Marine heatwaves and decreased light availability interact to erode the ecophysiological performance of habitat-forming kelp species

Coastal marine ecosystems are threatened by a range of anthropogenic stressors, operating at global, local, and temporal scales. We investigated the impact of marine heatwaves (MHWs) combined with decreased light availability over two seasons on the ecophysiological responses of three kelp species (Laminaria digitata, L. hyperborea, and L. ochroleuca). These species function as important habitat-forming foundation organisms in the northeast Atlantic and have distinct but overlapping latitudinal distributions and thermal niches. Under low-light conditions, summertime MHWs induced significant declines in biomass, blade surface area, and Fv/Fm values (a measure of photosynthetic efficiency) in the cool-water kelps L. digitata and L. hyperborea, albeit to varying degrees. Under high-light conditions, all species were largely resistant to simulated MHW activity. In springtime, MHWs had minimal impacts and in some cases promoted kelp performance, while reduced light availability resulted in lower growth rates. While some species were negatively affected by summer MHWs under low-light conditions (particularly L. digitata), they were generally resilient to MHWs under high-light conditions. As such, maintaining good environmental quality and water clarity may increase resilience of populations to summertime MHWs. Our study informs predictions of how habitat-forming foundation kelp species will be affected by interacting, concurrent stressors, typical of compound events that are intensifying under anthropogenic climate change.     

Heat stress responses and population genetics of the kelp Laminaria digitata (Phaeophyceae) across latitudes reveal differentiation among North Atlantic populations

To understand the thermal plasticity of a coastal foundation species across its latitudinal distribution, we assess physiological responses to high temperature stress in the kelp Laminaria digitata in combination with population genetic characteristics and relate heat resilience to genetic features and phylogeography. We hypothesize that populations from Arctic and cold‐temperate locations are less heat resilient than populations from warm distributional edges. Using meristems of natural L. digitata populations from six locations ranging between Kongsfjorden, Spitsbergen (79°N), and Quiberon, France (47°N), we performed a common‐garden heat stress experiment applying 15°C to 23°C over eight days. We assessed growth, photosynthetic quantum yield, carbon and nitrogen storage, and xanthophyll pigment contents as response traits. Population connectivity and genetic diversity were analyzed with microsatellite markers. Results from the heat stress experiment suggest that the upper temperature limit of L. digitata is nearly identical across its distribution range, but subtle differences in growth and stress responses were revealed for three populations from the species’ ecological range margins. Two populations at the species’ warm distribution limit showed higher temperature tolerance compared to other populations in growth at 19°C and recovery from 21°C (Quiberon, France), and photosynthetic quantum yield and xanthophyll pigment responses at 23°C (Helgoland, Germany). In L. digitata from the northernmost population (Spitsbergen, Norway), quantum yield indicated the highest heat sensitivity. Microsatellite genotyping revealed all sampled populations to be genetically distinct, with a strong hierarchical structure between southern and northern clades. Genetic diversity was lowest in the isolated population of the North Sea island of Helgoland and highest in Roscoff in the English Channel. All together, these results support the hypothesis of moderate local differentiation across L. digitata's European distribution, whereas effects are likely too weak to ameliorate the species’ capacity to withstand ocean warming and marine heatwaves at the southern range edge.     

Contrasting genetic diversity patterns in two sister kelp species co‐distributed along the coast of Brittany,France

We investigated patterns of genetic structure in two sister kelp species to explore how distribution width along the shore, zonation, latitudinal distribution and historical factors contribute to contrasting patterns of genetic diversity. We implemented a hierarchical sampling scheme to compare patterns of genetic diversity and structure in these two kelp species co‐distributed along the coasts of Brittany (France) using a total of 12 microsatellites, nine for Laminaria hyperborea and 11 for Laminaria digitata, of which eight amplified in both species. The genetic diversity and connectivity of L. hyperborea populations were greater than those of L. digitata populations in accordance with the larger cross‐shore distribution width along the coast and the greater depth occupied by L. hyperborea populations in contrast to L. digitata populations. In addition, marginal populations showed reduced genetic diversity and connectivity, which erased isolation‐by‐distance patterns in both species. As L. digitata encounters its southern range limit in southern Brittany (SBr) while L. hyperborea extends down to mid‐Portugal, it was possible to distinguish the effect of habitat continuity from range edge effects. We found that L. digitata did not harbour high regional diversity at its southern edge, as expected in a typical rear edge, suggesting that refuges from the last glacial maximum for L. digitata were probably not located in SBr, but most likely further north. For both species, the highest levels of genetic diversity were found in the Iroise Sea and Morlaix Bay, the two regions in which they are being currently harvested. Preserving genetic diversity of these two foundation species in these areas should, thus, be a priority for the management of this resource in Brittany.     

Intra-Annual Variability in Responses of a Canopy Forming Kelp to Cumulative Low Tide Heat Stress: Implications for Populations at the Trailing Range Edge

Anthropogenic climate change is driving the redistribution of species at a global scale. For marine species, populations at trailing edges often live very close to their upper thermal limits and, as such, poleward range contractions are one of the most pervasive effects of ongoing and predicted warming. However, the mechanics of processes driving such contractions are poorly understood. Here, we examined the response of the habitat forming kelp, Laminaria digitata, to realistic terrestrial heatwave simulations akin to those experienced by intertidal populations persisting at the trailing range edge in the northeast Atlantic (SW England). We conducted experiments in both spring and autumn to determine temporal variability in the effects of heatwaves. In spring, heatwave scenarios caused minimal stress to L. digitata but in autumn all scenarios tested resulted in tissue being nonviable by the end of each assay. The effects of heatwave scenarios were only apparent after consecutive exposures, indicating erosion of resilience over time. Monthly field surveys corroborated experimental evidence as the prevalence of bleaching (an indication of physiological stress and tissue damage) in natural populations was greatest in autumn and early winter. Overall, our data showed that L. digitata populations in SW England persist close to their upper physiological limits for emersion stress in autumn. As the intensity of extreme warming events is likely to increase with anthropogenic climate change, thermal conditions experienced during periods of emersion will soon exceed physiological thresholds and will likely induce widespread mortality and consequent changes at the population level.     

Ecophysiological plasticity of annual populations of giant kelp (Macrocystis pyrifera) in a seasonally variable coastal environment in the Northern Patagonian Inner Seas of Southern Chile

Annual populations of Macrocystis pyrifera in Southern Chile have been the main focus of studies intending to understand how these populations can couple consecutive sporophytic generations. Research has included studying the population dynamics and gametophytic responses to environmental conditions and the role of recruitment, grazing, and the use of benthic filter feeders as secondary substrate. Adult sporophytes undergo senescence due to changes in abiotic factors during summer and autumn producing 100 % mortality. This study provides evidence about the environmental factors driving the decline in sporophyte populations occurring in summer and fall by monitoring two independent kelp populations and also by running experiments using 400 L tubular photobioreactors with semicontrolled environmental factors for testing the capacity for new recruits to recover population levels under winter conditions. The study of natural populations of giant kelp indicates that high temperatures (>15–17 °C) explain the high mortality of adult plants in summer. On the other hand, the sporophytes established in late winter/early spring are able, under high nitrogen availability, to increase their chlorophyll content significantly, allowing the individuals to reduce their light saturation point and thus allow a higher productivity under the low light conditions that exist in late winter and early spring. These results, in addition to the recruitment facilitation produced by filter feeders, help to explain how giant kelp can deal with, and couple sporophytic generations, in variable environments. These results also emphasize the highly plastic physiology of giant kelp that enables this species to colonize diverse habitats across its large distributional range.     

The effects of warming on the ecophysiology of two co-existing kelp species with contrasting distributions

The northeast Atlantic has warmed significantly since the early 1980s, leading to shifts in species distributions and changes in the structure and functioning of communities and ecosystems. This study investigated the effects of increased temperature on two co-existing habitat-forming kelps: Laminaria digitata, a northern boreal species, and Laminaria ochroleuca, a southern Lusitanian species, to shed light on mechanisms underpinning responses of trailing and leading edge populations to warming. Kelp sporophytes collected from southwest United Kingdom were maintained under 3 treatments: ambient temperature (12 °C), +3 °C (15 °C) and +6 °C (18 °C) for 16 days. At higher temperatures, L. digitata showed a decline in growth rates and Fv/Fm, an increase in chemical defence production and a decrease in palatability. In contrast, L. ochroleuca demonstrated superior growth and photosynthesis at temperatures higher than current ambient levels, and was more heavily grazed. Whilst the observed decreased palatability of L. digitata held at higher temperatures could reduce top-down pressure on marginal populations, field observations of grazer densities suggest that this may be unimportant within the study system. Overall, our study suggests that shifts in trailing edge populations will be primarily driven by ecophysiological responses to high temperatures experienced during current and predicted thermal maxima, and although compensatory mechanisms may reduce top-down pressure on marginal populations, this is unlikely to be important within the current biogeographical context. Better understanding of the mechanisms underpinning climate-driven range shifts is important for habitat-forming species like kelps, which provide organic matter, create biogenic structure and alter environmental conditions for associated communities.     

Waterborne signaling primes the expression of elicitor-induced genes and buffers the oxidative responses in the brown alga Laminaria digitata

As marine sessile organisms, seaweeds must respond efficiently to biotic and abiotic challenges in their natural environment to reduce the fitness consequences of wounds and oxidative stress. This study explores the early steps of the defense responses of a large marine brown alga (the tangle kelp Laminaria digitata) and investigates its ability to transmit a warning message to neighboring conspecifics. We compared the early responses to elicitation with oligoguluronates in laboratory-grown and harvested wild individuals of L. digitata. We followed the release of H₂O₂ and the concomitant production of volatile organic compounds. We also monitored the kinetics of expression of defense-related genes following the oxidative burst. Laboratory-grown algae were transplanted in kelp habitats to further evaluate their responses to elicitation after a transient immersion in natural seawater. In addition, a novel conditioning procedure was established to mimic field conditions in the laboratory. Our experiments showed that L. digitata integrates waterborne cues present in the kelp bed and/or released from elicited neighboring plants. Indeed, the exposure to elicited conspecifics changes the patterns of oxidative burst and volatile emissions and potentiates this kelp for faster induction of genes specifically regulated in response to oligoguluronates. Thus, waterborne signals shape the elicitor-induced responses of kelps through a yet unknown mechanism reminiscent of priming in land plants.     

Physiological and morphological short-term responses to light and temperature in two Nothofagus species of Patagonia, South America

The study of plant responses to environmental stress factors is essential for management of plant systems and for anticipating their response to climate change. The main goal of this study was to determine morphological and physiological responses of Nothofagus obliqua and N. nervosa seedlings to light and temperature, two of the main stress factors acting in their current natural distribution in NW Patagonia. Responses to light were evaluated analyzing growth and survival, as well as morphological and physiological traits related to them, in seedlings subjected to three contrasting light conditions (full-sun conditions, 50% of sunlight and 20% of sunlight) during one growth season. Temperature photosynthetic responses were evaluated in seedlings subjected to temperature treatments between ?5 and 40°C for 2 and 4 h. Growth rate and biomass partition were similar between light treatments in both species. High apical meristem damage and decreased photosynthetic capacity of preformed leaves were observed under full-sun conditions, suggesting that high light levels have a deleterious effect on plant yield. Both species produced neoformed leaves during the growing season with better photosynthetic capacity than preformed leaves under full sun conditions, contributing to plant acclimation. Almost no plasticity was observed in morphological traits in response to shade. Both species differed in optimum temperature for photosynthesis, with a wider temperature range at which high photosynthesis is maintained in N. obliqua. In both species the higher values of net photosynthetic rate were found at higher temperatures than the mean annual temperature of its current natural distribution range. Under no water-stress conditions, future higher temperatures could increase carbon fixation of these species, with a little advantage of N. obliqua if temperature variance is high. Synergy effect of various environmental stress factors, particularly considering cultivation of these species outside their current natural distribution sites require further studies.     

Physiological studies of subtidal red algae

The net photosynthesis of the subtidal red algae Euthora cristata (C. Agardh) J. Agardh, Phycodrys rubens (L. ) Batters, Phyllophora truncata (Pallas) Newroth et Taylor and Ptilota serrata Kützing were determined under a variety of different light and temperature regimes. The optimal light requirements for net photosynthesis of the species are relatively low, mostly ranging from 465 to 747 ft-c at 5 ° and 15 °C. Seasonal and spatial differences were found in the photosynthesis-light responses of Phyllophora truncata and Ptilota serrata; winter plants exhibited lower light optima for net photosynthesis than spring plants. Deep-water populations of Ptilota showed lower light optima and reduced net photosynthesis as compared with shallow subtidal populations. Summer plants of Euthora cristata, Phycodrys rubens and Phyllophora truncata showed a greater tolerance to high temperatures and higher temperature optima than winter plants. It is suggested that optimal temperature and light requirements of seaweeds are adjusted in an adaptive fashion to the environmental regimes of their habitats. The temperature requirements of the four species are discussed in relation to their local estuarine distributions in New England; eurythermal species have the widest estuarine distributions. Cystocarpic and tetrasporic plants of Euthora cristata and Ptilota serrata show differential physiological responses and vertical distributions. The significance of higher rates of net photosynthesis and lower light optima are discussed in relation to vertical stratification of different generations.     

EFFECTS OF LIGHT AND TEMPERATURE ON NET PHOTOSYNTHESIS AND DARK RESPIRATION OF GAMETOPHYTES AND EMBRYONIC SPOROPHYTES OF MACROCYSTIS PYRIFERA1

The rates of net photosynthesis as a function of irradiance and temperature were determined for gametophytes and embryonic sporophytes of the kelp, Macrocystis pyrifera (L.) C. Ag. Gametophytes exhibited higher net photosynthetic rates based on oxygen and pH measurements than their derived embryonic sporophytes, but reached light saturation at comparable irradiance levels. The net photosynthesis of gametophytes reached a maximum of 66.4 mg O₂ g dry wt^?1 h^?1 (86.5 mg CO₂ g dry wt^?1 h^?1), a value approximately seven times the rate reported previously for the adult sporophyte blades. Gametophytes were light saturated at 70 μE m^?2 s^?1 and exhibited a significant decline in photosynthetic performance at irradiances 140 μE m^?1 s^?1. Embryonic sporophytes revealed a maximum photosynthetic capacity of 20.6 mg O₂ g dry wt^?1 h^?1 (25.3 mg CO₂ g dry wt^?1 h^?1), a rate about twice that reported for adult sporophyte blades. Embryonic sporophytes also became light saturated at 70 μE m^?2 s^?1, but unlike their parental gametophytes, failed to exhibit lesser photosynthetic rates at the highest irradiance levels studied; light compensation occurred at 2.8 μE m^?2 s^?1. Light-saturated net photosynthetic rates of gametophytes and embryonic sporophytes varied significantly with temperature. Gametophytes exhibited maximal photosynthesis at 15° to 20° C, whereas embryonic sporophytes maintained comparable rates between 10° and 20° C. Both gametophytes and embryonic sporophytes declined in photosynthetic capacity at 30° C. Dark respiration of gametophytes was uniform from 10° to 25° C, but increased six-fold at 30° C; the rates for embryonic sporophytes were comparable over the entire range of temperatures examined. The broader light and temperature tolerances of the embryonic sporophytes suggest that this stage in the life history of M. pyrifera is well suited for the subtidal benthic environment and for the conditions in the upper levels of the water column.     

Phenotypic plasticity in Philonotis fontana (Bryopsida: Bartramiaceae)

Abstract

Gametophytes from six populations of the moss Philonotis fontana (Hedw.) Brid. were grown under two light and two water regimes in order to assess the effects of these environmental factors on gametophytic architecture and leaf and leaf-cell dimensions. Both light and water affectedgrowth, but the light treatments had a greater effect, and on more characters, than did the water treatments. Significant population effects under common garden conditions point to genetic variation for several traits, and population × environmental treatment interactions demonstratedgenetic variation for patterns of phenotypic plasticity, i.e. plants differed in their 'norms of reaction'. Variation among populations in leaf dimensions tended to have a strong genetic component (20–30% of the total variation), whereas cell dimensions showed relatively little geneticvariation (<10% of the total).     

Loss of a chloroplast encoded function could influence species range in kelp

Kelps are important providers and constituents of marine ecological niches, the coastal kelp forests. Kelp species have differing distribution ranges, but mainly thrive in temperate and arctic regions. Although the principal factors determining biogeographic distribution ranges are known, genomics could provide additional answers to this question. We sequenced DNA from two Laminaria species with contrasting distribution ranges, Laminaria digitata and Laminaria solidungula. Laminaria digitata is found in the Northern Atlantic with a southern boundary in Brittany (France) or Massachusetts (USA) and a northern boundary in the Arctic, whereas L. solidungula is endemic to the Arctic only. From the raw reads of DNA, we reconstructed both chloroplast genomes and annotated them. A concatenated data set of all available brown algae chloroplast sequences was used for the calculation of a robust phylogeny, and sequence variations were analyzed. The two Laminaria chloroplast genomes are collinear to previously analyzed kelp chloroplast genomes with important exceptions. Rearrangements at the inverted repeat regions led to the pseudogenization of ycf37 in L. solidungula, a gene possibly required under high light conditions. This defunct gene might be one of the reasons why the habitat range of L. solidungula is restricted to lowlight sublittoral sites in the Arctic. The inheritance pattern of single nucleotide polymorphisms suggests incomplete lineage sorting of chloroplast genomes in kelp species. Our analysis of kelp chloroplast genomes shows that not only evolutionary information could be gleaned from sequence data. Concomitantly, those sequences can also tell us something about the ecological conditions which are required for species well‐being.     

Examining the bank of microscopic stages in kelps using culturing and barcoding

This paper describes a method to study the diversity of young kelp sporophytes that are recruited from the bank of microscopic stages. Small samples of rocky substratum (0.5 cm²) were collected from the low intertidal zone, which was dominated by the kelp Laminaria digitata. Samples were cultivated in the laboratory under conditions permitting gametogenesis. Sporophyte recruits in the cultures were isolated and identified at the species level using the barcoding mitochondrial marker rpl31–rns. Sixty per cent of the collected samples had at least one to a maximum of 30 kelp recruits, belonging to five different species (L. digitata, L. hyperborea, L. ochroleuca, Saccharina latissima and Sacchorhiza polyschides). As the examination of freshly collected rocky samples under a stereo microscope did not reveal any kelp sporophytes, the recruitment in these samples after culture probably occurred from the bank of microscopic forms present on the substratum. Despite the dominance of L. digitata in the field, the young sporophytes obtained after culturing were mainly S. polyschides. This study illustrates the suitability of culturing in combination with molecular identification of young sporophytes to address several key aspects of kelp ecology related to the existence of a bank of microscopic stages in the field.     

Influence of waves and currents on the growth rate of the kelp Laminaria digitata (Phaeophyceae)

The kelp Laminaria digitata growing in the low intertidal region along energetic coastlines are exposed to a range of hydrodynamic environments. Macroalgae in the intertidal zone can experience both waves and currents independently, but it is unknown how they influence growth rate. Relative growth rate of the meristematic region and the entire blade of L. digitata were measured to assess the influence of wave and current motion from three hydrodynamic environments: low current and low wave (LCLW), high current and low wave (HCLW), and high wave and low current (HWLC). Alongside hydrodynamic data, seawater nutrient concentrations and temperature were also collected and analyzed. Results suggest that differences in L. digitata relative growth rates were not attributed to the seawater nutrient concentrations or temperature, but attributed to the hydrodynamic environments. At the high current condition, kelp growth rate of the meristematic region was enhanced by 45% compared to the high wave condition. When including the entire blade growth rate, an average increase of 25% was observed between the high current and high wave condition. Potentially, the division in growth rate observed between the wave and current motion is related to the frequency and magnitude at which the hydrodynamic forces act. These findings highlight the complexity of the hydrodynamic environment and that forces associated with waves and currents may have a significant role on the productivity of kelp.     

Abiotic influences on bicarbonate use in the giant kelp,Macrocystis pyrifera,in the Monterey Bay

In the Monterey Bay region of central California, the giant kelp Macrocystis pyrifera experiences broad fluctuations in wave forces, temperature, light availability, nutrient availability, and seawater carbonate chemistry, all of which may impact their productivity. In particular, current velocities and light intensity may strongly regulate the supply and demand of inorganic carbon (Ci) as substrates for photosynthesis. Macrocystis pyrifera can acquire and utilize both CO₂ and bicarbonate (HCO₃^?) as Ci substrates for photosynthesis and growth. Given the variability in carbon delivery (due to current velocities and varying [DIC]) and demand (in the form of saturating irradiance), we hypothesized that the proportion of CO₂ and bicarbonate utilized is not constant for M. pyrifera, but a variable function of their fluctuating environment. We further hypothesized that populations acclimated to different wave exposure and irradiance habitats would display different patterns of bicarbonate uptake. To test these hypotheses, we carried out oxygen evolution trials in the laboratory to measure the proportion of bicarbonate utilized by M. pyrifera via external CA under an orthogonal cross of velocity, irradiance, and acclimation treatments. Our Monterey Bay populations of M. pyrifera exhibited proportionally higher external bicarbonate utilization in high irradiance and high flow velocity conditions than in sub‐saturating irradiance or low flow velocity conditions. However, there was no significant difference in proportional bicarbonate use between deep blades and canopy blades, nor between individuals from wave‐exposed versus wave‐protected sites. This study contributes a new field‐oriented perspective on the abiotic controls of carbon utilization physiology in macroalgae.     

Assessment of photosynthetic performance in the two life history stages of Alaria crassifolia (Laminariales,Phaeophyceae)

Understanding of the physiological responses of kelp to environmental parameters is crucial, especially in the context of environmental change that may have contributed to the decline of kelp forests all over the world. The current study presents the photosynthetic characteristics of the macroscopic sporophyte and microscopic gametophyte stages of the brown alga Alaria crassifolia from Hokkaido, Japan, as determined by examining their photosynthetic responses over a range of temperature and irradiance using dissolved oxygen and chlorophyll fluorescence measurements. Net photosynthetic rates of the sporophyte were consistently higher than those of gametophyte across temperature gradients and irradiance levels. Photosynthesis–irradiance curves at 8°C, 16°C, and 20°C revealed similar initial slopes (α = 0.4–0.9) on the two life history stages, but higher compensation (E _c = 4–7 μmol photons m^?2 s^?1) and saturation irradiances (E _k = 53–103 μmol photons m^?2 s^?1) for the sporophyte than for the gametophyte (E _c = 0–7 μmol photons m^?2 s^?1; E _k = 7–10 μmol photons m^?2 s^?1). Both stages exhibited chronic photoinhibition, as shown by the failure of recovery in their maximum quantum yields (F _v/F _m) following high irradiance stress, with greater possibility of photodamage at low temperature. Gametophytes were less sensitive to low temperatures than sporophytes, given their relatively stable F _v/F _m response. Nevertheless, temperature optima for photosynthesis of both stages coincide with each other at 20–23°C, which correspond to the growth and maturation periods of A. crassifolia in Japan. This species is also likely to suffer from thermal inhibition as both GP rates and F _v/F _m decreased above 24°C.     

Decline in Kelp in West Europe and Climate

Kelp ecosystems form widespread underwater forests playing a major role in structuring the biodiversity at a regional scale. Some seaweeds such as Laminaria digitata are also economically important, being exploited for their alginate and iodine content. Although some studies have shown that kelp ecosystems are regressing and that multiple causes are likely to be at the origin of the disappearance of certain populations, the extent to which global climate change may play a role remains speculative. Here we show that many populations of L. digitata along European coasts are on the verge of local extinction due to a climate-caused increase in sea temperature. By modeling the spatial distribution of the seaweed, we evaluate the possible implications of global climate change for the geographical patterns of the species using temperature data from the Coupled Model Intercomparison Project phase 5 (CMIP5). Projections of the future range of L. digitata throughout the 21st century show large shifts in the suitable habitat of the kelp and a northward retreat of the southern limit of its current geographic distribution from France to Danish coasts and the southern regions of the United Kingdom. However, these projections depend on the intensity of warming. A medium to high warming is expected to lead to the extirpation of the species as early as the first half of the 21st century and there is high confidence that regional extinction will spread northwards by the end of this century. These changes are likely to cause the decline of species whose life cycle is closely dependent upon L. digitata and lead to the establishment of new ecosystems with lower ecological and economic values.     

Elevated Temperature Affects Phenotypic Plasticity in the Bull Kelp (Nereocystis luetkeana,Phaeophyceae)

The sensitivity of kelps to elevated temperatures has been linked to recent declines in some kelp populations, with cascading impacts on marine communities. However, it remains unclear how thermal stress affects the ability of kelps to respond to other environmental factors, which could influence their vulnerability to climate change. We investigated the effect of thermal stress on the ability of the bull kelp Nereocystis luetkeana to acclimate to its surrounding hydrodynamic environment through tension-regulated plasticity in blade morphology. We first determined optimal and stressful temperatures for N. luetkeana by measuring growth over nine temperatures from 5°C to 22°C. We then exposed N. luetkeana blades to a factorial combination of temperature (13°C and 20°C) and tension (0.5 N and 2.0 N) simulating different flow conditions, and measured changes in blade length and width after 7 days. The temperature at which N. luetkeana exhibited maximum growth was estimated to be ~11.9°C, though growth was high over a relatively wide temperature range. When thermally stressed, N. luetkeana maintained morphological responses to simulated high flow, but were inhibited from acclimating to low flow, indicated by an inability of blades to widen. Our results suggest that N. luetkeana in sheltered habitats may be particularly vulnerable to climate warming, where an inability to adjust blade morphology to local hydrodynamic conditions could drive declines at sublethal levels of warming. As ecologically important foundation species, declines in sheltered kelp populations could result in major biodiversity loss and disrupt ecosystem function.     

UV‐radiation and elevated temperatures induce formation of reactive oxygen species in gametophytes of cold‐temperate/Arctic kelps (Laminariales,Phaeophyceae)

Enhanced UV‐radiation (UVR) through stratospheric ozone depletion and global warming are crucial stressors to marine macroalgae. Damages may arise through formation of reactive oxygen species (ROS) in gametophytes of ecologically important kelps, brown algae of the order Laminariales, Such stress‐induced damages may have a negative impact on their fitness and further impact their following life stages. In our study, gametophytes of three kelp species Alaria esculenta (L.) Grev., Laminaria digitata (Huds.) Lamour., Saccharina latissima (L.) Lane, Mayes, Druehl, Saunders from the Arctic, and of L. hyperborea (Gunnerus) Foslie from the North Sea were exposed to photosynthetically active radiation, UV‐A, and UV‐B radiation and four temperatures (2–18°C). ROS are formed predominantly in the peripheral cytoplasm and in chloroplasts especially after exposure to UVR. Superoxide (O₂*^‐) is additionally formed in small, globular cytoplasmic structures, possibly mitochondria. In the surrounding medium O₂*^‐‐concentration increased markedly at elevated temperatures and under UV stress in some cases. Ultrastructural damage was negligible pointing to a high stress tolerance of this developmental stage. Our data indicate that stress tolerant gametophytes of three Arctic kelp species should sustain their crucial function as seed bank for kelp populations even under prospective rising environmental perturbations.     

Photosynthetic acclimation of Chlorella saccharophila to heat stress

Photosynthesis is one of the most important metabolic processes of algae; which is altered as a stress response. During mass cultivation of algae, temperature rise and high light are major factors that affect biomass productivity. High temperature affects photosystem II (PSII) complex irreversibly, damaging intermolecular interactions in it. However, the impact of high temperature on photosynthesis is highly variable among different algal species, depending on the prior acclimation to environmental conditions they were exposed to. The acclimation plays an important role in combating high temperature stress via regulation of photosynthetic responses. Chlorophyll a fluorescence is a highly sensitive, non‐destructive and reliable tool for such measurements of photosynthetic parameters, which provides information about algal photosynthetic performance under given conditions. To understand the effect of heat stress on the responses of high light acclimated alga Chlorella saccharophila, chlorophyll a fluorescence transients were measured after heat exposure at 40°C. Our study demonstrates that rise in temperature for short duration; during open field cultivation reversibly affects the efficiency of PSII in light acclimated alga C. saccharophila. The effects of heat stress on chlorophyll a fluorescence in this alga, grown under high light (max‐1600 μmol photons m^?2 s^?1) are presented here; they are used to infer changes in photosynthetic process during its exposure to heat, as well as their recovery after 72 h. We speculate that heat resistance may have been acquired due to prior exposures to high light.