An examination of species dispersion patterns along the intertidal gradient on Macquarie Island (sub-Antarctica) using a restricted occupancy model

It has been widely held that intertidal zonation boundaries and the tidal emersion levels are causally related: the critical tidal level hypothesis. In order to evaluate this hypothesis, the dispersion patterns of species boundaries on the intertidal rocky shores of Macquarie Island (sub-Antarctica) were examined using a restricted occupancy model to test the null hypothesis that the observed species were randomly dispersed along the tidal emersion gradient. Subsequently this investigation demonstrated that the intertidal species boundaries on Macquarie Island shores were randomly dispersed with respect to the tidal emersion gradient. Hence no prima facie evidence was found to support the critical tidal level hypothesis. This suggests that tidal emersion is not a significant factor structuring intertidal communities on Macquarie Island.     

140 Ecophysiology of The Red Alga Porphyra Umbilicalis Kützing (Rhodophyta,Bangalies): Responses to Abiotic Stress

The effects of tidal elevation, emersion, sun exposure, and season on several antioxidant enzymes (ascorbate peroxidase, glutathione reductase, and catalase), pigments (phycoerythrin, phycocyanin, chlorophyll a and total carotene) and photosynthetic efficiency of photosystem II (Fv/Fm) in Porphyra umbilicalis were evaluated. Plants were collected monthly from sun‐exposed and shaded locations in the high, mid, and low intertidal following periods of tidal emersion ranging from 0–6 hours. Glutathione reductase activity was greatly affected by emersion during summer months, while ascorbate peroxidase and catalase activities showed no seasonal patterns. Differences in glutathione reductase and catalase levels occurred between sun‐exposed and shaded plants in the high and mid intertidal during summer. At all elevations, photosynthetic pigments showed a strong seasonal trend, with the effect of sun exposure being most apparent during summer. While total carotene increased with emersion during summer months, the combined effects of emersion and season were inconsistent for phycoerythrin, phycocyanin and chl a. Photosynthetic efficiency (Fv/Fm) decreased following emersion in summer and fall. During most months, sun exposed plants had lower Fv/Fm values compared to plants growing in the shade. This study emphasizes the importance of examining the effects of abiotic stresses simultaneously in order to reveal interactive relationships.     

Environmental correlates of phenotypic variation: do variable tidal regimes influence morphology in intertidal seaweeds?

Seaweed morphology is often shaped by the hydrodynamic environment. However, exposure to air at low tide represents an additional factor potentially affecting the morphology of intertidal species. Here, we examined the relationships between the morphology of Hormosira banksii, an important intertidal habitat‐forming seaweed in southern Australia, and environmental factors across multiple spatial scales around the island of Tasmania, Australia. Tasmania is surrounded by a diverse coastline with differences in wave exposure, tidal parameters, and temperature. We sampled Hormosira from four regions (100s km apart), three sites (10s km apart) within each region, and two zones (meters apart; eulittoral and sublittoral) at each site, and measured multiple morphological variables to test for differences in morphology at those different spatial scales. Thirteen environmental variables reflecting wave exposure, tidal conditions, and temperature for each site were generated to assess the relationship between Hormosira morphology and environmental variation. Morphology varied at all spatial scales examined. Most notably, north coast individuals had a distinct morphology, generally having smaller vesicles and shorter fronds, compared to other regions. Tidal conditions were the main environmental factors separating north coast sites from other sites and tidal regime was identified as the best predictor of morphological differences between regions. In contrast to other studies, we found little evidence that wave exposure was associated with morphological variation. Overall, our study emphasizes the role of tidal conditions, associated with emersion stress during low tide, in affecting the morphology of intertidal seaweeds.     

Sublethal stress in the intertidal zone: tidal emersion inhibits photosynthesis and retards development in embryos of the brown alga Pelvetia fastigiata

The effect of tidal emersion on survivorship, photosynthesis and embryonic development was studied in 8 h old zygotes and 7 d old embryos of the intertidal brown alga Pelvetia fastigiata (J. Ag.) DeToni. Zygotes and embryos were outplanted for single low tides in the intertidal zone on the central coast of California (U.S.A.) during June, 1990. Both zygotes and embryos exhibited close to 100% survival when outplanted beneath the canopy of adult P. fastigiata. Embryos (7 d old) also exhibited high survival when outplanted in a red algal turf, the microhabitat where most successful recruitment occurs. However, zygotes (8 h old) experienced high mortality (65–90%) when outplanted in the turf microhabitat. Embryos and zygotes that survived emersion experienced sub-lethal stress that temporarily impaired light-saturated photosynthesis when plants were reimmersed in seawater. The effects of sub-lethal stress were more pronounced in 8 h old zygotes than 7 d embryos, and more severe in the turf microhabitat than beneath the adult Pelvetia canopy. Zygotes outplanted in the red algal turf did not re-establish net photosynthesis until at least 6 h after re-immersion. Photosynthesis was less inhibited in 8 h old zygotes outplanted beneath the adult Pelvetia canopy, and recovered to control (non-emersed) levels within 3 h of re-immersion. Embryos (7 d old) were able to achieve positive net photosynthesis immediately on re-immersion after emersion in the turf or canopy microhabitats. Emersion also retarded the rate of embryonic development in 8 h old zygotes, delaying the formation of primary rhizoids, which help to attach the plant to the substrate. For example, at 19 h post-fertilization, 75% of control (non-emersed) zygotes had developed rhizoids, compared to 3% and 30% for zygotes outplanted in the turf and canopy microhabitats. The different emersion responses of 8 h old zygotes and 7 d old embryos appeared to be related to their ability to tolerate cellular dehydration. Overall, our data suggest that the effects of sub-lethal stresses may have been underestimated in studies of intertidal ecology.     

141 New Entities in the Order Chaetopeltidales

The effects of tidal elevation, emersion, sun exposure, and season on several antioxidant enzymes (ascorbate peroxidase, glutathione reductase, and catalase), pigments (phycoerythrin, phycocyanin, chlorophyll a and total carotene) and photosynthetic efficiency of photosystem II (Fv/Fm) in Porphyra umbilicalis were evaluated. Plants were collected monthly from sun-exposed and shaded locations in the high, mid, and low intertidal following periods of tidal emersion ranging from 0–6 hours. Glutathione reductase activity was greatly affected by emersion during summer months, while ascorbate peroxidase and catalase activities showed no seasonal patterns. Differences in glutathione reductase and catalase levels occurred between sun-exposed and shaded plants in the high and mid intertidal during summer. At all elevations, photosynthetic pigments showed a strong seasonal trend, with the effect of sun exposure being most apparent during summer. While total carotene increased with emersion during summer months, the combined effects of emersion and season were inconsistent for phycoerythrin, phycocyanin and chl a. Photosynthetic efficiency (Fv/Fm) decreased following emersion in summer and fall. During most months, sun exposed plants had lower Fv/Fm values compared to plants growing in the shade. This study emphasizes the importance of examining the effects of abiotic stresses simultaneously in order to reveal interactive relationships.     

ISOPOD TRACKWAYS FROM THE CRAYSSAC LAGERSTÄTTE, UPPER JURASSIC, FRANCE

Abstract:  Well-preserved arthropod trackways are described from the laminated limestones of the Crayssac Lagerstätte (south-west France, Lower Tithonian). They occur in sediments deposited in the temporary coastal mudflats of intertidal to supratidal zones. The trackways are referred to Pterichnus isopodicus isp. nov., and are interpreted as the locomotion traces of isopods. Different trackway morphotypes are recognized and clearly resulted from variations in the original consistency of the sediment. Sinuous trackways may correspond to vagrant activity on wet mud whereas numerous straight ones indicate a more rapid crawling on a soft-to-firm substratum (e.g. tidal flat during emersion). The preferred orientation of trackways indicates that isopods were crawling in a direction perpendicular to shoreline as a result of possible taxis induced by sediment wetness and/or by a migratory behaviour controlled by tidal rhythm. Unusually long emergence of the sediments may have favoured the preservation of dense networks of trackways. An isopod identity is supported by the general morphology of the tracks and the association of trackways with isopod body fossils. Archaeoniscus , which occurs abundantly in Late Jurassic deposits of England and France, was probably the trace-maker.     

SURVIVAL OF FUCOID EMBRYOS IN THE INTERTIDAL ZONE DEPENDS UPON DEVELOPMENTAL STAGE AND MICROHABITAT1

Embryos of the fucoid alga Pelvetia fastigiata (J. Ag.) DeToni were outplanted into the intertidal zone to assess survival during the physical stress brought about by emersion during a single low tide. Survival varied among microhabitats. Under the adult Pelvetia canopy, survival of 6-h-, 24-h-, 48-h-, and 1-wk-old embryos was nearly 100%. Almost all embryos of all ages died in exposed habitats on bare rock or within habitats where the Pelvetia canopy was removed experimentally. However, within red algal turfs, where most juvenile Pelvetia occur, survival was unusually age specific: 24- to 48-h-old embryos survived poorly compared to younger (6 h old) or older embryos (1 wk old). Survival patterns reflected microhabitat temperatures during the experiments. The fate of young post-settlement stages must be studied at these fine temporal and spatial scales to understand the organization of intertidal communities.     

Seasonal effects of sun exposure and emersion on intertidal seaweed physiology: Fluctuations in antioxidant contents, photosynthetic pigments and photosynthetic efficiency in the red alga Porphyra umbilicalis Kützing (Rhodophyta, Bangiales)

There is a great deal of speculation regarding the physiological and biochemical mechanisms that give certain seaweed species the ability to colonize the intertidal zone. Frequent exposure to ambient temperatures and high irradiance levels in addition to dehydration during tidal emersion generates acute physiological stress. The ability of seaweeds like Porphya to overcome these challenges and survive in such a harsh environment has been linked to elevated reactive oxygen metabolism. The current study focused on measuring seasonal changes in antioxidant enzymes plus alterations in pigment contents and photosynthetic efficiency of P. umbilicalis plants found growing in the uppermost intertidal zone.Our results suggest that P. umbilicalis exhibits increased antioxidant metabolism, which could contribute to its success in colonizing such a stressful habitat. Elevated levels of glutathione reductase GTR, catalase and carotenoid contents during emersion suggested heightened protection against reactive oxygen species ROS damage is a necessary attribute for species in the upper intertidal regions. This hypothesis was further strengthened by the finding that the greatest antioxidant increases were observed during summer months when irradiance levels and temperatures were at their peak. Winter emersion did not elicit the same physiological response, as antioxidant levels were similar in submersed and emersed plants.For the most part, photosynthetic pigments were largely affected by sun exposure and less by emersion stress. Shaded blades maintained higher concentrations of photosynthetic pigments compared to sun exposed thalli concurring with established research. Photosynthetic efficiency measurements indicated emersion and not sun exposure was the greater facilitator of photoinhibitory damage and ROS generation at PSII. The findings of this field study strengthen previous assertions that protection via elevated antioxidant metabolism and increased PSII repair are involved in providing relief from the acute environmental stresses in the intertidal zone.     

Linking Physiology To Ecological Function: Environmental Conditions Affect Performance And Size Of The Intertidal Kelp Hedophyllum Sessile (Laminariales,Phaeophyceae)

For autogenic ecosystem engineers, body size is an aspect of individual performance that has direct connections to community structure; yet the complex morphology of these species can make it difficult to draw clear connections between the environment and performance. We combined laboratory experiments and field surveys to test the hypothesis that individual body size was determined by disparate localized physiological responses to environmental conditions across the complex thallus of the intertidal kelp Hedophyllum sessile, a canopy‐forming physical ecosystem engineer. We documented substantial (> 40%) declines in whole‐thallus photosynthetic potential (as Maximum Quantum Yield, MQY) as a consequence of emersion, which were related to greater than 10‐fold increases in intra‐thallus MQY variability (as Coefficient of Variation). In laboratory experiments, desiccation and high light levels during emersion led to lasting impairment of photosynthetic potential and an immediate > 25% reduction in area due to tissue contraction, which was followed by complete loss of structural integrity after three days of submersion. Tissue exposed to desiccation and high light during emersion had higher nitrogen concentrations and lower phlorotannin concentrations than tissue in control treatments (on average 1.36 and 0.1x controls, respectively), suggesting that conditions during emersion have the potential to affect food quality for consumers. Our data indicate that the complex thallus morphology of H. sessile may be critical to this kelp’s ability to persist in the intertidal zone despite the physiological challenges of emersion and encourage a more nuanced view of the concept of “sub‐lethal stress” on the scale of the whole individual.     

Guidelines for evaluating performance of oyster habitat restoration should include tidal emersion: reply to Baggett et al.

Baggett et al. (2015) identified a set of three universal environmental variables to be monitored for evaluating all oyster habitat restoration projects: salinity, temperature, and dissolved oxygen. Perhaps evidencing a bias toward subtidal reefs, this set of parameters omits another first‐order environmental factor, tidal emersion. Intertidal oyster reefs can be the dominant reef habitat in estuaries, with clear zonation in oyster performance across the intertidal exposure gradient. Therefore, we propose to include tidal emersion as a fourth universal environmental parameter when designing and evaluating oyster restoration projects to better encompass the whole environmental spectrum along which reefs occur.     

The role of niche measures in explaining the abundance–distribution relationship in tropical lotic chironomids

In situ measurements of both community metabolism (primary production and respiration) and PAM fluorometry were conducted during emersion on intertidal sediments in the Mont Saint-Michel Bay, in areas where oysters and mussels were cultivated. Results highlighted a low benthic metabolism compared to other intertidal areas previously investigated with the same methods. Comparisons between gross community primary production and relative electron transport rates confirmed this statement. More specifically, primary productivity remained very low all over the year, whereas the associated microalgal biomass was estimated to be high. We suggest that the microphytobenthic community studied was characterized by a self-limitation of its primary productivity by its own biomass, as previously shown in Marennes-Oléron Bay for example. The almost permanent high biomass would represent a limiting factor for micromigration processes within the first millimetres of the sediment. This could be explained by very low resuspension processes occurring in the western part of the bay, enhanced by the occurrence of numerous aquaculture structures that could decrease tidal currents in the benthic boundary layer. Handling editor: N. Desroy     

The effects of prolonged emersion and submersion by tidal manipulation on marine macrobenthos

Effects of tidal manipulation, resulting in prolonged periods of emersion and submersion or in protracted tidal cycles, on estuarine benthic animals are reviewed.Prolonged submersion periods did not show effects on mortality of most benthic animals tested, with the exception of the crumb-of-bread sponge Halichondrea panicea, which, at low water-flow rates, was covered with a layer of bacteria and subsequently died.Protracted low-water periods of 18 hours during several weeks hardly caused any mortality. However, protracted low-water periods of 30 hours during some weeks or emersion during several days caused a strong increase in mortality, depending on: the duration of emersion, temperature, condition of the animals, species and age. At temperatures below –1 °C and above 24 °C mortality was generally high. Animals with a low glycogen content were more sensitive to emersion than those with a high content. Species with a shell and those that are relatively big were less sensitive than those without a shell or of small size.The reproductive cycle of benthic animals could be delayed or accelerated by both emersion and submersion.     

Physiological responses to emersion in the intertidal green crab,Carcinus maenas (L.)

The green shore crab, Carcinus maenas, undergoes on average 6?h periods of emersion during each low-tide cycle during the summer months. Under those conditions, the crab is cut off from its normal water environment and is exposed to potential stress from a suite of environmental and physiological changes: dehydration, compromised gas exchange and resultant internal hypoxia and hypercapnia, thermal stress, and ammonia toxicity. This study examined the comprehensive responses of the green crab in water and to a 6?h emersion period laboratory simulation of a tidal cycle followed by a 1?h re-immersion period, measuring indicators of dehydration, hemolymph osmolality, oxygen uptake, hemolymph acid–base status, heart and ventilatory rate, and hemolymph ammonia and ammonia excretion. Green crabs showed physiological responses of varying magnitude to 6?h of emersion. Individuals were found in the field exclusively under rocks and large clumps of seaweed where temperatures were approximately half those of exposed surfaces and relative humidity was about twice as high as ambient air. During emersion, crabs lost less than 5% of their wet weight, and hemolymph osmolality did not increase significantly. Oxygen uptake continued in air at about 50% of the control, aquatic values; and the gills continued to be ventilated by the scaphognathite, albeit at lower rates. Hemolymph lactate concentrations increased, indicating a shift to a greater reliance on anaerobic metabolism to support energetic needs. A slight acidosis developed in the hemolymph after 1?h of emersion, but it did not increase thereafter. Ammonia concentrations in the hemolymph were unchanged, as ammonia was volatilized by the gills and excreted into the air as NH₃ gas. These results show that the green crab copes with emersion by seeking refuge in microhabitats that mitigate the changes in the physical parameters of intertidal emersion. Physiologically, desiccation is avoided, cardio-respiratory processes are maintained at reduced levels, and hemolymph acid–base balance is minimally affected. Ammonia toxicity appears to be avoided by a shift to excreting NH₃ gas directly or indirectly to air.     

WATER MOTION AND MORPHOLOGY IN CHONDRUS CRISPUS (RHODOPHYTA)1

Morphological variability of intertidal Chondrus crispus Stackh. fronds along a small open rocky coast was related to wave exposure and emersion. Cluster analysis revealed two well-defined morphologies: filiform and planiform, named the N morphotype and B morphotype, respectively. We propose a rapid method of classifying fronds based on the morphology of the cross section at half the height on the thallus. The N morphotype is characterized by fewer dichotomies per unit length, a circular cross section with a large inner cortex, and narrow fronds. It is abundant at low intertidal and exposed sites. The B morphotype is characterized by more dichotomies, smaller sizes, a subelliptical or flattened cross section, and broad fronds. It is abundant at high intertidal sites in sheltered areas. Regression analysis revealed a major effect of water movements on frond morphology with respect to tidal level, which was more evident at high intertidal levels. No relationships were observed between morphology and life history phases.     

The maintenance of shore-level size gradients in an intertidal snail (Littorina sitkana)

Summary The size of many intertidal animals varies with tidal height. These size gradients could be produced by growth or survival varying with tidal height, or by animals moving to a preferred tidal level. The body size of the snail, Littorina sitkana, increases steadily with tidal height in rocky high intertidal habitats of British Columbia. To determine how size gradients were maintained in L. sitkana, I quantified how growth, survival, and snail movement varied with tidal height. I studied populations of L. sitkana found on sheltered pebble beach and exposed basaltic shelf habitats. Mark-recapture studies and experimental transplants showed that growth could not have produced the size gradients because snail growth rates in both habitats were as fast or faster at low tidal levels (where the snails were the smallest) than at high tidal levels. However, survival rates were lowest at low tidal levels. On pebble beaches, this was due to size selective predation on large snails by the pile perch, Rhacochilus vacca. On basaltic shelves, heavy wave action at low tidal levels may have caused the poor survival rates. Transplanted snails moved homeward on pebble beaches, but not on basaltic shelves. Reduced survival rates at low tidal levels cause size gradients in both habitats, and snail movement helps to maintain size gradients on pebble beaches.     

Fortnightly light and temperature variability in estuarine intertidal sediments and implications for microphytobenthos primary productivity

Photosynthetically active radiation (PAR) and temperature were measured continuously at the surface of estuarine intertidal sediments in the Tagus estuary, Portugal, along two spring-neap tidal cycles. PAR and temperature were strongly conditioned by the periodic tidal inundation, with large and abrupt variations occurring during flooding and ebbing. PAR levels reaching the sediment surface decreased very rapidly to zero or very low values during most of the daytime immersion. Inundation during high tide had the general effect of attenuating the amplitude of daily temperature fluctuation, with the incoming water usually warmer than the sediment during the night or early morning and cooler during the day. The daily progression of tidal emersion resulted in a clear fortnightly variation in total daily PAR reaching the sediment surface, while both daily mean temperature and mean temperature of diurnal low tide periods failed to exhibit a well-defined fortnightly periodicity. The obtained results indicate that the estuarine intertidal environment is dominated, at sub-seasonal time scales, by fortnightly periodicity in irradiance and temperature conditions favourable for benthic photosynthesis.     

A refutation of critical tidal levels as determinants of the structure of intertidal communities on British shores

Early hypotheses to account for the pattern of zonation and vertical distribution of species on rocky shores in Britain invoked the concept of critical tidal levels. These levels were at heights on the shore where the upper or lower boundaries of distribution of a number of species coincided. Critical levels were correlated with heights on the shore where there were changes in the rate of change, with height, of the annual proportion of the time spent emersed. This was calculated from predicted tide tables. One implications of this was that different assemblages of intertidal species were present between particular levels on the shore. Futhermore, this hypothesis depended on patterns of vertical distribution of several species being identically controlled by physical factors associated with the rise and fall of the tide. This conflicts with more recent hypotheses, based upon experimental evidence, which include the effects of biological interactions among species on their patterns of distribution.The methods of calculation of the annual emersion curve were approximate, with a high degree of extrapolation. The hypothesis that critical levels exist has not been quantitatively tested.New, more accurate calculations of the emersion curve, from predicted tidal heights, indicate a smooth, monotonic curve against height on the shore. This eliminates the possibility of correlation between the coincident boundaries of distribution of a group of species and a height at which the curve changes in slope.The upper and lower boundaries of species were recorded in transects on five shores in different parts of Britain. If critical level exist, the upper or lower boundaries of a set of species must depart from random dispersion up and down the shore, to be clumped or aggregated at the critical levels. A test for non-random dispersion of the boundaries indicated no significant departure from random on any of the shores sampled.There is thus no evidence that critical tidal levels exist, because there is no evidence that the upper, or lower, boundaries of vertical distribution of intertidal species are in any way aggregated. There is no possibility of correlation between changes of slope of the emersion curve at particular heights on the shore and the patterns of distribution of intertidal species on British shores. The hypothesis of critical tidal levels must be abandoned and the rôle of the effects of tidal rise and fall on the vertical distribution of intertidal species must be re-examined in any new hypotheses to account for observed patterns of zonation.     

Characterizing biogeochemical fluctuations in a world of extremes: A synthesis for temperate intertidal habitats in the face of global change

Coastal and intertidal habitats are at the forefront of anthropogenic influence and environmental change. The species occupying these habitats are adapted to a world of extremes, which may render them robust to the changing climate or more vulnerable if they are at their physiological limits. We characterized the diurnal, seasonal and interannual patterns of flux in biogeochemistry across an intertidal gradient on a temperate sandstone platform in eastern Australia over 6 years (2009–2015) and present a synthesis of our current understanding of this habitat in context with global change. We used rock pools as natural mesocosms to determine biogeochemistry dynamics and patterns of eco‐stress experienced by resident biota. In situ measurements and discrete water samples were collected night and day during neap low tide events to capture diurnal biogeochemistry cycles. Calculation of pH_T using total alkalinity (TA) and dissolved inorganic carbon (DIC) revealed that the mid‐intertidal habitat exhibited the greatest flux over the years (pH_T 7.52–8.87), and over a single tidal cycle (1.11 pH_T units), while the low‐intertidal (pH_T 7.82–8.30) and subtidal (pH_T 7.87–8.30) were less variable. Temperature flux was also greatest in the mid‐intertidal (8.0–34.5°C) and over a single tidal event (14°C range), as typical of temperate rocky shores. Mean TA and DIC increased at night and decreased during the day, with the most extreme conditions measured in the mid‐intertidal owing to prolonged emersion periods. Temporal sampling revealed that net ecosystem calcification and production were highest during the day and lowest at night, particularly in the mid‐intertidal. Characterization of biogeochemical fluctuations in a world of extremes demonstrates the variable conditions that intertidal biota routinely experience and highlight potential microhabitat‐specific vulnerabilities and climate change refugia.     

Seasonality in nutrient concentrations and stable isotope ratios of Halophila ovalis growing on the intertidal flat of SW Thailand

Halophila ovalis on the intertidal flat of SW Thailand maintains its presence under heavy grazing by dugongs; nevertheless, the site provides lower availability of nutrients in the water column to the seagrasses due to emersion during ebb tide. To examine the seasonality of the nutrient status of fast-growing H. ovalis in the intertidal flat, and to assess whether emersion at low tide affects the availability of nutrients and inorganic carbon, monthly changes in carbon (C), nitrogen (N), and phosphorus (P) concentrations of H. ovalis were determined. Carbon and nitrogen stable isotope ratios were also determined to examine the carbon and nitrogen sources. CNP concentrations of the leaves and petioles of H. ovalis at the intertidal flat were considerably lower than those of the same species previously reported from a subtidal environment throughout the observation period of 8 months. The δ¹³C of the above-ground parts of H. ovalis was lower than in previous reports, suggesting that CO₂ was sufficient for its growth. The CNP concentrations of H. ovalis on the intertidal flat increased after a decrease in grazing pressure by dugongs. It is suggested that H. ovalis sustained rapid growth on the intertidal flat by lowering its nutrient concentrations under heavy grazing by dugongs to maintain its presence. Received: March 2, 2001 / Accepted: June 6, 2001     

Effects of UV radiation on five marine microphytobenthic Wadden sea diatoms isolated from the Solthörn tidal flat (Lower Saxony,southern North Sea) – Part I: growth and antioxidative defence strategies

The unconsolidated sediment of intertidal mudflats constitutes a highly unstable environment, due to continuously changing water levels and currents as well as temporary exposure to the air. Therefore, diatoms inhabiting marine intertidal areas are subjected to strongly changing surface light and UV intensities due to exposure at low tide. Five marine intertidal diatoms (Achnanthes exigua, Cocconeis peltoides, Diploneis littoralis, Navicula digitoradiata and Amphora exigua) were isolated from the Solthörn tidal flat (Lower Saxony, southern North Sea). Semi-continuous cultures were used to determine the effect of UV radiation (photosynthetically active radiation only [PAR], PAR+UV-B, PAR+UV-A, PAR+UV-B+UV-A) during short- and long-term exposure (6 h or 30 days). Growth rates, chlorophyll a (chl a), antioxidant capacities, accumulation of phenolic compounds (e.g. flavonoids) and DMSP, and activities of antioxidant enzymes (superoxide dismutase, ascorbate peroxidase, monodehydroascorbate reductase and glutathione reductase) were assessed. UV-A had only minor effects on cells, while growth rate, chl a content and protein content were significantly reduced after long-term UV-B exposure. Achnanthes exigua extracts showed the highest antioxidant capacity. The highest activity of SOD, APX and MDHAR was found under long-term combined UV exposure (PAR+UV-B+UV-A). Overall, the antioxidative defence of the five isolates was stimulated during exposure to UV radiation, as may be found during emersion. Emersion induces oxidative stress and, as a result, growth of the five diatom taxa was inhibited to suit changing environmental conditions. All five taxa tested in the present study showed species-specific acclimatization potentials, providing possible explanations for variability in population, species composition and ecosystem structures in the face of climatic variations.