Christian Wiencke

The genus Laminaria sensu lato : recent insights and developments

This review about the genus Laminaria sensu lato summarizes the extensive literature that has been published since the overview of the genus given by Kain in 1979. The recent proposal to divide the genus into the two genera Laminaria and Saccharina is acknowledged, but the published data are discussed under a ‘sensu lato’ concept, introduced here. This includes all species which have been considered to be ‘Laminaria’ before the division of the genus. In detail, after an introduction the review covers recent insights into phylogeny and taxonomy, and discusses morphotypes, ecotypes, population genetics and demography. It describes growth and photosynthetic performance of sporophytes with special paragraphs on the regulation of sporogenesis, regulation by endogenous rhythms, nutrient metabolism, storage products, and salinity tolerance. The biology of microstages is discussed separately. The ecology of these kelps is described with a focus on stress defence against abiotic and biotic factors and the role of Laminaria as habitat, its trophic interactions and its competition is discussed. Finally, recent developments in aquaculture are summarized. In conclusion to each section, as a perspective and guide to future research, we draw attention to the remaining gaps in the knowledge about the genus and kelps in general.

Interactive effects of radiation, temperature and salinity on different life history stages of the Arctic kelp Alaria esculenta (Phaeophyceae)

To estimate the potential effects of climate change on polar marine macroalgae, studies on interactive stress effects of multiple climate-related parameters are essential. Interactions of temperature, radiation and salinity on two different life history stages of Alaria esculenta (L.) Greville from the Kongsfjord (Spitsbergen) were investigated for the first time within this study. Adult macroscopic sporophytes of A. esculenta were exposed to different temperatures between 4 and 21°C combined with artificial irradiation conditions [photosynthetically active radiation, ultraviolet (UV) radiation: UV-A/UV-B, first experiment] and with different salinities [34, 28, 20 practical salinity units (p.s.u.)¸ second experiment]. Effects of photosynthetic activity were determined by measuring variable chlorophyll fluorescence of photosystem II. Germination success of young microscopic zoospores of A. esculenta was studied under multifactorial stress. Zoospore suspensions were exposed to the three different salinities and irradiation conditions at four temperatures between 2 and 16°C. Overall, A. esculenta exhibited a highly stage-specific susceptibility towards the experimental treatments. In both experiments using sporophytes, photosynthetic activity showed significant temperature effects and only very few significant radiation and salinity effects. Microscopic stages of A. esculenta were shown to be more sensitive than the adult macroscopic stages, since germination capacity of zoospores was significantly affected by temperature and salinity changes, and interactions of both. These results suggest that multiple stress factors interact synergistically. Temperature seems to be a predominant environmental parameter for the kelp A. esculenta. Overall, A. esculenta proved to be relatively tolerant and adaptable to increasing temperature and UV radiation, as well as to diluted salinities, but only up to a specific limit.

Thallus morphology and optical characteristics affect growth and DNA damage by UV radiation in juvenile Arctic Laminaria sporophytes

Growth of young sporophytes of the brown algae Laminaria digitata, L. saccharina and L. solidungula from Spitsbergen were measured in the laboratory after being exposed for 21xa0days to either photosynthetically active radiation (PAR=P) or to full light spectrum (PAR + UV-A + UV-B=PAB) using of cutoff glass filters. The plants were grown at 8±2°C and 16xa0h light : 8xa0h dark cycles with 6xa0h additional ultraviolet radiation (UVR) exposure in the middle of the light period. Growth was measured every 10xa0min using growth chambers with online video measuring technique. Tissue morphology and absorption spectra were measured in untreated young sporophytes while chlorophyll (Chl) a content and DNA damage were measured in treated thalli at the end of the experiment. In all species, growth rates were significantly higher in sporophytes exposed to P alone compared to sporophytes exposed to PAB. Tissue DNA damage is dependent on thallus thickness and absorption spectra characteristics of pigments and UV-absorbing compounds. In sporophytes exposed to UVR, energy demands for repair of DNA damage and synthesis of UV-absorbing compounds for protection effectively diverts photosynthate at the expense of growth. Photosynthetic pigment was not significantly different between treatments suggesting a capacity for acclimation to moderate UVR fluence. The general growth pattern in sporophytes exposed to P alone showed an increasing growth rate from the onset of light (0500–0900xa0hours) to a peak at the middle of the light phase (0900–1500xa0hours), a decline towards the end of the light phase (1500–2100xa0hours) and a minimum “low” growth in the dark (2100–0500xa0hours) relative to growth during the entire light phase. Under PAB, different growth patterns were observed such as growth compensation at night in L. digitata, delayed growth recovery in L. saccharina and minimal but continuous growth in L. solidungula. Growth as an integrative parameter of all physiological processes showed that the effect of UVR is correlated to the depth distribution of these species.

Effects of ultraviolet radiation and temperature on the ultrastructure of zoospores of the brown macroalga Laminaria hyperborea

The interactive effects of an 8 h exposure to UV radiation and altered temperatures on the ultrastructure and germination of zoospores of the sublittoral brown alga Laminaria hyperborea (Gunn.) Foslie were investigated for the first time. Spores were exposed to four temperatures (2, 7, 12 and 17 degrees C) and three light regimes (PAR, PAR + UV-A, PAR + UV-A+UV-B). Freshly-released spores of L. hyperborea lack a cell wall and contain a nucleus with fine granular nucleoplasm and a nucleolus, one chloroplast, several mitochondria, dictyosomes and an endoplasmatic reticulum. Further, several kinds of so-called adhesive vesicles, lipid globuli and physodes containing UV-absorbing phlorotannins are embedded in the cytoplasm. No eye-spot is present. Physodes were found but they were rare and small. After an 8 h exposure to UV-B, the nucleoplasm had a mottled structure, chloroplasts contained plastoglobuli, the structure of the mitochondria changed from crista- to sacculus-type and germination was strongly inhibited at all temperatures. UV-A only had an impact on the ultrastructure at the highest temperature tested. The strongest effects were found at 17 degrees C, where germination was reduced to 35%, 32% and 9% after exposure to PAR, PAR+UV-A and PAR + UV-A + UV-B, respectively. This study indicates that UV-B radiation has strong damaging effects on the physiology and ultrastructure of zoospores of L. hyperborea. The results are important for developing scenarios for the effect of enhanced UV radiation and increasing temperatures caused by global climate changes.

Effects of water temperatures, UV radiation and low vs high PAR on phlorotannin content and germination in zoospores of Saccorhiza dermatodea (Tilopteridales, Phaeophyceae)

Steinhoff F.S., Wiencke C., Wuttke S. and Bischof K. 2011. Effects of water temperatures, UV radiation and low vs high PAR on phlorotannin content and germination in zoospores of Saccorhiza dermatodea (Tilopteridales, Phaeophyceae). Phycologia 50: 256–263. DOI: 10.2216/09-107.1 Global climate change will have multiple effects on our environment and might especially change marine coastal ecosystems and their communities due to rising water temperatures and changing light regimes in the water column. Due to the key role of zoospores in the life-history cycle of kelps, we investigated how zoospores of the shallow water species Saccorhiza dermatodea (B. de la Pylaie) J.E. Areschoug respond to interactive effects of changing water temperatures, low/high photosynthetically active radiation (PAR) and UV radiation (UVR). Zoospores were examined with respect to germination pattern (germination rates, light micrographs) and phlorotannin content (Folin–Ciocalteu method). In summer 2007, we exposed S. dermatodea zoospores, obtained from fertile thalli in Kongsfjorden (Svalbard, Norway), to low PAR laboratory [UV-B∶UV-A∶PAR (W m−2) 8∶114∶100; 2–17°C] and high PAR outdoor experimental conditions [UV-B∶UV-A∶PAR (W m−2) 0.1∶5∶100; 7°C] for 8 h. Under low PAR+UVR conditions and at 2–12°C, almost 98% of zoospores germinated in all light treatments. The germination rate was reduced under PAR+UV-A+UV-B only in the 17°C and 7°C outdoor treatment, to 21.4% and 9.6%, respectively. Light microscopic investigations showed differences in germling appearance, growth and pigmentation in the outdoor treatment and at 17°C low PAR conditions, compared with all other treatments, after an exposure time of 8 h. However, phlorotannin content within zoospores remained unaffected by changing UV and low/high PAR conditions. These results suggest that combined effects of present water temperatures, high PAR and UVR can exhibit similar effects on germination as the combined effects of unnatural high temperatures (17°C), low PAR and UVR. With respect to global climate change, the ecological implications of the present study illustrate that zoospores of S. dermatodea are able to survive enhanced water temperatures up to 12°C. Moreover, high PAR in combination with UVR might potentially affect zoospores of S. dermatodea stronger than expected in clear waters.

UV-susceptibility of zoospores of the brown macroalga Laminaria digitata from Spitsbergen

The UV-susceptibility of zoospores of the lower sublittoral kelp Laminaria digitata was studied in the laboratory under varying fluence of spectral irradiance consisting of photosynthetically active radiation (PAR, 400–700xa0nm;xa0=xa0P), PARxa0+xa0UV-A radiation (UV-A, 320–400xa0nm;xa0=xa0PA), and PARxa0+xa0UV-Axa0+xa0UV-B radiation (UV-B, 280–320xa0nm;xa0=xa0PAB). In vivo absorption of phlorotannin, localisation of phlorotannin-containing physodes, structural changes, DNA damage and repair, photosynthesis and germination of zoospores were measured after exposure treatments and after 2–6xa0days of recovery in dim white light. Photodegradation of phlorotannins was observed after extended exposure to ultraviolet radiation (UVR). The UV-protective function of extra- and intracellular phlorotannins was, therefore, observed only after 8xa0h, but not after 16-h UVR exposure. The energetic cost of photoprotection may have caused the delay in ontogenic development of zoospores after 8-h exposure to PA and PAB treatment; longer exposure time corresponding to 16-h PA and PAB treatment eventually lead to cell degeneration at 6xa0days post-cultivation. The formation of cyclobutane–pyrimidine dimers (CPDs), as indicator of DNA damage, was not blocked by the UV-absorbing phlorotannins during the 16-h PAB exposure and the inability for DNA damage repair was likely responsible for low photosynthetic recovery and spore mortality. The higher sensitivity of L. digitata zoospores to UVR compared to other kelps such as Saccorhiza dermatodea and Alaria esculenta confirmed our hypothesis that the depth distribution of adult sporophytes in the field correlates to the sensitivity of their corresponding early life history stages to different stress factors in general and UVR in particular.