Variable responses of native eelgrass Zostera marina to a non-indigenous bivalve Musculista senhousia

The transport and establishment of non-indigenous species in coastal marine environments are increasing worldwide, yet few studies have experimentally addressed the interactions between potentially dominant non-native species and native organisms. We studied the effects of the introduced mussel Musculista senhousia on leaf and rhizome growth and shoot density of eelgrass Zostera marina in San Diego Bay, California. We added M. senhousia over a natural range in biomass (0–1200 g dry mass/m²) to eelgrass in transplanted and established beds. The effects of the non-indigenous mussel varied from facilitation to interference depending on time, the abundance of M. senhousia, and the response variable considered. Consistent results were that mussel additions linearly inhibited eelgrass rhizome elongation rates. With 800 g dry mass/m² of M. senhousia, eelgrass rhizomes grew 40% less than controls in two eelgrass transplantations and in one established eelgrass bed. These results indicate that M. senhousia, could both impair the success of transplantations of eelgrass, which spread vegetatively by rhizomes, and the spread of established Z. marina beds to areas inhabited by M. senhousia. Although effects on leaf growth were not always significant, in August in both eelgrass transplantations and established meadows leaf growth was fertilized by mussels, and showed a saturation-type relationship to sediment ammonium concentrations. Ammonium concentrations and sediment organic content were linear functions of mussel biomass. We found only small, non-consistent effects of M. senhousia on shoot density of eelgrass over 6-month periods. In established eelgrass beds, but not in transplanted eelgrass patches (≈0.8 m in diameter), added mussels suffered large declines. Hence, eelgrass is likely to be affected by M. senhousia primarily where Z. marina beds are patchy and sparse. Our study has management and conservation implications for eelgrass because many beds are already seriously degraded and limited in southern California where the mussel is very abundant. Received: 31 May 1997 / Accepted: 4 September 1997     

Factors Influencing Seedling Establishment Rates in Zostera marina and Their Implications for Seagrass Restoration

Selection of strategies to efficiently utilize limited seed supplies in efforts to restore the seagrass Zostera marina (eelgrass) requires a better understanding of the processes that limit seedling establishment at potential restoration sites. We investigated the effect of seed distribution timing on seedling establishment and tested for interactive effects of seed burial and distribution timing. We also investigated the effect of habitat type on seedling establishment by distributing Z. marina seeds inside and outside of established Ruppia maritima (widgeongrass) patches and examined mechanisms causing habitat differences by manipulating seed position (buried or unburied) and vulnerability to seed predators (unprotected or protected in packets). Seeds distributed on the sediment surface in the summer (July or August) produced fewer seedlings than seeds distributed in fall (October) in five of six trials over 3 years. Seed burial increased success rates for seeds distributed in summer at one of two sites tested, eliminating the effect of season, but reduced success at the other site. Seeds placed in R. maritima generally produced fewer seedlings than seeds in bare sand, and although seed burial and protection in packets increased success in bare sand at three of four sites, the effect was less consistent in R. maritima. We conclude that seed predation and physical interactions were influential in reducing seedling establishment in R. maritima, contrary to hypotheses positing a nursery role for existing vegetation. Efficient restoration efforts with Z. marina seeds should target unvegetated areas after summertime sources of mortality have diminished. Direct seed burial may enhance seedling establishment rates.     

Large‐Scale Zostera marina (eelgrass) Restoration in Chesapeake Bay,Maryland, USA. Part II: A Comparison of Restoration Methods in the Patuxent and Potomac Rivers

In response to systemic losses of submerged aquatic vegetation (SAV) in the Chesapeake Bay (east coast of North America), the U.S. Environmental Protection Agency's (EPA) Chesapeake Bay Program (CBP) and Maryland Department of Natural Resources (MD DNR) have considered SAV restoration a critical component in Bay restoration programs. In 2003, the CBP created the “Strategy to Accelerate the Protection and Restoration of Submerged Aquatic Vegetation in the Chesapeake Bay” in an effort to increase SAV area. As part of this strategy, large‐scale eelgrass (Zostera marina) restoration efforts were initiated in the Patuxent and Potomac Rivers in Maryland. From 2004 to 2007, nearly 4 million Z. marina seeds were dispersed over 10 ha on the Patuxent River and almost 9 million seeds over 16 ha on the Potomac River. Z. marina seedling establishment was consistent throughout the project (<4%); however, restored eelgrass survival was highly dependent on restoration site. Restoration locations on the Patuxent River experienced initial Z. marina seedling germination, but no long‐term plant survival. Restored Z. marina on the Potomac River has persisted and expanded, both vegetatively and sexually, beyond initial seeding areas. Healthy Z. marina beds now cover approximately five acres of the Potomac River bottom for the first time in decades. The differential success of Z. marina restoration efforts in the two rivers is evidence for the necessity of carefully considering site‐specific characteristics when using large‐scale seeding methods to achieve successful SAV restoration.     

The Roles of Competition and Disturbance in a Marine Invasion

Two hypotheses for the decline of native species are the superior exploitation of disturbance by exotic species and the competitive displacement of native species by their exotic counterparts. Theory predicts that functional similarity will increase the intensity of competition between native and invasive species. Ecologically important “foundation” species, Zostera marina and other seagrasses have globally declined during the past century. This study used transplant and vegetation removal experiments to test the hypotheses that disturbance and competitive interactions with an invasive congener (Z. japonica) are contributing to the decline of native Z. marina in the northeastern Pacific. Interspecific competition reduced Z. marina and Z. japonica above-ground biomass by 44 and 96%, respectively, relative to intraspecific competition. Disturbance substantially enhanced Z. japonica productivity and fitness, and concomitantly decreased Z. marina performance, effects that persisted two years following substratum disturbance. These results demonstrate that disturbance and competitive interactions with Z. japonica reduce Z. marina performance, and suggest that Z. japonica’s success as an invasive species stems dually from its ability to persist in competition with Z. marina and its positive response to disturbance. These results highlight the importance of understanding the interconnected roles of species interactions and disturbance in the decline of seagrass habitats, and provide a rationale for amending conservation policy in Washington State. In the interest of conserving native eelgrass populations, the current policy of protecting both native and invasive Zostera spp. should be refined to differentiate between native and invader, and to rescind the protection of invasive eelgrass.     

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.].     

Effects of copper enrichment on survival,growth and photosynthetic pigment of seedlings and young plants of the eelgrass Zostera marina

Copper (Cu²⁺) is an essential nutrient for plants but toxic at high concentrations. We subjected seedlings and young plants of eelgrass Zostera marina to different seawater Cu concentrations (3, 4, 5, 10, 30 and 50?µg?l^?1) for over 30 days under controlled laboratory conditions. Natural seawater without added Cu (3?µg?l^?1) was used as reference seawater. We measured plant response in terms of survivorship, morphology, growth, productivity and leaf pigment concentration. Survival analysis combined with morphological, dynamic and productive assessment suggested that the optimum seawater Cu concentration for the establishment of Z. marina seedlings and young plants is 4?μg?l^?1. The photosynthetic response of young plants to copper enrichment, including an increase in chlorophyll content under low Cu concentration treatment but significant decrease when treated with high concentrations of Cu, is similar to those reported for other seagrass species. NOEC (no observed effect concentration), LOEC (lowest observed effect concentration) and LC₅₀ (lethal concentration that caused an increase in mortality to 50% of that of the control) values of seedlings were significantly lower than those of young plants, implying a reduced Cu tolerance to high concentrations (>10?μg?l^?1). This study provides data that could prove helpful in the development of successful eelgrass restoration and conservation.     

Eelgrass (Zostera spp.) associated phytomyxids are host-specific congeneric parasites and predominant eukaryotes in the eelgrass rhizosphere on a global scale

Together with increasing environmental and anthropogenic pressures, pathogenic diseases are one of the important factors contributing to the ongoing decline of seagrass meadows worldwide; yet the diversity and ecology of the microorganisms acknowledged as seagrass parasites remain critically understudied. Here, we investigate phytomyxid parasites (Rhizaria: Endomyxa: Phytomyxea) of three different eelgrass (Zostera spp.) species found in the Northern hemisphere. We present molecular evidence that Plasmodiophora bicaudata, a long-recognized parasite of dwarf eelgrass taxa, is closely related to the novel phytomyxid recently discovered in root hairs of Zostera marina, and together they form a distinct clade within the order Phagomyxida, proposed here as Feldmanniella gen. nov. A full life cycle is systematically described in a phagomyxid representative for the first time, proving its conformity with the generalized phytomyxid life history, despite previous uncertainties. The presence of primary infection stages in nearly all collected eelgrass specimens, and subsequent analysis of amplicon sequences from a global Z. marina dataset, reveal phytomyxids to be ubiquitous and one of the predominant microeukaryotes associated with eelgrass roots on a global scale. Our discoveries challenge the current view of Phytomyxea as rare entities in seagrass meadows and suggest their generally low pathogenicity in natural ecosystems.     

Form–function relationships in a marine foundation species depend on scale: a shoot to global perspective from a distributed ecological experiment

Form–function relationships in plants underlie their ecosystem roles in supporting higher trophic levels through primary production, detrital pathways, and habitat provision. For widespread, phenotypically‐variable plants, productivity may differ not only across abiotic conditions, but also from distinct morphological or demographic traits. A single foundation species, eelgrass Zostera marina, typically dominates north temperate seagrass meadows, which we studied across 14 sites spanning 32–61°N latitude and two ocean basins. Body size varied by nearly two orders of magnitude through this range, and was largest at mid‐latitudes and in the Pacific Ocean. At the global scale, neither latitude, site‐level environmental conditions, nor body size helped predict productivity (relative growth rate 1–2% day^‐1 at most sites), suggesting a remarkable capacity of Z. marina to achieve similar productivity in summer. Furthermore, among a suite of stressors applied within sites, only ambient leaf damage reduced productivity; grazer reduction and nutrient addition had no effect on eelgrass size or growth. Scale‐dependence was evident in different allometric relationships within and across sites for productivity and for modules (leaf count) relative to size. Zostera marina provides a range of ecosystem functions related to both body size (habitat provision, water flow) and growth rates (food, carbon dynamics). Our observed decoupling of body size and maximum production suggests that geographic variation in these ecosystem functions may be independent, with a future need to resolve how local adaptation or plasticity of body size might actually enable more consistent peak productivity across disparate environmental conditions.     

Habitat suitability of the Wadden Sea for restoration of Zostera marina beds

A conceptual model is proposed, describing potential Zostera marina habitats in the Wadden Sea, based on reported data from laboratory, mesocosm and field studies. Controlling factors in the model are dynamics, degree of desiccation, turbidity, nutrients and salinity. A distinction has been made between a higher and a lower zone of potential habitats, each suitable for different morphotypes of Z. marina. The model relates the decline of Z. marina in the Wadden Sea to increased sediment and water dynamics, turbidity, drainage of sediments (resulting in increased degree of desiccation) and total nutrient loads during the twentieth century. The upper and lower delineation of both the higher and the lower zone of potential Z. marina habitats appear to be determined by one or a combination of several of these factors. Environmental changes in one of these factors will therefore influence the borderlines of the zones. The lower zone of Z. marina will be mainly affected by increased turbidity, sediment dynamics, degree of desiccation during low tide and nutrient load. The higher zone will be affected by increases in water and sediment dynamics, desiccation rates and nutrient loads. Potential Z. marina habitats are located above approx. –0.80 m mean sea level (when turbidity remains at the same level as in the early 1990s) in sheltered, undisturbed locations, and preferably where some freshwater influence is present. At locations with a high, near-marine, salinity, the nutrient load has to be low to allow the growth of Z. marina. The sediment should retain enough water during low tide to keep the plants moist. Our results suggest that the return of Z. marina beds within a reasonable time-scale will require not only suitable habitat conditions, but also revegetation measures, as the changes in the environment resulting from the disappearance of Z. marina may impede its recovery, and the natural import of propagules will be unlikely. Furthermore, the lower zone of Z. marina may require a genotype that is no longer found in the Wadden Sea. Received: 26 April 1999 / Received in revised form: 15 October 1999 / Accepted: 16 October 1999     

Top-down impact through a bottom-up mechanism: the effect of limpet grazing on growth,productivity and carbon allocation of Zostera marina L. (eelgrass)

The unusual appearance of a commensal eelgrass limpet [Tectura depicta (Berry)] from southern California at high density (up to 10 shoot^–1) has coincided with the catastrophic decline of a subtidal Zostera marina L. meadow in Monterey Bay, California. Some commensal limpets graze the chloroplast-rich epidermis of eelgrass leaves, but were not known to affect seagrass growth or productivity. We evaluated the effect on eelgrass productivity of grazing by limpets maintained at natural densities (8±2 shoot^–1) in a natural light mesocosm for 45 days. Growth rates, carbon reserves, root proliferation and net photosynthesis of grazed plants were 50–80% below those of ungrazed plants, but biomass-specific respiration was unaffected. The daily period of irradiance-saturated photosynthesis (H _sat) needed to maintain positive carbon balance in grazed plants approached 13.5 h, compared with 5–6 h for ungrazed plants. The amount of carbon allocated to roots of ungrazed plants was 800% higher than for grazed plants. By grazing the chlorophyll-rich epidermis, T. depicta induced carbon limitation in eelgrass growing in an other-wise light-replete environment. Continued northward movement of T. depicta, may have significant impacts on eelgrass production and population dynamics in the northeast Pacific, even thought this limpet consumes very little plant biomass. This interaction is a dramatic example of top-down control (grazing/predation) of eelgrass productivity and survival operating via a bottom-up mechanism (photosynthesis limitation).     

Photoadaptation and growth of Zostera marina L. (eelgrass) transplants along a depth gradient

Photosynthetic and growth responses were assessed in Zostera marina L. transplants within and beyond the natural extent of an eelgrass meadow in Great Harbor, Woods Hole, MA. Transplant survival and rapid growth inshore of the shallow edge of the meadow (0.5 and 0.8 m water depth) indicated a periodic disturbance factor maintaining the shallow edge of the meadow. Transplant mortality, reduced growth, and a negative carbon balance of eelgrass transplanted offshore the deep edge of the meadow (7 and 10m) supported the hypothesis of light-limited eelgrass growth in the deep regions of the Great Harbor meadow. Photoadaptive responses occurred along the water depth gradient, and both photosynthesis and growth responses were used to assess the genetic vs. phenotypic components of eelgrass response to the water depth gradient. Reciprocal transplants between shallow (1.3 m) and deep (5.5 m) areas within the eelgrass meadow indicated photosynthetic and growth responses were primarily a result of growth habitat rather than genetic differentiation within the eelgrass meadow.     

Characterization of microsatellite loci in the dwarf eelgrass Zostera noltii (Zosteraceae) and cross‐reactivity with Z. japonica

Zostera noltii is an important species of eelgrass occurring along European, north African, Mediterranean, Black Sea and Azov Sea coasts. Nine microsatellite loci were developed and no linkage disequilibrium was observed. Cross‐amplification was observed for all loci (polymorphic) in Z. japonica; only four loci amplified (monomorphic) in Z. marina.     

An analysis of the population genetics of restored Zostera marina plantings in Barnegat Bay, New Jersey

Within Barnegat Bay, New Jersey, eelgrass (Zostera marina) populations have declined by 62 % over the last 20 years. To better understand the consequences of this devastation, we have previously employed microsatellite DNA polymorphisms to analyze the population structure of Z. marina within Barnegat Bay, as well as along the eastern United States seaboard. We have restored populations of Z. marina in Barnegat Bay over the last 10 years to help assess the best planting conditions and ecotypes that might be used in long-term restoration strategies. In this study, we examined the genetic health of the restored populations compared to that of the donor eelgrass populations within the bay. Using microsatellites, we can identify which parental founding ecotypes survived the restoration process over multiple generations. The frequency of observed heterozygotes, although higher than in the natural populations, still indicates reduced levels of diversity and connectivity. The inbreeding frequency is high in the restored populations, but lower than what is seen in the native populations. All restored populations have effective population values >50, suggesting a high probability of survival in the short term.     

Small-scale removal of seagrass (Zostera marina L.): effects on the infaunal community

Eelgrass meadows are a common feature in shallow waters along the Norwegian coast, where they provide a habitat for a diverse infaunal community. Recreational boat anchoring and moorings physically scour seagrass and may affect the ecosystem functioning and resilience of the system to natural and anthropogenic disturbances. A small-scale eelgrass (Zostera marina) removal experiment was conducted to study the effects on macro- and meiofauna. Entire plants, including the rhizomes, were removed from 4?m² patches in three eelgrass meadows in the inner Oslofjord in October 2010. Core samples were taken after a recovery period of 10 months, from the removed patches as well as from the surrounding meadow. Macrofauna (>500?μm) and meiofauna (63–500?μm) in the sediment were investigated for possible effects of the eelgrass removal. Macrofauna and meiofauna composition were site specific and therefore location was identified as the main determinant for the infaunal community. The eelgrass did not regrow within the recovery period and bare sediment patches with only single eelgrass shoots were present during the sampling. Our analyses support an influence of the removal on individual species, but not the complete community. In particular one species, the gastropod Peringia ulvae, was encountered in higher numbers in samples from the removed patches than in control samples. From a management perspective, such minor removal of eelgrass, on the scale of square metres, appears to have no long-lasting detrimental effect to the infaunal community in sheltered meadows with muddy sediments.     

Landscape-level seagrass–sediment relations in a coastal lagoon

We investigated the relationships between sediment (subaqueous soil) properties and eelgrass (Zostera marina L.) distribution to develop landscape-level soil-based strategies for choosing eelgrass restoration locations. Subaqueous soils were sampled and eelgrass cover determined for 14 soil-landscape units within a 116 ha area of Ninigret Pond, a coastal lagoon in Rhode Island, USA. Of the 14 soil-landscape units sampled for eelgrass cover, 52% had virtually no eelgrass cover (<10%), while 18% had high eelgrass cover (>90%). The Lagoon Bottom, Shallow Lagoon Bottom, Flood-tidal Delta Slope, and Barrier Cove subaqueous soil-landscape units had the highest eelgrass cover (66–100%). A weak relationship between eelgrass cover and water depths (r² = 0.10) was observed suggesting that properties other than water depth may also control eelgrass distribution. Subaqueous soils on landscapes with >60% eelgrass cover had relatively high levels of acid-volatile sulfides (>90 μg/g), high soil salinity levels (34–44 ppt), fine textures (silt loam), and relatively high total nitrogen levels (>0.15%). Four principal components accounted for 81% of the variability in eelgrass cover. The first component reflected particle-size distribution (i.e. sand, silt, and clay contents) effects and accounted for 43% of the variability. The other components suggested that eelgrass cover is correlated to carbonaceous remains, non-calcareous rock fragments and soil salinity. These data suggest that the current distribution of eelgrass within the study area is strongly influenced by physical and chemical subaqueous soil characteristics. Soil survey techniques proved useful for the delineation of sediment characteristics (e.g. texture, salinity) that influence eelgrass distribution patterns at landscape-level scales.     

Growing Zostera marina (eelgrass) from Seeds in Land‐Based Culture Systems for Use in Restoration Projects

The use of aquaculture systems to grow the seagrass Zostera marina (eelgrass) from seeds for restoration projects was evaluated through laboratory and mesocosm studies. Along the mid‐Atlantic coast of North America Z. marina seeds are shed from late spring through early summer, but seeds typically do not begin to germinate until the late fall. Fall is the optimal season to plant both seeds and shoots in this region. We conducted studies to determine if Z. marina seeds can be induced to germinate in the summer and seedlings grown in mesocosms to a size sufficiently large enough for out‐planting in the fall. Seeds in soil‐less culture germinated in the summer when held at 14°C, with percent germination increasing with lower salinities. Cold storage (4°C) of seeds prior to planting in sediments enhanced germination and seedling survival. Growth rates of seedlings were significantly higher in nutrient enriched estuarine sediments. Results from preliminary studies were used in designing a large‐scale culture project in which 15,000 shoots were grown and out‐planted into the Potomac River estuary in the Chesapeake Bay and compared with an equal number of transplanted shoots. These studies demonstrate that growing Z. marina from seeds is an alternative approach to harvesting plants from donor beds when vegetative shoots are required for restoration projects.     

The role of seedlings and seed bank viability in the recovery of Chesapeake Bay,USA, <Emphasis Type="Italic">Zostera marina</Emphasis> populations following a large-scale decline

The objective of this study was to quantify the spatial and temporal recolonization characteristics of Zostera marina beds in the lower Chesapeake Bay following large scale declines in the late summer of 2005. Transects were established and monitored monthly for changes in eelgrass abundance at three sites (two downriver, one upriver) in the York River from April–October 2006 and 2007. Measurements included percent bottom cover, above ground biomass, shoot density, shoot origin (seedling or vegetative), seed bank abundance and seed viability. During 2006, the eelgrass beds at all sites recovered with seedlings providing the largest proportion of the total shoot abundance. This trend shifted in 2007 and surviving vegetative shoots were the dominant component of shoot standing crop. A second consecutive decline related to low light conditions occurred during the summer of 2006 in the upriver site and recovery there was minimal in 2007. These results highlight that after a single die off event, seed germination with subsequent seedling growth is the principal method for revegetation in lower Chesapeake Bay Z. marina beds. However, no viable seeds remain in the seed bank during this first year of recovery and shoots produced by the seedling growth do not flower and produce seeds until their second year of growth. Therefore the seed-bank density is low and is not immediately replenished. This suggests that the resiliency of perennial Chesapeake Bay Z. marina populations to repeated disturbances is restricted and repeated annual stress may result in much longer term bed loss.     

Large‐Scale Zostera marina (eelgrass) Restoration in Chesapeake Bay,Maryland, USA. Part I: A Comparison of Techniques and Associated Costs

The Chesapeake Bay, like many other temperate estuaries, has exhibited dramatic declines in the abundance of submerged aquatic vegetation (SAV) during the later half of the twentieth century. Because of the functions SAV serve in maintaining a healthy estuarine ecosystem, SAV restoration has become an important component of Chesapeake Bay restoration. Specifically, recent water quality improvements in areas from which populations of Zostera marina (eelgrass) have been extirpated have suggested that Z. marina restoration could succeed. Early restoration efforts involved transplanting Z. marina plants from healthy source beds to restoration locations, but this was labor intensive, time consuming, expensive, and potentially detrimental to donor beds. This multi‐year project investigated new techniques for large‐scale Z. marina seed collection and processing and compared two seed dispersal methods to evaluate cost effectiveness. Tens of millions of mature Z. marina seeds were collected through snorkeling, SCUBA, or with a mechanical harvester. Seed storage conditions and processing techniques were manipulated in order to maximize seed yield. Seeds were dispersed using two methods: spring seed buoys and fall seed broadcasts. Our costs for planting 1 ha of bottom with Z. marina seeds ranged from $6,674 to $165,699 depending on seeding density and dispersal method used. The average cost per Z. marina seed was $0.17. Interannual variations in seed collection yield and seed viability after summer storage had great impact on final costs. Our results suggest that the use of seeds for large‐scale Z. marina restoration offers a competitive advantage to more traditional transplanting methods.     

EPIPHYTE-GRAZER INTERACTIONS ON ZOSTERA MARINA (ANTHOPHYTA: MONOCOTYLEDONES): EFFECTS OF DENSITY ON COMMUNITY FUNCTION1

Interactions between algal epiphytes and their grazers can have a significant impact on the structure and function of eelgrass (Zostera marina L.) meadows. In Puget Sound, the herbivorous gastropod Lacuna variegata Carpenter and its congeners appear to remove large quantities of the epiphytic community from eelgrass blades. When snails at typical field densities were used in microcosms, Lacuna significantly reduced epiphytic biomass and areal productivity. Biomass-specific productivity of the epiphytic community showed an increasing trend with increasing snail density. Epiphytic productivity increased nonlinearly with increasing epiphytic biomass. The commonly used logistic population growth formula adequately described this relationship. Grazing rate also increased nonlinearly with increasing epiphytic biomass. The Hailing equation adequately described the relationship between grazing rate and epiphytic biomass. The proportion of the epiphytic biomass found on the oldest blade of an eelgrass shoot was related linearly to epiphytic biomass, suggesting that a constant fraction of the epiphytic community is lost regardless of epiphytic density. Lacuna clearly removed large quantities of epiphytic material from eelgrass blades, significantly altering community function. Modified Lotka-Volterra equations, incorporating the logistic growth form and Hailing grazing equation, should prove useful in modeling the epiphyte–grazer interaction.     

Simultaneous measurements of H+ and O2 fluxes in Zostera marina and its physiological implications

Zostera marina (eelgrass) is an important ecological component of many shallow, temperate lagoons. Evidence suggests that Z. marina has a high bicarbonate utilization capability, which could be promoted by possible proton extrusion and the consequent formation of an ‘acid zone’ in the apoplastic space (unstirred layer) of its leaves. It has been found that 50 mM of the buffer Tris significantly inhibited the photosynthetic O₂ evolution of Z. marina and it was proposed that this was because of Tris's ability to bond with protons outside the cell wall. To investigate if H⁺ played an important role in the photosynthetic carbon utilization of Z. marina, it is very important to simultaneously monitor the photosynthesis status and possible H⁺ fluxes. However, probably because of the lack of suitable techniques, this has never been attempted. In this study, experiments were undertaken on Z. marina by monitoring H⁺ and O₂ fluxes and the relative electron transport rates during light–dark transition. During stable photosynthesis, in addition to an obvious O₂ outflow, there was a significant net H⁺ influx connected to Z. marina photosynthesis. The inhibitory effects of both Tris and respiration inhibitors on apparent O₂ evolution of Z. marina were confirmed. However, evidence did not support the proposed Tris inhibition mechanism.