterise the composition of seaweed species and to evaluate the efficacy of aqueous extracts as plant biostimulants. Five species (Ascophyllum nodosum, Fucus serratus, Fucus vesiculosus, Laminaria hyperborea and Sargassum muticum) of seaweed were harvested from Strangford Lough, Northern Ireland for the evaluation of polysaccharides, indole-3-acetic acid (IAA), carbon, nitrogen, lipid, ash and mineral contents. The compositional analyses of the five species and their freeze-dried extracts were also carried out using thermogravimetric analysis, scanning electron microscopy and X-ray microanalysis. The concentration of IAA in the acid extracts of the five species ranged between 2.74 and 46.8?ng?g^?1. The carbon, nitrogen, lipid and ash contents ranged between 25.0 and 38.6, 1.37%, and 3.16, 0.83%, and 3.98 and 18.10 and 47.68%, respectively. L. hyperborea and S. muticum contained the highest amounts of minerals. The biostimulant activities of acidic (pH?3.0), neutral (pH?6.5) and alkaline (pH?9.0) extracts were determined by mung bean bioassay. The alkaline extracts from F. vesiculosus and A. nodosum stimulated significantly (P?<?0.001) higher dry matter (DM, %) yield of the mung bean plants. The majority of the acidic extracts significantly (P?<?0.001) enhanced root formation on the mung bean stem cuttings compared to alkaline or neutral extracts. The acidic extracts of the five species, water control and a commercial product were evaluated as foliar feeds for pak choi plants using a hydroponic production system. The interaction of species, e.g. A. nodosum and F. vesiculosus and the two treatment dilutions on DM yield increases of pak choi were significant (P?<?0.05).     

Density dependent grazing effects of the isopod Idotea baltica Pallas on Fucus vesiculosus L in the Baltic Sea

In the Baltic Sea, abiotic factors are often supposed to explain the distribution of the key species Fucus vesiculosus. Still, in many areas, decline of F. vesiculosus has coincided with mass occurrence of the herbivorous isopod Idotea baltica. The aim of this work was to examine whether, how and at what densities I. baltica can affect the distribution of F. vesiculosus in the central Baltic proper. Both large-scale field surveys and a two-week grazing experiment have been performed.In the field survey there was a correlation between density of I. baltica and reduction in depth penetration of F. vesiculosus. At 80 animals per 100 g F. vesiculosus wet weight, the depth penetration of the F. vesiculosus belt was reduced by 2.5 m within a year. In the grazing experiment there was a correlation between density of I. baltica and loss of F. vesiculosus biomass and meristems. In the controls biomass and number of meristems increased by 50%, while at 20 animals per 100 g of F. vesiculosus there was no net growth of F. vesiculosus. Intensity of grazing did not differ between isopod densities of 20, 40 and 60/100 g. At isopod densities of 80 and 100/100 g though, biomass and meristems decreased by 50%, indicating a threshold for the survival of F. vesiculosus in the experiment. At all densities the isopods preferred younger tissue to older.Our results indicate that grazing by Idotea baltica is an important structuring factor in the Baltic Fucus vesiculosus populations.     

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.     

The harvesting of macroalgae in New Zealand

Several species of algae have been commercially harvested in New Zealand, mainly for extraction of agar and alginates. In the past, the harvest was comprised mostly of shore-cast plants. There has been more recent interest, however, in harvesting attached plants of Pterocladia spp., Porphyra spp., Gracilaria sordida, Durvillaea spp., Macrocystis pyrifera, and Ecklonia radiata. The ecological effects of harvesting attached algae depend largely on the sizes of plants, the season of removal, the patch size of clearances, and the proximity and identity of mature plants. These have not been well-studied for seaweeds in New Zealand, but population and life history studies indicate that harvesting methods affect the continuity of algal resources, at least on a local scale, and are crucial factors in their management.     

Functional properties of photosynthetic apparatus in three <Emphasis Type="Italic">Fucus</Emphasis> species inhabiting the White Sea: Effect of dehydration

Three species of Fucus inhabiting different zones at the littoral and sublittoral of the White Sea (F. vesiculosus L., F. serratus L., and F. distichus L.) were compared with regard to performance of the photosynthetic apparatus (PSA) and PSA changes upon prolonged desiccation of algae. The content of chlorophyll a and the total content of carotenoids were significantly higher in F. serratus than in F. vesiculosus. The potential quantum yield (F _v/F _m) of photosystem II (PSII) was 0.69–0.74 in all species. The highest effective quantum yield of PSII (Y _II) was observed in F. vesiculosus plants inhabiting zones with the most intense insolation. The ratio of non-photochemical quenching to photochemical quenching (qN/qP) increased with the depth of algal habitats in the row from F. vesiculosus to F. serratus. The fluorescence parameters F. serratus deviated during desiccation from their normal values and did not recover upon the subsequent return to water. By contrast, the fluorescence parameters of F. vesiculosus and F. distichus recovered gradually after the return of algae from air to sea water. It is supposed that F. serratus plants, unlike other species, are characterized by comparatively low physiological plasticity.     

Seasonal patterns of acid fluctuations and resource storage in the arborescent cactus Opuntia excelsa in relation to light availability and size

Summary We investigated relationships between light availability, diel acid fluctuation, and resource storage in the arborescent cactus Opuntia excelsa growing in western Mexico. We compared canopy and understory individuals from a deciduous forest as well as open-grown plants of the same approximate size as those in the understory. During the wet season light availability and daily fluctuations in titratable acidity (an index of carbon uptake) were lower in the understory than in unshaded habitats. In the dry season all plants had reduced levels of acid fluctuation, with the smallest individuals, regardless of habitat, showing the greatest reduction. These data suggest that light availability in the forest understory constrains carbon assimilation during the wet season, but that a factor associated with plant size, possibly water status, limits carbon gain during the dry season. Plants in all habitats remained physiologically active for at least five months into the dry season. We suggest that this was possible due to the maintenance of constant concentrations of water and nitrogen in the photosynthetically active chlorenchyma. Parenchyma in terminal cladodes showed a different seasonal pattern of resource storage; water content and nitrogen concentration were reduced from the wet to the dry season in the parenchyma. Using the parenchyma to supply photosynthetic tissues during times of reduced resource availability allows O. excelsa to assimilate carbon during times of the year when most other trees in the forest are leafless.