KELP PATCH DYNAMICS IN THE FACE OF INTENSE HERBIVORY: STABILITY OF AGARUM CLATHRATUM (PHAEOPHYTA) STANDS AND ASSOCIATED FLORA ON URCHIN BARRENS1

The brown alga Agarum clathratum (Dumortier) is the only large, perennial, fleshy macrophyte commonly found on urchin‐dominated barrens in the northwestern North Atlantic. We examined the spatial and temporal stability of A. clathratum stands and their impact on algal recruitment in the Mingan Islands, northern Gulf of St. Lawrence. The stands were highly stable in space and time, with only small intersite variations. The percent cover of A. clathratum in 144‐m² areas increased by 6.5%–11.4% over a 2‐year period, and most changes in abundance occurred at the edge of the stands. The surface area of small (<13 m²) single stands of A. clathratum increased by approximately 1.8%·month^?1, although marked increases (>95%) occurred during winter, largely because adjacent stands merged into larger single stands. Mature stands of A. clathratum appear to enhance algal recruitment, as juvenile A. clathratum and the understory red alga Ptilota serrata (Kützing) were orders of magnitude more abundant inside than outside the stands. The experimental removal of the A. clathratum canopy (1‐m² portions) had no long‐term effect on the abundance of A. clathratum, which within 14 months had recolonized most of the cleared areas. In contrast to juvenile A. clathratum, the abundance of P. serrata rapidly decreased after canopy removal. Our results demonstrate that A. clathratum stands are a stable component of urchin barrens in spite of the heavy grazing that typically occurs there. Maintenance and expansion of A. clathratum stands and associated flora appear to depend on positive interactions with self‐defended adult A. clathratum.     

Exposure to ultraviolet radiation delays photosynthetic recovery in Arctic kelp zoospores

Seasonal reproduction in some Arctic Laminariales coincides with increased UV-B radiation due to stratospheric ozone depletion and relatively high water temperatures during polar spring. To find out the capacity to cope with different spectral irradiance, the kinetics of photosynthetic recovery was investigated in zoospores of four Arctic species of the order Laminariales, the kelps Saccorhiza dermatodea, Alaria esculenta, Laminaria digitata, and Laminaria saccharina. The physiology of light harvesting, changes in photosynthetic efficiency and kinetics of photosynthetic recovery were measured by in vivo fluorescence changes of Photosystem II (PSII). Saturation irradiance of freshly released spores showed minimal I _k values (photon fluence rate where initial slope intersects horizontal asymptote of the curve) values ranging from 13 to 18 μmol photons m⁻² s⁻¹ among species collected at different depths, confirming that spores are low-light adapted. Exposure to different radiation spectra consisting of photosynthetically active radiation (PAR; 400–700 nm), PAR+UV-A radiation (UV-A; 320–400 nm), and PAR+ UV-A+UV-B radiation (UV-B; 280–320 nm) showed that the cumulative effects of increasing PAR fluence and the additional effect of UV-A and UV-B radiations on photoinhibition of photosynthesis are species specific. After long exposures, Laminaria saccharina was more sensitive to the different light treatments than the other three species investigated. Kinetics of recovery in zoospores showed a fast phase in S. dermatodea, which indicates a reduction of the photoprotective process while a slow phase in L. saccharina indicates recovery from severe photodamage. This first attempt to study photoinhibition and kinetics of recovery in zoospores showed that zoospores are the stage in the life history of seaweeds most susceptible to light stress and that ultraviolet radiation (UVR) effectively delays photosynthetic recovery. The viability of spores is important on the recruitment of the gametophytic and sporophytic life stages. The impact of UVR on the zoospores is related to the vertical depth distribution of the large sporophytes in the field.     

Population biology of the intertidal kelp, Alaria marginata Postels and Ruprecht: A non-fugitive annual

Persistence of annual plant populations requires sufficient seeds and suitable habitat for development and growth each year. Competition with perennials may prevent within site persistence and result in “fugitive” annual populations. Comparisons have been made between the population biology of annual macroalgae and terrestrial plants, but demographic information necessary to make strong comparisons is lacking for most of these algae, and life history differences may make such comparisons questionable. We studied population dynamics of the kelp Alaria marginata to determine if it was an annual and, if so, how populations persisted. This kelp is the dominant macroalga on exposed mid to low rocky intertidal shores along the Big Sur coast of California. Experimental clearings at two sites were used to assess recruitment timing and survivorship. Sporophytes were collected monthly to determine growth and fecundity. Recruitment occurred in late winter to early spring, primarily on geniculate corallines and residual A. marginata holdfasts. Thinning was inversely related to density, and occurred during the February through July growing season as larger thalli rapidly increased in length (up to 1.4 m month^− 1) and formed a thick canopy. Sorus development was positively related to size, began as early as March, peaked in late August-October, and decreased as adults were removed by winter surf. Spore release was generally highest (10⁸-10⁹ spores individual^− 11 h^− 1) between October and January and associated with high water motion. Survivorship of sporophytes beyond one year was < 1%, showing the populations were annual.Field observations and experiments on effects of canopy clearing, season of clearing, and influence of substrate type on recruitment were done to assess how these annual populations persist. Massive spore production at the onset of fall storms, survival of microscopic stages for 3-4 months facilitated by microhabitat refuges, rapid growth, large size and rapid maturation of sporophytes contributed to persistence. Furthermore, the dense stands with thick canopies may suppress potential competitors via shading and abrasion. Rather than being a fugitive, this combination of growth and life history features enables A. marginata and perhaps other large, annual kelps to maintain perennial populations.     

Isolation and description of eight polymorphic microsatellite loci for the winged-kelp Alaria nana

Here I report on the isolation of eight microsatellites from the winged-kelp, Alaria nana, using an enrichment protocol. Using 55 individuals from Tatoosh, WA and complete data at all loci, loci displayed 6-11 alleles with observed and expected heterozygosities of 0.396-0.754, and 0.769-0.872 respectively. Using an isolated, randomly breeding subset, six of the eight loci were determined to be in Hardy-Weinberg equilibrium. These markers will be useful in assessing population structure in the intertidal and inferring patterns of dispersal.     

Regulation of Sorus Formation by Auxin in Laminariales Sporophyte

Young sporophytes of Laminaria japonica Areshoug were cultured in six indole-acetic acid (IAA) concentrations (0, 10⁻⁸, 10⁻⁷, 10⁻⁶, 10⁻⁵, 10⁻⁴ M) to examine the effect of auxin on growth. The effects of auxin on sorus formation were also examined by using discs taken from the adult sporophyte. The auxin contents and IAA oxidase activities in the thallus and sorus parts of the sporophyte were determined with the blade and sporophyll of other Laminariales plants, Undaria pinnatifida (Harvey) Suringar and Alaria crassifolia Kjellman. The young sporophytes of L. japonica showed highest elongation rate in 10⁻⁵ M IAA. In contrast, the sorus formation on the discs cultured in 10⁻⁵ M IAA was markedly delayed in comparison with other concentrations, indicating that sorus formation was suppressed by IAA. Free and conjugated auxin contents were lower in the reproductive parts than in the vegetative parts. In three Laminariales sporophytes, IAA oxidase activity was about 3–9 times higher in the reproductive parts than in the vegetative parts. Taken together these results suggest that the growth and reproduction of Laminariales sporophytes are regulated by internal auxin levels. Elucidating the regulation mechanism is likely to provide information that is important for the management of plant production and the assessment of the physiological status of plants in the field.     

PHYSIOLOGICAL INVESTIGATIONS ON ALARM ESCULENTA (LAMINARIALES. PHAEOPHYCEAE). IV. INORGANIC AND ORGANIC NITROGEN IN THE BLADE1,2

Seasonal variations in nitrate and organic nitrogen content along the wing and midrib of Alaria esculenta (L.) Grev. lamina have been compared with the NO₃^- cycle in the sea and yearly growth pattern of the blade. Throughout the year, organic N is highest in blade meristem, while NO₃^- distribution is less consistent. NO₃^-in blades reaches a peak in March (ca. 25–28 μM), whereas maximum relative accumulation, 3,300X ambient seawater level, occurs in October. Content of NO₃^- and organic N in the blade decreases in concert with the decline of seawater NO₃^- in April. The three periods of rapid blade growth are not correlated with a specific organic N content in the blade meristem. Laboratory experiments suggest that low NO₃^- and elevated seawater temperature are not the major factors retarding Alaria blade growth during summer and early fall in nature.     

AUXIN,AN ENDOGENOUS REGULATOR OF GROWTH IN ALGAE?1,2

The effects of 5 pairs of synthetic auxins and chemically related, but biologically inactive, derivatives (e.g., 2,4- and 3,5-dichlorophenoxyacetic acid, respectively) on blade growth of the brown alga, Alaria esculenta (L.) Grev.(Laminariales) were studied. The significant result was the inability to divide the growth responses into auxin and non-auxin groups. At high concentrations (10^?3 M) 9 test substances inhibited blade growth; lower concentrations (<10^?3 M) were either slightly inhibitory or showed no effect. The data suggest that a higher plant auxin-specific recognition mechanism is lacking in this alga. As this mechanism is central to higher plant concepts of growth control by auxin, the role of auxin as an endogenous regulator of growth in Alaria, and perhaps other algae as well, seems unlikely. This interpretation is consistent with recent negative biochemical data from auxin binding studies using subcellular preparations from algae.     

Effects of chelated iron on oogenesis and vegetative growth of kelp gametophytes (Phaeophyceae)

The supply of iron has been reported to affect gametogenesis in the gametophytes of some species of kelps (order Laminariales). Spores of the kelps Alaria marginata Postels & Ruprecht, Dictyoneurum californicum Ruprecht, Egregia menziesii (Turner) Areschoug, Laminaria setchellii Silva, and Macrocystis pyrifera (L.) C. Agardh were cultured in enriched seawater with and without added chelated iron (Fe‐ethylenediaminetetraacetate) to determine the effects of iron on oogenesis. All species showed a decrease in oogenesis without added Fe‐ethylenediaminetetraacetic acid (EDTA). Gametophytes of E. menziesii showed predominant gametogenesis with or without supplied iron, resulting in all cells being converted to gametes so that vegetative growth did not continue. Vegetative gametophytes were obtained in the other species used. Gametophytes of M. pyrifera did not show any oogenesis without added Fe‐EDTA, while those of L. setchellii, A. marginata and D. californicum were intermediate in their response, showing some gametogenesis without added Fe‐EDTA. When Fe‐EDTA supply was delayed by 6, 13 and 20 days with spores of M. pyrifera, the gametophytes produced fewer eggs, with a greater decrease as the delay grew longer. A range of Fe‐EDTA concentrations was investigated using isolated female gametophytes of two strains of M. pyrifera and one of Macrocystis integrifolia Bory. None of these three strains produced gametes without the addition of Fe‐EDTA. Gametophytes of M. integrifolia required the least amount of added Fe‐EDTA to achieve gametogenesis while gametophytes of M. pyrifera required higher amounts, with the two strains showing somewhat different responses. Iron nutrition appears to be an essential factor for gametogenesis in several species of kelps.     

New data in France on the trematode Alaria alata (Goeze, 1792) obtained during Trichinella inspections

The trematode Alaria alata is a cosmopolite parasite found in red foxes (Vulpes vulpes), the main definitive host in Europe. In contrast only few data are reported in wild boars (Sus scrofa), a paratenic host. The aim of this paper is to describe the importance and distribution of Alaria alata mesocercariae in wild boars, information is given by findings of these larvae during Trichinella mandatory meat inspection on wild boars' carcasses aimed for human consumption. More than a hundred cases of mesocercariae positive animals are found every year in the East of France. First investigations on the parasite's resistance to deep-freezing in meat are presented in this work.     

INTERACTIONS BETWEEN UV RADIATION AND TEMPERATURE LIMIT INFERENCES FROM SINGLE‐FACTOR EXPERIMENTS1

The global environment is changing. Substantial shifts in temperature, rainfall, cloud cover, and UV radiation (UVR) are all predicted as a result of anthropogenic activity. Although the actual and potential effects of changes in single environmental variables are being studied intensively, the interactive effects of multiple stressors have received little attention. Here we offer the first experimental evidence of interactive effects between UVR and temperature on germination and growth in multicellular organisms. To address the question of how temperature affects survival and growth of organisms in the presence of UVR, we exposed early life stages of two species of intertidal algae, Alaria marginata Postels et Ruprecht and Fucus gardneri Silva, to four levels of UVR at three temperatures for 56 h. PAR and day length (12:12‐h light:dark) were held constant across all treatments. UVR levels bracketed natural levels, and temperatures were within the range of ambient temperatures. Designated endpoints were germination rate and cell number, and we recorded mortality where survival was nil. Our results support the hypothesis that temperature mediates the net biological effect of UVR and vice versa. For instance, spores of A. marginata were able to survive and grow at 15° C at all UV levels and at 10° C in the absence of UVR but were unable to survive at 10° C in the presence of high levels of UVR. Our results suggest that the ability to predict the effects of global change hinges on understanding interactions among environmental variables, imposing strict limits on inferences made from single‐factor experiments.