Photosynthetic and morphological responses of eelgrass (Zostera marina L.) to a gradient of light conditions

Zostera marina L. (eelgrass) from Great Bay Estuary, New Hampshire and Maine (USA), was transplanted in outdoor mesocosms and subjected to four light treatments (100, 58, 34 and 11% surface irradiance, SI) between May and September 2003 to investigate the relationship between light availability and the growth and survival of eelgrass. Evaluating eelgrass seedlings and adult mature plants demonstrated no differences in photosynthetic response after 22 days of acclimation. During at least the first 19 days of shading, maximum electron transport rate (ETR_max) rate of eelgrass did not differ significantly between light treatments. After 40 days, a significant reduction in ETR_max and minimum saturating light was observed in plants growing at 34% SI and below. Morphological responses exhibited a linear increasing trend with greater light. 34% SI exhibited drastic reductions (to less than 25% of control) in rhizome growth, shoot density, shoot production, number of nodes per plant and plant weight at the end of the study (81 days). Shoot to root ratio at 34% SI increased by > 50%. Plants shaded to 58% SI showed no significant difference from the control in plant parameters except an increased rate of rhizome elongation. Our results link the lower shoot densities with shading to the slow growth rate of horizontal rhizomes and a total lack of lateral expansion at 11% SI. ETR_max declined over time in plants at 11% SI resulting in 81% mortality, no lateral branching and no morphological development, indicating that the minimum light required for long-term eelgrass growth and survival is greater than the previously suggested 11% SI. We demonstrate that eelgrass plants at these latitudes can persist at light levels of 58% SI and above, and are light-limited at 34% SI and below.     

Population dynamics and life history adaptations of Littorina neglecta Bean in an eelgrass meadow (Zostera marina L.) in Nova Scotia

The density of Littorina neglecta Bean from an eelgrass meadow in Nova Scotia was highest in mid-summer (July 1979; 45059 per m²) but fell rapidly during late summer and early autumn due to high mortality of new recruits and two year old individuals. Density remained constant until the following spring when it increased, due to movement of some adults into the study area and the release of young by females (up to ≈ 34000 per m²). During most of the year individuals were found on vegetation (live and dead eelgrass and drift algae) but nearly all moved down to the mud surface during winter to avoid ice (up to 25 cm thick) which was present from December to early March. Movement of ice was not a source of mortality for L. neglecta. Size-frequency histograms showed distinct cohorts and it was possible to follow the growth of year classes. Maximum age was 3 yr and maximum shell height was 4.56 mm. Males and females matured when 1 yr old at 1.7 and 1.8 mm shell height respectively. Size-specific fecundity was very high for such a small littorinid. These life history attributes appeared to be related to the low probability of survival to the maximum age. L. neglecta consumed epiphytes and detrital particles from the surface of eelgrass leaves, and the proportion of eelgrass plants with epiphytes was always low. Grazer exclusion experiments showed that L. neglecta controlled the amount of epiphytes and detritus on eelgrass leaves, and the population thus had a limited food supply. There were significant positive relations between surface area of plants and the numbers of adult and 0 + aged L. neglecta during summer and at that time 0 + individuals were restricted to the base of plants where epiphyte biomass was lowest. Post-recruitment mortality was probably related to intraspecific competition for food. Predation by a naticid gastropod probably accounted for much of the mortality of 2 + aged individuals. L. neglecta has a major positive influence on the growth of eelgrass and probably contributes significantly to nutrient recycling in the eelgrass community.     

Dissolved and particulate organic matter in contrasting Zostera marina (eelgrass) sediments

The influence of seagrass Zostera marina on sediment characteristics was examined in two contrasting sediments, one organic-rich and one organic-poor. The presence of plants leads to reduced sediment redox potential in both sediment types compared to bare sediment with the largest effects in the organic-poor sediment. Z. marina stimulated the sulfate reduction rates in organic-poor sediment with ∼50% and higher pools of dissolved organic carbon (DOC) were found. In contrast, sulfate reduction rates were lower in vegetated compared to bare sites in the organic-rich sediment. Despite a low contribution of dissolved carbohydrate (DCHO) to the DOC pool (<5%), the seagrass vegetation was responsible for an increase of ∼50% in DCHO pools with a peak in the root zone suggesting that Z. marina supplied DCHO to the pore waters. The Z. marina meadows also enhanced the contribution of particulate carbohydrate (PCHO) to sedimentary particulate organic carbon (POC) pools by 6-14% compared to bare sediment. Although the PCHO pools were higher in organic-rich than organic-poor sediments, the analyses of carbohydrate composition revealed that three groups of neutral sugars including glucose, galactose and mannose+xylose were the major compounds of PCHO and contributed with >60% to sedimentary carbohydrate pools at both sites. Only glucose showed depletion with depth in the vegetated sediments, whereas the percentage of ribose and rhamnose increased indicating a selective degradation of labile carbohydrates in the meadows. Galactose and mannose+xylose appeared to represent a refractory part of carbohydrate that remained after degradation of the more labile components. The sugar content was rather constant with depth at the bare organic-rich sediment indicating that only recalcitrant carbohydrate pools were buried. There was less difference in the PCHO composition profiles between vegetated and bare organic-poor sediments.     

Assessing within habitat variability in plant demography, faunal density, and secondary production in an eelgrass (Zostera marina L.) bed

This investigation addressed faunal relationships with habitat structure within a Zostera marina community targeting differences between seagrass bed edge and interior. Z. marina biomass was significantly higher from the interior portions of the bed compared to the edge, but shoot density did not vary. Additionally, leaf width and length were significantly greater in the interior of the bed, suggesting greater total leaf area. Densities of larger organisms (> 0.85 mm) were significantly greater in vegetated samples (Z. marina edge and interior) compared to unvegetated, but an analysis of similarities demonstrated significant faunal community differences among each of the identified habitats. Densities of small organisms (0.25-0.85 mm), however, were significantly greater at Z. marina edge compared to unvegetated samples and Z. marina interior. Additionally, secondary production (μg AFDW day^− 1) was estimated based on the size distribution of taxa and showed significantly greater production from samples gathered in Z. marina compared to unvegetated samples. The relative size distribution of taxa was assessed using regression analysis and results showed that the size distribution was similar for samples collected at edge and interior Z. marina, but these distributions differed significantly when compared to unvegetated samples. The results of this study suggest that although similarities exist between edge and interior portions of Z. marina beds, especially compared to unvegetated habitats, noteworthy differences in faunal density, species composition, size distribution, and secondary production exist between edge and interior Z. marina.     

Seed rafting as a dispersal strategy for eelgrass (Zostera marina)

Using observations, experiments and model simulations we investigate the possibility and range of long-distance dispersal by seed rafting in eelgrass, Zostera marina, at the Swedish west coast. A field experiment showed that the breaking strength of post-flowering rhipidia decreased significantly over a 30-day period, indicating that ontogenetic changes of the morphology facilitate dislodgement of the rhipidia at the optimal time for dispersal. We recorded positive buoyancy of detached post-flowering rhipidia for at least 26 days. Finally, we found evidence for sexual reproduction via seeds in the study area. These results, together with field measurements of wind-driven surface-transport velocity of rhipidia and wind data from 7 years, allowed us to model a dispersal potential of up to 150 km in one season along the Swedish west coast, which corresponds well with published estimates of dispersal potential from both observational and genetic studies. The results add to evidence that rafting of seeds on floating reproductive shoots is an important dispersal strategy for eelgrass, allowing for colonisation of new areas and for recolonisation after die-back events.     

Seasonal acclimation in metabolism reduces light requirements of eelgrass (Zostera marina)

We investigated the ability of eelgrass (Zostera marina) to adjust light requirements to seasonal changes in temperature, light and nutrient conditions through changes in metabolism, pigment and nutrient content. In agreement with expectations we found that rates of respiration and light saturated photosynthesis of summer acclimated plants peaked at higher temperatures (5 °C and 2 °C higher, respectively), and were lower than of winter acclimated plants, both at sub- and supra-optimal temperatures. Moreover respiration rates were generally more sensitive to increasing temperatures than photosynthetic rates, especially so for cold acclimated plants in February (36% higher Q₁₀-values). These changes were accompanied by a reduction in chlorophyll a and nitrogen concentrations in leaves by 35% and 60% respectively from February to August. The critical light requirement (E_C) of Z. marina to maintain a positive carbon balance increased exponentially with increasing temperature but less so for summer-acclimated than for winter-acclimated plants. However, combining E_C vs temperature models for whole-plants with data on daily light availability showed that seasonal acclimation in metabolism increased the annual period, when light requirements were meet at the 2-5 m depth interval, by 32-66 days. Hence, acclimation is an important mechanism allowing eelgrass to grow faster and penetrate to deeper waters. Critical depth limits estimated for different combinations of summer temperatures and water clarity in a future climate scenario, suggested that expected increases in temperature and nutrient run-off have synergistic negative effects, especially in clear waters, stressing the importance of continued efforts to improve water clarity of coastal waters.     

Phenology of eelgrass, Zostera marina L., along latitudinal gradients in North America

Eelgrass, Zostera marina L., was collected monthly from 1976 to 1979 and its phenology compared in the distributional range on both coastlines of North America. Each of three reproductive phenophases (initial appearance of macroscopically visible flower buds, initial anthesis, initial appearance of visible fruits) differed significantly among the sites in their dates of occurrence. Among the sites, flowering at the same latitude was later in the Atlantic than in the Pacific and flowering along a latitudinal gradient occurred increasingly later at more northern latitudes in the Atlantic than in the Pacific. Although the reproductive periodicity probably is controlled primarily by water temperature at the different sites, the variation in timing at the same latitude among Pacific and Atlantic sites indicates that Z. marina may include genotypes with different temperature requirements that are selectively adapted to different habitats along the latitudinal gradients.     

The ecology of eelgrass, Zostera marina (L.), fish communities. I. Structural analysis

Fish populations utilizing eelgrass, Zostera marina (L.), beds in two different estuarine areas near Beaufort, North Carolina were analysed and compared to determine aspects of their community structure. The fish community of the eelgrass beds was characterized by low diversity and high standing crops of biomass and energy, both of which showed seasonal variation. Wide temperature fluctuations related to the overall shallowness of the beds probably regulated the diversity of fishes utilizing the beds. This community was dominated by pinfish, Lagodon rhombiodes (L.), which comprised 45% and 67% of the fish biomass in the Phillips Island and Bogue Sound beds, respectively.Changes in total body caloric content were probably related to developmental stages and changes in diet. Adult fish often had significantly higher weight-specific caloric contents than juvenile fish. Monthly or seasonal variations in caloric content of the organic matter of pinfish had little influence on the caloric content within the various sizes of pinfish.There was a significant correlation between fish biomass, temperature, and Zostera biomass. Fish biomass was highest when temperature and grass biomass were at a maximum. In general, water depth over the beds had little effect on the standing crop of fish within the bed, but cooler waters which occurred at night, darkness, or both, had a large effect.     

Effects of filamentous algal mats on sulfide invasion in eelgrass (Zostera marina)

The effect of filamentous algae invasion into Zostera marina meadows on water quality, sediment sulfur pools and sulfide invasion into plant tissues was studied experimentally. Sulfide invasion was assessed through analysis of sulfur isotopic composition (δ³⁴S) and total sulfur (TS) concentrations in plant tissues. The algal mats (5 and 10 cm thickness) depleted oxygen in the mats and increased the pools of sulfides in the sediments. Plants exposed to algal mats had δ³⁴S signals closer to the δ³⁴S of sediment sulfide, whereas plants with no mats present had δ³⁴S signals closer to the δ³⁴S of seawater sulfate, indicating a higher sulfide invasion in plants exposed to algal mats. The δ³⁴S varied between the plant tissues with the leaves having more positive δ³⁴S signals than roots and rhizomes, indicating that sulfide was invading into the roots and moved to the other tissues through the lacunae. TS concentrations were higher in plants exposed to algal mats suggesting that sulfur derived from sediment sulfide accumulated in the plants. F_sulfide showed that up to 50% of the sulfides in the plants were derived from sedimentary sulfides. The combined effect of water column anoxia in the lower parts of the meadow and high sulfide invasion into the plants lead to significantly reduced growth rates after 3 weeks and the below-ground tissues showed signs of degradation suggesting that algal mats invasion in to Zostera marina meadows can result in seagrass decline.     

The ecology of eelgrass, Zostera marina (L.), fish communities. II. Functional analysis

Consumption, production, and respiration of fish communities utilizing two eelgrass beds in a shallow estuarine system near Beaufort, North Carolina have been estimated for 1971–1972: annual production was 21.7 kcal/m² in each bed with pinfish accounting for 45 and 68% of the production in the Phillips Island and Bogue Sound beds, respectively. Annual community respiration was 57.9 and 69.7 kcal/m² in the two beds with pinfish accounting for 62.6 and 26.7% of the total in the Bogue Sound and Phillips Island beds, respectively. Estimation of the annual food energy consumed by the eelgrass fish community using the Winberg and daily ration methods gave values within 6% of each other.Energy turnover was high (2.8), and the efficiency of energy dissipation low for the two eelgrass fish communities, suggesting that the resident fish populations were adapted to the temperature extremes within the bed. High ecological efficiencies of 0.24 and 0.23 and the high overall efficiency of the eelgrass system (production/solar radiation) of 0.0051 and 0.0086% indicate that the eelgrass beds are efficient systems for converting consumed energy and solar radiation into fish.     

Evaluation of a mechanical seed planter for transplanting Zostera marina (eelgrass) seeds

Few seagrass transplant projects worldwide have relied on seeds, and those projects using Zostera marina (eelgrass) seeds have generally found low rates of seedling establishment (<10%). We compared seedling establishment achieved by a mechanical seed planter with seeds broadcast on the sediment surface by hand. The planter injected seeds into the sediment by pumping the seeds, suspended in a gelatin-based matrix, to a benthic sled with eight planting nozzles. As a control for the gel, seeds were also injected into the sediment without gel using a hand-held pipette. Seeds were planted at a density of 300 m⁻² into six replicate plots at each of three sites in the Chesapeake Bay region in September 2005, with seedling establishment measured in April 2006. Burying seeds, either with or without gel, had an overall positive effect on seedling establishment, but the effectiveness and the best method varied among sites. Mean seedling establishment for machine-planted seeds was significantly greater than for broadcast seeds at the Piankatank River site (4% vs. 1%), but not at the York (1.2% vs. 1.4%) or Spider Crab Bay (10.1% vs. 7.4%) sites. The effect of the gel was inconsistent among sites, with the highest seedling establishment (18.8%), resulting from seeds injected by pipette without gel at the Spider Crab site. Seed burial shows promise for increasing seedling establishment relative to seed broadcasting in the Chesapeake region, but further investigation of seed–sediment interactions at specific restoration sites is necessary. Low seedling establishment rates remain a bottleneck for seed-based eelgrass restoration.     

Application and assessment of a nutrient pollution indicator using eelgrass (Zostera marina L.) in Barnegat Bay-Little Egg Harbor estuary, New Jersey

Eutrophication degrades numerous estuaries worldwide and a myriad of assessment metrics have been developed. Here, we apply an example of a previously developed metric (Lee et al., 2004) designed to indicate incipient estuarine eutrophication to validate this technique in an already eutrophic estuary end-member, Barnegat Bay-Little Egg Harbor, New Jersey. The metric, termed ‘Nutrient Pollution Indicator’ (NPI) uses eelgrass (Zostera marina L.) as a bioindicator and is calculated as the ratio of leaf nitrogen content (%N) to area normalized leaf mass (mg dry wt cm⁻²). Eelgrass samples were collected along the entire length of the Barnegat Bay-Little Egg Harbor from June to October 2008 to determine if leaf chemistry and morphology reflect eutrophication status and a north-south gradient of nitrogen loading from the Barnegat Bay watershed. Nitrogen content, area normalized leaf mass, and NPI values all significantly (p < 0.05) varied temporally but not spatially. NPI values did not significantly correspond to the north-south gradient of nitrogen loading from the Barnegat Bay watershed. The NPI metric is therefore not deemed to reliably indicate estuarine eutrophic status. Differences between sampling effort (number of stations) and replication did not bias the overall conclusions.     

Effects of aerobic versus anoxic conditions on glutamine synthetase activity in eelgrass (Zostera marina L.) roots: regulation of ammonium assimilation potential

Root glutamine synthetase (GS; EC 6.3.1.2) activity was measured daily (0 to 4 days) for eelgrass (Zostera marina L.) plants held under continuous darkness rooted in sediments, continuous darkness without sediments, continuous light without sediments, and control light/dark cycle (Control L/D). Roots experiencing prolonged aerobiosis exhibited lower activity in vitro than controls, whereas roots experiencing prolonged anoxia exhibited increased activity. Plants held in darkness without sediments had activity intermediate between controls and anoxic roots. One-hour pretreatment of root extracts with ATP slightly reduced in vitro glutamine synthetase activity, whereas pretreatment with ADP and AMP increased activity ≈50%. While glutamine synthetase activity increased with higher adenylate energy charge (AEC) in the reaction mixture, pretreatment of enzyme extracts at high adenylate energy charges decreased subsequent activity relative to pretreatment at lower energy charges. One-hour pretreatment with l-alanine (Ala) had little effect on enzyme activity. Pretreatment with l-glutamine (Gln), l-glutamate (Glu), and γ-amino butyric acid (GABA) increased activity ≈75%. Incubation of excised roots under anoxic conditions for 24 h nearly doubled enzyme activity. However, addition of cycloheximide to anoxic root incubations lessened or prevented the increase in activity. It appears that enhanced glutamine synthetase activity following periods of root anoxia results from interactions with metabolites that fluctuate between aerobic and anoxic conditions, particularly adenylates, and from de novo synthesis of glutamine synthetase or some other protein synthesis-dependent process.     

Experimental analysis of the response and recovery of Zostera marina (L.) and Halodule wrightii (Ascher.) to repeated light-limitation stress

Seagrasses are considered important indicators of decline in water quality resulting in increased light attenuation that negatively influences their growth and survival. Chronic light-limitation interspersed with unpredictable acute attenuation events have had poorly understood effects on seagrass recovery dynamics. Zostera marina (eelgrass) and Halodule wrightii (shoalgrass) were subject to a matrix of light-deprivation events followed by recovery periods to mimic repeated acute shading events. Plant survival, morphology, biomass, chlorophyll content, and Fv/Fm were assessed over time to determine recovery. At the end of the experiment, all plants were harvested and species-specific treatment effects were determined. Significant differences due to treatments were noted in all parameters measured. In general, responses were similar for both life-stages and between species, suggesting similar physiological tolerance to repeated acute light-attenuation events. Only plants in treatments where light-deprivation was followed by a recovery interval of at least the same duration showed signs of long-term survival. Chlorophyll fluorescence (Fv/Fm) was an important metric for assessing recovery, but it failed to detect the onset of mortality in many plants. Other metrics of plant condition need to be assessed and coupled with chlorophyll fluorescence data to assess seagrass “health”. This is of particular importance in field studies, where the history of the plants is largely unknown.     

Food web analysis of an eelgrass (Zostera marina L.) meadow and neighbouring sites in Mitsukuchi Bay (Seto Inland Sea,Japan) using carbon and nitrogen stable isotope ratios

The contribution of benthic microalgal production has been compared both within and outside a coastal eelgrass (Zostera marina L.) meadow. Carbon and nitrogen stable isotope ratios of suspended particulate organic matter (POM), epiphytic and epilithic organic matter (EOM), leaves of Z. marina (inside the meadow only) and two secondary consumer species (small crustaceans and fish) were measured inside and outside a meadow in Mitsukuchi Bay, Northwest Seto Inland Sea, Japan. Inside the meadow, primary producers (epiphyton) and consumers showed higher δ¹³C signatures than outside. Primary and secondary consumers inside the meadow were mainly dependent on epiphyton on the leaves of Z. marina, while consumer species outside the meadow were basically dependent on epilithon.     

Cloning of TPS gene from eelgrass species Zostera marina and its functional identification by genetic transformation in rice

The full-length cDNA sequence (2613 bp) of the trehalose-6-phosphate synthase (TPS) gene of eelgrass Zostera marina (ZmTPS) was identified and cloned. Z. marina is a kind of seed-plant growing in sea water during its whole life history. The open reading frame (ORF) region of ZmTPS gene encodes a protein of 870 amino acid residues and a stop codon. The corresponding genomic DNA sequence is 3770 bp in length, which contains 3 exons and 2 introns. The ZmTPS gene was transformed into rice variety ZH11 via Agrobacterium-mediated transformation method. After antibiotic screening, molecular characterization, salt-tolerance and trehalose content determinations, two transgenic lines resistant to 150 mM NaCL solutions were screened. Our study results indicated that the ZmTPS gene was integrated into the genomic DNA of the two transgenic rice lines and could be expressed well. Moreover, the detection of the transformed ZmTPS gene in the progenies of the two transgenic lines was performed from T₁ to T₄ generations; and results suggested that the transformed ZmTPS gene can be transmitted from parent to the progeny in transgenic rice.     

Experimental manipulation of sediment organic content and water column aeration reduces Zostera marina (eelgrass) growth and survival

The effect of synergy between sediment organic enrichment and lack of night oxygen renewal in the water column on growth and survival of Zostera marina, and how it is reflected in the sulfur parameters in the plants (δ³⁴S, TS and S⁰) was studied experimentally. An experiment consisting of Z. marina mesocosms with different levels of organic enrichment and water column aeration was established, and the effects on sediment conditions, sulfide invasion and growth and survival of Z. marina were examined over a 4 week period. Shoots growing in Ambient Organic matter-sediments showed signs of sulfide invasion, as TS increased in all plant compartments and δ³⁴S of the plant tissues decreased during the experiment, but neither growth rate nor survival were significantly affected. The lack of night oxygen renewal had no evident effects in non-enriched sediments as porewater sulfide concentrations, AVS- and CRS-pools were not different from the corresponding 24 h aeration treatment. Plant growth rate and survival were neither different from the corresponding 24 h aeration treatment. On the contrary, shoots growing on High Organic matter-sediments suffered a massive sulfide invasion and it was directly correlated to the observed decrease in growth rates. Even though the lack of night oxygen renewal had no evident effects on sediment variables there were, however, strong indications that the different aeration levels affected plant performance, suggesting a lower sulfide oxidation capacity and confirming that low water column oxygen concentrations reduces the defense capacity of the shoots against sulfide invasion.Although δ³⁴S, TS and S⁰ concentrations together provided a powerful set of indicators to detect the invasion of sulfide in Z. marina shoots, this study enlightens the need for a deeper investigation of sulfide intrusion in seagrasses and the relation between plant sulfur parameters and sediment conditions.     

Experimental test of biodeposition and ammonium excretion from blue mussels (Mytilus edulis) on eelgrass (Zostera marina) performance

Blue mussels and eelgrass have been found to coexist in many locations. However, knowledge of the interactions between these species is limited. Two experiments were conducted in the laboratory, a “Deposit” and an “Epiphyte” experiment. The Deposit experiment examined possible effects of increasing load of blue mussel (Mytilus edulis) biodeposits on sediment biogeochemistry and eelgrass (Zostera marina) performance. Z. marina mesocosms received normal or high loads of mussel biodeposits (Normal and High), while no biodeposits were added to the Control. High dosage had overall negative effects on Z. marina, which was reflected as lower leaf numbers and biomass and accumulation of elemental sulphur in rhizomes. The sediment biogeochemical conditions were altered, as the mussel biodeposits enhanced sulphate reduction rates and increased sulphide concentrations in the porewater, which may result in sulphide invasion and reduced growth of Z. marina.In the Epiphyte experiment effects of mussel excretion, with particular emphasis on ammonium, on the growth of Z. marina and their epiphytes were examined. A thick cover of epiphytes developed on Z. marina growing together with M. edulis, and the relative growth rate was reduced with 20% compared to plants from control without mussels. Overall the experiments showed negative effects on Z. marina growing together with M. edulis, thereby supporting a preceding field study by Vinther et al. [Vinther, H.F., Laursen, J.S., Holmer, M. 2008. Negative effects of blue mussel (Mytilus edulis) presence in eelgrass (Zostera marina) beds in Flensborg fjord, Denmark. Est. Coast Shelf. Sci. 77, 91-103.].     

Seasonal changes in the concentrations of Ca,Fe, K,Mg, Mn and Na in eelgrass (Zostera marina L.) in the Limfjord,Denmark

The seasonal changes in concentrations of calcium, iron, potassium, magnesium, manganese and sodium in above- and below-ground parts of eelgrass (Zostera marina L.) were studied at three locations in the Limfjord, Denmark.The concentrations of calcium, iron and manganese in eelgrass differed significantly at the three stations. Above-ground parts of eelgrass contained significantly higher concentrations of potassium and manganese than below-ground parts, whereas the concentrations of iron and sodium were highest in the below-ground parts. Calcium and mangnesium concentrations in the two fractions were not significantly different.Significant seasonal variations were observed in the concentrations of calcium, potassium, manganese and sodium in above-ground parts of eelgrass and in the concentrations of iron, magnesium and sodium in below-ground parts. Iron, potassium, manganese and, to an extent, sodium showed a similar seasonal pattern with maximum concentrations in the summer and minimum concentrations in the spring or fall, whereas calcium concentrations in the above-ground parts showed the opposite pattern.The observed seasonal variation patterns are discussed in relation to the respective metals and changes in environmental factors.