Size and estimated age of genets in eelgrass, Zostera marina, assessed with microsatellite markers

We examined the spatial distribution of genotypes in a perennial population of eelgrass, Zostera marina L., at two spatial scales. We mapped and sampled 80 ramets in a subtidal area of 20 × 80 m, and an additional 15 ramets in two 1-m² sub-quadrats. Ramets were genotyped for seven polymorphic microsatellite loci. Among a total number of 54 genotypes detected, 12 occurred more than once. The ramets of ten of these genotypes occurred in clusters and represented genets based on their expected multi-locus genotype frequencies. The size distribution of genets was uneven with estimated ramet numbers ranging from 2 to 5000. Whereas some areas displayed a high genet diversity, which may indicate past disturbances, large genets (≥10 m²) predominated in other locations of the sampled plot. Spatial heterogeneity in clone distribution was also obvious at the smaller sampling scale (15 ramets sampled within 1 m²) where the clonal diversity (genets identified/ramets sampled) was 0.24 in one quadrat, and 0.077 in the other. Ramets belonging to the largest clone were maximally 17 m apart, which corresponds to a genet age of 67 yr based on annual rhizome growth rates. We conclude that the spatial arrangement of clones in seagrasses allows inferences about the past demography and the disturbance regime at a given site which may prove useful for coastal zone management of ecologically valuable seagrass meadows. Received: 15 September 1998 / Accepted: 14 November 1998     

Significance of salinity and silicon levels for growth of a formerly estuarine eelgrass (Zostera marina) population (Lake Grevelingen, The Netherlands)

Since the early 1980s, the eelgrass, Zostera marina L., population in the saline Lake Gevelingen, The Netherlands, is rapidly declining. An earlier study, in which long-term data on eelgrass coverage in this former estuary were correlated with several environmental variables, showed only one significant correlation: coverage was positively related to water column silicon levels. In addition, a negative correlation with salinity was observed, but this was not significant. In the present study, the effect of silicon and the effect of salinity on the development of Z. marina were investigated experimentally. Enhancement of dissolved silicon concentrations in the water did not stimulate Z. marina above-ground production or an increase in final above- and below-ground biomass. The highly significant correlation between eelgrass coverage and water column silicon levels, thus, remains to be explained. The results of the growth experiments did, however, demonstrate a clear effect of salinity on Z. marina growth. Plants cultured at 22 psu showed a higher production of shoots and leaves, resulting in more above-ground biomass, than plants grown at 32 psu. In addition, below-ground biomass was also higher at 22 psu. Measurements of chlorophyll a fluorescence, performed with a PAM-fluorometer, indicated a reduction of photosynthesis in the high-salinity treatments. Thus, low salinity stimulates development of Z. marina from Lake Grevelingen. Eelgrass from such a historically estuarine area may be more sensitive to high salinities than other, more marine populations. Recovery of the autochthonous eelgrass population is expected to be favoured when the estuarine conditions of the seagrass area are re-established, or when restoration programmes are carried out with allochthonous ecotypes that are less sensitive to high salinities. Received: 23 June 1998 / Accepted: 19 November 1998     

Estimates of outcrossing rates and of inbreeding depression in a population of the marine angiosperm Zostera marina

Despite the great diversity of pollination and fertilization mechanisms observed in marine plants, little is known about the causes or maintenance of this variation. In this study, I estimated outcrossing rates and levels of inbreeding depression in Zostera marina L. (eelgrass), providing the first empirical test of hypotheses about the evolution of breeding systems in plants with submerged flowers. This study also addressed temporal separation of female and male flowering (dichogamy) in eelgrass as a mechanism promoting an outcrossing mating system, and whether the mating system in eelgrass is related to the degree of dichogamy in the field. Outcrossing rates (0<t<1) estimated from two polymorphic allozyme loci indicate that the Z. marina population in intertidal and subtidal habitats in False Bay, Washington, USA, was highly outcrossing in both 1991 (intertidal t=0.905, subtidal t=1.0) and 1992 (intertidal t=0.775, subtidal t=0.611). The outcrossing rates were positively associated with the degree of dichogamy in 1992; intertidal plants exhibited a greater temporal separation of female and male flowering and a higher outcrossing rate than did subtidal plants. Inbreeding depression at seed set was estimated from hand pollinations (self- and outcross) on 20 reproductive individuals from the False Bay population. Averaged across all maternal parents, a greater proportion of outcrossed flowers set seed than selfed flowers; i.e., inbreeding depression was detected. Plants exhibited genetic variation for inbreeding depression, detected as a significant pollination treatment × maternal family interaction in a log-likelihood analysis. By the end of the seed-maturation period (7 mo after intial seed set) some families showed outbreeding depression, i.e., greater fitness in progeny derived from selfing than in progeny from outcrossing. The inbreeding depression in the False Bay population may be an important selective factor contributing to the maintenance of dichogamy and an outcrossing mating system, as proposed for aquatic plants.     

Characterization of disjunct populations of Zostera marina (eelgrass) from California: genetic differences resolved by restriction-fragment length polymorphisms

Comparative restriction-fragment analysis was used to analyze the nuclear ribosomal DNA, and alcohol dehydrogenase-1 loci of Zostera marina L., for variation within and among populations. Eelgrass is a perennial marine flowering plant that is widespread and ecologically significant throughout the temperate northern hemisphere. A chemical method was developed to obtain restriction-quality DNA without CsCl fractionation from experimentally relevant quantities of seagrass tissues (0.5 to 1.0 g). The yield was 25 g g^-1 fresh weight. The three morphologically distinct forms of Z. marina from disjunct populations examined in this study were found to be genetically distinct; morphologically similar populations were indistinguishable genetically. Genetic distinction also correlated with habitat depth, as subtidal and intertidal populations were clearly divergent. Homologous probes for the 17S and 28S ribosomal DNA genes were used to map 24 restriction sites on the rDNA repeat of Z. marina, which was determined to be about 14 kb in length. At least 1 length mutation and 5 restriction-site changes were identified that distinguished Z. marina populations from San Diego and Monterey Bay (Del Monte Beach) from Z. marina populations from Elkhorn Slough and Tomales Bay. Estimated sequence variation (100×p) between eelgrass populations ranged from 0.00 to 0.69. Individual plants were observed to contain as many as four different rDNA-repeat length variants. The mean number of rDNA-repeat length variants per individual in Z. marina was about two. Intrapopulation variation in rDNA-repeat type was observed in only one individual from the Tomales Bay population.     

Effects of anaerobiosis on root metabolism of Zostera marina (eelgrass): implications for survival in reducing sediments

The temperate seagrass Zostera marina L. typically grows in highly reducing sediments. Photosynthesis-mediated O₂ supplied to below-ground tissues sustains aerobic respiration during photosynthetic periods. Roots, however, experience daily periods of anoxia and/or hypoxia at night and under conditions that reduce photosynthesis. Rhizosphere cores of Z. marina were collected in August 1984 from Great Harbor, Massachusetts, USA. We examined short-term anaerobic metabolism of [U-¹⁴C]sucrose in excised roots and roots of intact plants. Under anaerobic conditions roots showed appreciable labeling of CO₂, ethanol and lactate, and slight labeling of alanine and other metabolites. Over 95% of the ¹⁴C-ethanol was recovered in the root exudate. Release of other metabolites from the roots was minimal. Ethanol was also released from hypoxic/anoxic roots of intact plants and none of this ethanol was transported to the shoot under any experimental conditions. Loss of ethanol from roots prevented tissue levels of this phytotoxin from increasing during anaerobiosis despite increased synthesis of ethanol. Anaerobic metabolism of [U-¹⁴C]glutamate in excised roots led to appreciable labelling of -aminobutyrate, which was known to accumulate in eelgrass roots. Roots recovered to fully aerobic metabolism within 4 h after re-establishment of aerobic conditions. The contributions of these root metabolic responses to the ability of Z. marina to grow in reducing marine sediments are related to light-regulated interactions of shoots and roots that likely dictate depth penetration, distribution and ecological success of eelgrass.     

Irradiance reduction: Effects on standing crops of the eelgrass Zostera marina in a coastal lagoon

Abundance of the eelgrass Zostera marina L. was studied in a coastal lagoon in southern California (USA), and was found to correlate with the level of irradiance at depths greater than 0.5 m below tidal datum. Results of controlled field experiments, using canopies to reduce downwelling illuminance by 63%, confirmed that turion density is a function of the irradiance the plants receive. By Day 18 of the experiment, turion density in the shaded experimental areas had decreased compared to the density of adjacent unshaded controls. Turion densities were continually lower throughout the 9-month study in the experimental areas, which at the end of the study had a turion density only 5% that of the control areas. Flowering in the experimental areas was also inhibited by shading. The biological implications of these findings are discussed with respect to seasonal changes in incident solar radiation, water transparency, and changes in water quality due to man's increased intervention in the natural processes of coastal lagoons.Contribution No. 4 from the Center for Marine Studies, San Diego State University, San Diego, California 92182, USA.     

Sediment ammonium availability and eelgrass (Zostera marina) growth

The interaction of sediment ammonium (NH ₄ ⁺ ) availability and eelgrass (Zostera marina L.) growth, biomass and photosynthesis was investigated using controlled environment and in-situ manipulations of pore water ammonium concentrations. Sediment diffusers were used to create pore water diffusion gradients to fertilize and deplete ammonium levels in sediments with intact eelgrass rhizospheres. Between October, 1982 and September, 1983 controlled environment experiments using plants from shallow (1.3 m) and deep (5.5 m) stations in a Great Harbor, Woods Hole, Massachusetts, USA eelgrass meadow along with in-situ experiments at these stations provided a range of sediment ammonium concentrations between 0.1 and 10 mM (adsorbed+interstitial NH ₄ ⁺ ). The results of the in-situ experiments indicate that nitrogen limitation of eelgrass growth does not occur in the Great Harbor eelgrass meadow. A comparison of NH ₄ ⁺ regeneration rates and eelgrass nitrogen requirements indicates an excess of nitrogen supply over demand and provides an explanation for the lack of response to the manipulations. Results of controlled environment experiments combined with in-situ results suggest that sediment ammonium pool concentrations above approximately 100 mol NH ₄ ⁺ per liter of sediment (interstitial only) saturate the growth response of Zostera marina.     

Evidence that reef-wide patterns of coral bleaching may be the result of the distribution of bleaching-susceptible clones

The hypothesis that intraspecific variation in coral bleaching is a result of the distribution of bleaching-susceptible clonal genotypes (genets) was addressed using photoquadrats recorded during the 1987 Caribbean bleaching event on a reef dominated by Montastraea annularis (Morphotype I), together with manipulative experiments with Porites porites. Nearest-neighbor analysis showed that bleached colonies (ramets) of M. annularis at 10 m depth had a high probability (0.80) of having a nearest bleached neighbor of the same genet rather than a bleached ramet of a different genet. Furthermore, the frequency distributions of bleached ramets of M. annularis in the photoquadrats was significantly different from a Poisson distribution, suggesting that bleached ramets were aggregated on the reef. Manipulative experiments with P. porites from 15 m depth showed that some genets were more susceptible to thermal bleaching than others, since three genets had significantly different rates of zooxanthellae loss when exposed to elevated temperatures in tanks receiving irradiances similar to those found in situ. These results suggest that the in situ patchy distribution of bleached ramets could correspond to the distribution of certain genets, and that adjacent genets can exhibit sufficiently different phenotypes to account for intraspecific variation in bleaching. Further studies of genet-specific coral bleaching may provide valuable insights into the causes and consequences of bleaching.     

Genetic structure and gene flow of eelgrass <Emphasis Type="Italic">Zostera marina</Emphasis> populations in Tokyo Bay,Japan: implications for their restoration

Massive losses of eelgrass Zostera marina beds in Japan have occurred over the past 100 years. Toward their restoration, transplantation of eelgrass has been attempted in some areas, including Tokyo Bay. This study examined population genetic structures and gene flow in eelgrass in Tokyo Bay to establish guidelines for conducting restoration. Genotypes of a total of 360 individuals from 12 beds were determined using five microsatellite markers. The eelgrass beds in inner bay had above-average genetic diversity. A neighbor-joining tree based on F _ST values among beds revealed that a strong gene flow had occurred among six beds in the inner bay. Genetic assignment testing of drifting shoots indicated that those with seeds migrate in both directions between the inner and outer bay. We suggested that the restoration of eelgrass in the innermost part of Tokyo Bay, where natural habitats have been lost, should be conducted using the inner bay beds.     

Utilization of growth parameters of eelgrass,Zostera marina,for productivity estimation under laboratory and in situ conditions

Allometry was used for monitoring aboveground growth of the marine angiosperm Zostera marina L. Dry weight was regressed with leaf length and width, allowing estimation of aboveground net productivity and biomass of individual plants. At the termination of the experiment, rhizome productivity of the same plants was determined by harvesting. Plants in shaded and unshaded seawater tanks were monitored from June until September, 1976; in situ plants were also monitored at Point Judith Pond, Rhode Island, USA. Unshaded plants had shorter leaves, a lower net productivity, lower biomass, and a lower aboveground-torhizome productivity ratio than shaded plants. Unshaded plants had a higher rate of rhizome branching and the resulting new shoot formation than in situ plants.     

Ecology of two littoral species of caprellid amphipods (Crustacea) from Washington,USA

Ecology of 2 littoral species of caprellid amphipods is compared. Populations of Caprella laeviuscula, a periphyton scraper/filter-feeder, are most dense on eelgrass Zostera marina L. In the absence of C. laeviuscula, periphyton biomass increases 411% in protected Z. marina beds. The light absorption spectra of periphyton and Z. marina are similar, and C. laeviuscula, by its periphyton removal, may allow Z. marina to grow in areas where it would otherwise be excluded. C. laeviuscula is aggressively dominant over sympatric caprellids, and seasonal predators of C. laeviuscula are absent during winter, the period when periphyton would be most limiting to Z. marina. Populations of Deutella californica, a predator, are most dense on the hydroid Obelia dichotoma (L.). In the O. dichotoma epibiotic community, D. californica is the primary predator, but removal did not change the composition of the community structure. In the absence of other macropredators, the structure of the O. dichotoma epibiotic community depends more on the seasonality of O. dichotoma occurrence than on organismal interactions.     

Control of microbial growth and of amphipod grazing by water-soluble compounds from leaves of Zostera marina

The role of minor components of the leaves of Zostera marina L. in altering the activity of micro-organisms directly (and indirectly by affecting amphipod grazers) was investigated in laboratory experiments, using plants collected at Roberts Bank (49°2N; 123°8W) on the west coast of Canada. Water-soluble extracts of eelgrass leaves inhibited the growth of a micro-alga (Platymonas sp.) and many marine bacteria at concentrations equivalent to as little as 10 mg dry leaf l^-1. The potency of leaf extracts was higher (1) in young, actively metabolizing tissue than in older leaves, and (2) in leaves collected during rapid growth in summer 1980 than during the following winter. Water-soluble inhibitors (especially phenolic acids) may explain the low biomass of epiphytes on actively growing leaves. Three phenolic acids inhibited the test micro-organisms at concentrations as low as 0.3 mg l^-1; caffeic acid was more potent than either protocatechuic or gentisic acid. Extracts of young leaves also inhibited grazing by amphipods [Eogammarus confervicolus (Stimpson)] on dead leaves. The time required for leaching of soluble inhibitors may account for the delay between the loss of leaves from the plants and the onset of grazing. Thus, interactions among the biotic components of the detritus ecosystem may be significantly modified by minor compounds in the leaves of Z. marina     

Photosynthetic studies of Chondrus crispus

The net photosynthesis of intertidal, subtidal, carposporic, tetrasporic, and winter versus summer acclimatized plants of Chondrus crispus Stackhouse were evaluated under different temperatures and quantities of light. The optimum temperature and light conditions for net photosynthesis of C. crispus are seasonally and spatially variable, and there is an adaptive shift in the photosynthetic capacity at different seasons and positions on the shore. Plants collected during the fall and winter had lower light optima (465 to 747 ft-c) for net photosynthesis than spring and summer specimens (about 1000 ft-c). Intertidal populations exhibited a higher rate of net photosynthesis between 250 and 2819 ft-c than subtidal plants. Summer materials have a greater tolerance to high temperatures and a higher temperature optimum than winter materials. Shallow subtidal populations (-6m) exhibited a higher temperature optimum than deep subtidal plants (-12m). Tetrasporic plants (diploid) showed a higher rate of net photosynthesis than carposporic plants (haploid). It is suggested that the diploid plants of C. crispus may extend deeper in the subtidal zone, because they have a higher rate of net photosynthesis than carposporic plants. The results of the present studies are compared with previous physiological studies of C. crispus.Published with the approval of the Director of the New Hampshire Agriculture Experiment Station as Scientific Contribution Number 742.     

Uptake and translocation of phosphorus in eelgrass (Zostera marina)

The uptake and translocation of phosphorus in eelgrass (Zostera marina L.) was studied in two-compartment chambers using ³²P and autoradiography. Eelgrass was able to take up phosphorus both with leaves and rootrhizomes. Only a small amount (less than 4%) of the total amount of phosphorus taken up from either source was translocated within the plants during an incubation time of 120 h. Release of translocated phosphorus from the plant tissues to the surrounding water was not detected. The autoradiographs showed that the translocated phosphorus was localized mainly in the actively growing plant tissues, i.e. the stem and young leaves, and nodes of the rhizomes and young roots. The uptake of phosphorus in leaves and the bidirectional translocation in the plants was significantly lower in the dark. The uptake of phosphorus in rootrhizomes was unaffected by light-conditions. The concentration of phosphate in the water of one compartment did not affect the phosphorus uptake by the plant parts in the opposite compartment. However, the translocation was significantly reduced when both compartments were supplied with identical phosphate concentrations. The results of the present investigation indicate that the quantitative significance of eelgrass in the cycling of phosphorus between sediment and water may have been greatly overestimated in earlier studies. The amount of phosphorus taken up by root-rhizomes and subsequently translocated to leaves is probably insignificant during most of the growing season in comparison to the amount taken up by the leaves from the surrounding water. However the uptake of phosphorus by root-rhizomes may be significant in the nutrition of eelgrass during periods with low phosphate levels in the water.     

Increased macrophyte nitrate reductase activity as a consequence of groundwater input of nitrate through sandy beaches

While most marine macrophytes preferentially assimilate ammonium to meet growth demand for nitrogen, some also utilize nitrate and exhibit high nitrate reductase activity (NRA). Although nitrate concentrations are often low in coastal waters during the summer and sandy beaches are generally considered to be low nutrient-input habitats, we have observed elevated NRA in leaves of some eelgrass (Zostera marina L.) plants growing immediately adjacent to the shoreline. We postulated that nitrate may become available to eelgrass and macroalgae via groundwater inputs that enter the nearshore water column. To address this possibility, we investigated the availability of groundwater nitrate for the induction of NRA in the leaves of eelgrass and in the macroalgaeSargassum filipendula C. Agardh (Phaeophyceae) andEnteromorpha intestinalis L. Link (Chlorophyceae) collected adjacent to two sandy beaches in the vicinity of Woods Hole, Massachusetts, USA. Induction of NRA was determined in the laboratory for eelgrass collected from one of the beach sites and from an offshore site, Lackey's Bay, which is isolated from groundwater input. At the two beach locations, pore water nitrate concentrations were 100 to 400µM within a few meters inland from the waterline. Nitrate efflux into the nearshore water column was quite high and variable (2160±660µmol m^–2 h^–1) when associated with rapid percolation (37±11 1 m^–2 h^–1) of nitrate-enriched pore water. Turbulent wave mixing rapidly diluted the nitrate. Macroalgae and eelgrass growing adjacent to a beach with high nitrate efflux had NR activities three- to sevenfold higher than those of algae and eelgrass growing along a beach section with low nitrate efflux. NRA of eelgrass plants from Lackey's Bay and Great Harbor increased in response to low daily nitrate additions (10 to 25µM) in the laboratory, with higher nitrate additions (50 to 200µM) yielding less dramatic responses. The increase in NRA was roughly three times higher for Great Harbor than for Lackey's Bay eelgrass. It appears that groundwater input of nitrate is sufficient to induce NRA in marine macrophytes growing near some beaches, including those with turbulent wave mixing.     

Dynamics of benthic vegetation standing-stock,irradiance, and water properties in central Puget Sound

The standing stock of benthic macroalgae, sediment-associated microalgae and eelgrass (Zostera marina L.) was sampled in conjunction with irradiance and water properties from June 1982 through March 1984 to examine the relationship between the dynamics of benthic primary producers and environmental factors in central Puget Sound, Washington, USA. Sediment-associated microalgal standing stock (measured by chlorophylla) peaked in April and August. The seaweed assemblage, dominated by bladed green algae (e.g.Ulva fenestrata) and eelgrass exhibited maximum standing stocks in August. Although water temperature correlated best with changes in standing stock of all vegetation types, solar irradiance appeared to trigger the onset of biomass buildup and autumn die-back by the plants. Seasonal variations in dissolved oxygen reflected the buildup and loss of plant standing-stock. Nutrient concentrations, with the exception of ammonia, exhibited seasonal trends. Most nutrients were in greatest concentration in winter and reached minimum concentration in late spring-summer. Regeneration of nutrients in autumn followed shortly after the autumn dieback of the benthic vegetation. We concluded that irradiance was the primary controlling factor in the system. Nutrient limitation (primarily nitrate) may control standing-stock accumulations from the period May–October when light is not limiting. In contrast to phytoplankton systems in deep portions of Puget Sound, shallow nearshore systems may be more susceptible to the effects of increased inorganic nutrient-loadings from anthropogenic sources.Contribution No. 808, School of Fisheries, University of Washington     

Production dynamics of the eelgrass, <Emphasis Type="Italic">Zostera marina</Emphasis> in two bay systems on the south coast of the Korean peninsula

Production dynamics of eelgrass, Zostera marina was examined in two bay systems (Koje Bay and Kosung Bay) on the south coast of the Korean peninsula, where few seagrass studies have been conducted. Dramatically reduced eelgrass biomass and growth have been observed during summer period on the coast of Korea, and we hypothesized that the summer growth reduction is due to increased water temperature and/or reduced light and nutrient availabilities. Shoot density, biomass, morphological characteristics, leaf productivities, and tissue nutrient content of eelgrass were measured monthly from June 2001 to April 2003. Water column and sediment nutrient concentrations were also measured monthly, and water temperature and underwater irradiance were monitored continuously at seagrass canopy level. Eelgrass shoot density, biomass, and leaf productivities exhibited clear seasonal variations, which were strongly correlated with water temperature. Optimal water temperature for eelgrass growth in the present study sites was about 15–20°C during spring period, and eelgrass growths were inhibited at the water temperature above 20°C during summer. Daily maximum underwater photon flux density in the study sites was usually much higher than the light saturation point of Z. marina previously reported. Densities of each terminal, lateral, and reproductive shoot showed their unique seasonal peak. Seasonal trends of shoot densities suggest that new eelgrass shoots were created through formation of lateral shoots during spring and a part of the vegetative shoots was transformed into flowering shoots from March. Senescent reproductive shoots were detached around June, and contributed to reductions of shoot density and biomass during summer period. Ambient nutrient level appeared to provide an adequate reserve of nutrient for eelgrass growth throughout the experimental period. The relationships between eelgrass growth and water temperature suggested that rapid reductions of eelgrass biomass and growth during summer period on the south coast of the Korean peninsula were caused by high temperature inhibition effects on eelgrass growth during this season.     

Inhibition of the growth of micro-algae and bacteria by extracts of eelgrass (Zostera marina) leaves

The effects of the water-soluble fraction of dead leaves of the eelgrass Zostera marina L. on the growth of 8 species of micro-algae (pennate and centric diatoms, dinoflagellates, and a green flagellate) and a bacterium were studied on agar plates and in liquid culture. The extracts of leaves which had been dead from a few days to 2 wk inhibited growth and often killed cells in all test organisms. Extracts were lethal even at concentrations equivalent to as little as 0.25 mg dry leaf ml^-1, but inhibition decreased when extracts were prepared from leaves aged in the laboratory for 35 d (loss of anti-bacterial activity) or 90 d (loss of anti-algal activity). Extracts of leaves which had aged and dried several months in the field had no effect, except at very high concentrations (13 mg dry leaf ml^-1) when the lag phase of growth was prolonged several days in a culture of the chlorophyte Platymonas sp. The active fraction in eelgrass leaves may be important in controlling initial growth of micro-organisms on eelgrass detritus, and it could determine the composition and activity of the epiphytic community on living leaves.     

Nitrate reductase activity in Zostera marina

Eelgrass (Zostera marina L.) has access to nutrient pools in both the water column and sediments. We investigated the potential for eelgrass to utilize nitrate nitrogen by measuring nitrate reductase (NR) activity with an in vivo tissue assay. Optimal incubation media contained 60 mM nitrate, 100 mM phosphate, and 0.5% 1-propanol at pH 7.0. Leaves had significantly higher NR activity than roots (350 vs 50 nmoles NO ₂ ^– produced g FW^–1 h^–1). The effects of growing depth (0.8 m MLW, 1.2 m, 3.0 m, 5.0 m) and location within the eelgrass meadow (patch edge vs middle) on NR activity were examined using plants collected from three locations in the Woods Hole area, Massachusetts, USA, in July 1987. Neither depth nor position within the meadow appear to affect NR activity. Nitrate enrichment experiments (200 M NO ₃ ^– for 6 d) were conducted in the laboratory to determine if NR activity could be induced. Certain plants from shallow depth (1.2 m) showed a significant response to enrichment, with NR activity increasing from >100 up to 950 nmoles NO ₂ ^– g FW^–1 h^–1 over 6 d. It appears that Z. marina growing in very shallow water (0.8 m) near a shoreline may be affected by ground water or surface run-off enrichments, since plants from this area exhibited rates up to 1 600 nmol NO ₂ ^– g FW^–1 h^–1. Water samples from this location consistently had slightly higher NO ₃ ^– concentrations (1.4 M) than all other collection sites (0.7 M). Thus, it is possible that chronic run-off or localized groundwater inputs can create sufficient NO ₃ ^– enrichment in the water column to induce nitrate reductase activity in Zostera leaves.     

Predictive modelling of eelgrass (Zostera marina) depth limits

Empirical models relating secchi depths to maximum depth limits of eelgrass (Zostera marina L.) can describe basic differences in depth limits between areas or time periods exhibiting large differences in secchi depth. However, these models cannot predict the precise depth limit at a particular site at any specific time. In this study we aim to improve the ability of regression models to predict maximum depth limits by: (1) assuming that eelgrass depth limits respond to changes in secchi depth with a temporal delay of 1–2 years, (2) including other water-quality variables in addition to secchi depth, and (3) taking into account that factors regulating depth limits may vary between years and between sites. We were not able to improve the models by introducing a systematic delay in the response of depth limits to changes in secchi depths. The reason for this failure is likely to have been the systematic nature of our approach, since some sites responded with a delay, while others did not. The explanatory power of the models increased when additional water-quality variables were added in a multiple regression model. Where secchi depth alone explained 58% of the variations in depth limits, addition of winter [NH₄⁺] and maximum water depth as independent variables increased the explanatory power to 71%. These models applied to data from one specific year, but when data from several years (1989–1998) were included, only 35% of the variation in depth limits could be explained by the three factors. More detailed analyses showed that the regulation of eelgrass depth limits varied considerably between years and between sites, and the models were further improved by taking this information into account. Our results confirmed previous studies by showing light to be the most important parameter in the regulation of eelgrass depth limits, but also revealed a complexity in the regulation of depth limits not expressed in earlier studies. Limited colonisation potentials may delay the response to improved light conditions, and hypoxia/anoxia and indirect effects of nutrients may prevent eelgrass from attaining the depth limit that light levels would allow. The power to predict depth limits on the basis of secchi depths can therefore be improved by taking site-specific information on eelgrass growth conditions into account.Communicated by M. Kühl, Helsingør