Nitrogen Assimilation and Partitioning in the Mediterranean Seagrass Posidonia oceanica

Abstract. The supply of nitrogen (N) often limits the productivity of marine macrophytes. In vitro and in vivo assays of glutamine synthetase (GS) activity were employed to investigate patterns of N assimilation by the Mediterranean seagrass Posidonia oceanica (L.) DELILE. Biomass-specific GS activity wa measured in root tissue, in leaves within a shoot, in shoots collected at two sites during two season and over a depth range of 5–33 m. Root tissue was less important than shoot tissue in assimilating inorganic N in P. oceanica , due both to the small roots' biomass (ca. 3% of total plant biomass and greatly lower (10- to 50-fold) GS activities. While the GS activity and N assimilatory potentia (biomass × GS activity; μmol N-h-^-1) were greatest in leaf 2, leaves 1 and 3–5 assimilated N a significant rates. Shoots from a site characterized by elevated N availability in the winter water column and no significant sediment N reservoir exhibited GS activities that were 9-times higher than shoot from a more oligotrophic site. Shoot GS activities in July increased linearly from 5 to 33 m and wer correlated with light availability as defined by H^sat, (daily period during which photosynthetic reaction are light-saturated). This may represent metabolic compensation by P. oceanica to maintain N influx. Factors contributing to the ecological success of P. oceanica include the ability to assimilate N under conditions of severe light limitation (< 35 μmol photons. m^-2. s^-1), and metabolic plasticity to ensur the de novo generation of N-containing organic compounds.     

Antioxidant response of the seagrass Posidonia oceanica when epiphytized by the invasive macroalgae Lophocladia lallemandii

The aim was to evaluate the antioxidant defences and the occurrence of oxidative damage in Posidonia oceanica under a stress situation due to the epiphytism of the invasive macroalgae Lophocladia lallemandii. P. oceanica leaves were collected in the absence (control station) and in the presence of the epiphytic algae L. lallemandii and the antioxidant enzyme activities, markers of oxidative damage, and hydrogen peroxide production were determined. Antioxidant enzyme - catalase, glutathione peroxidase and superoxide dismutase - activities were significantly higher in Posidonia epiphytized by L. lallemandii. Malondialdehyde, protein carbonyl derivates, and glutathione levels were also higher in L. lallemandii epiphytized P. oceanica leaves compared to control samples. The production of hydrogen peroxide was also significantly increased when Posidonia was epiphytized by L. lallemandii. The invasion of P. oceanica meadows by L. lallemandii appeared to induce oxidative stress in the seagrass as evidenced by increased levels of oxidative stress markers and antioxidant defences.     

Architectural characteristics of two bed types of the seagrass Posidonia oceanica over different spatial scales

Seagrass beds occur in various morphological forms, ranging from small patches to continuous meadows. The endemic Mediterranean seagrass Posidonia oceanica forms dense and extensive stands that occur in several different morphotypes, including reticulate (seagrass interspersed with a different habitat type, such as bare sand) and continuous beds. This study, undertaken in the Maltese Islands, examined whether reticulate and continuous P. oceanica beds, located adjacent to each other and at similar depths, had different within-bed architectural characteristics. Five commonly used architectural measures (shoot density, number of leaves per shoot, mean leaf length, mean leaf width and shoot biomass) were measured from P. oceanica shoots collected from the two bed types at three different spatial scales: (1) tens of metres (‘small’ scale); (2) hundreds of metres (‘medium’ scale); and (3) kilometres (‘large’ scale). Results of 2-factor ANOVA (factor 1=bed type; factor 2=sampling locality) carried out at the three spatial scales indicated significant differences between the two bed types in shoot density (P<0.01) and leaf length (P<0.05) at the small scale, and in leaf number (P<0.05) at the large scale. Significant interactions were also apparent for shoot density (at the large scale) and for shoot biomass (at the medium scale). However, the results obtained did not indicate consistent architectural differences between the two P. oceanica bed types over the spatial scales considered. Spatial variations in within-bed architectural characteristics observed were therefore thought to be attributable mainly to the influence of local environmental factors. The findings are discussed with reference to the conservation and management of P. oceanica habitat.     

Proximate Constituents, Biomass, and Energy in Posidonia oceanica (Potamogetonaceae)

Abstract. The proximate composition of the various components of Posidonia oceanica is given in terms of gravimetric and energetic level and amount of plants from 2 depths at Port-Cros (Var. France). The level of proximate constituents differed little between the leaves (regardless of age) and the roots, but the rhizome contained much more soluble carbohydrate and less structural carbohydrate and ash. Because of this, the energy level in the leaves was more in terms of organic material and less in terms of total material than the energy level in the rhizome. The leaves of a P. oceanica shoot at 2 m depth in July contained 1.6 g organic material. 29.0 kJ. The weight and energy of the soluble carbohydrate in the rhizome from the base of the leaves to the 18th sheath scale decreased by ca. 40% from October to March and increased by ca. 100% from March to July.     

Enhalus acoroides responses to experimental shoot density reductions in Haad Chao Mai National Park,Trang Province,Thailand

The effect of self‐shading and competition for light in the seagrass Enhalus acoroides were investigated with a density reduction experiment in Haad Chao Mai National Park, Trang Province, Thailand. The study was carried out in a monospecific meadow with a natural density of 141.0 ± 8.7 shoots·m^?2. The intent was to determine the response of E. acoroides beds to loss of shoots and thinning, which often occur during typhoons and severe storm activity. Permanent quadrats were manipulated by clipping the seagrass shoots to 140, 72, 36 and 16 shoots·m^?2, to yield natural, 50%, 25% and 10% densities, respectively. Reducing shoot density in E. acoroides increased underwater light intensity below the canopy, generating increased leaf surface area and shoot weight. Seagrass leaf width, growth rate, and number of leaves per shoot also increased with greater light. The extent of flowering varied among treatments with no consistent trend. Our results demonstrate that increasing the available light to E. acoroides produces an increasing leaf size response as self‐shading in the bed is reduced.     

Ontogenic variation and effect of collection procedure on leaf biomechanical properties of Mediterranean seagrass Posidonia oceanica (L.) Delile

Leaf mechanical traits are important to understand how aquatic plants fracture and deform when subjected to abiotic (currents or waves) or biotic (herbivory attack) mechanical forces. The likely occurrence of variation during leaf ontogeny in these traits may thus have implications for hydrodynamic performance and vulnerability to herbivory damage, and may be associated with changes in morphologic and chemical traits. Seagrasses, marine flowering plants, consist of shoot bundles holding several leaves with different developmental stages, in which outer older leaves protect inner younger leaves. In this study we examined the long‐lived seagrass Posidonia oceanica to determine ontogenic variation in mechanical traits across leaf position within a shoot, representing different developmental stages. Moreover, we investigated whether or not the collection procedure (classical uprooted shoot versus non‐destructive shoot method: cutting the shoot without a portion of rhizome) and time span after collection influence mechanical measurements. Neither collection procedure nor time elapsed within 48 h of collection affected measurements of leaf biomechanical traits when seagrass shoots were kept moist in dark cool conditions. Ontogenic variation in mechanical traits in P. oceanica leaves over intermediate and adult developmental stages was observed: leaves weakened and lost stiffness with aging, while mid‐aged leaves (the longest and thickest ones) were able to withstand higher breaking forces. In addition, younger leaves had higher nitrogen content and lower fiber content than older leaves. The observed patterns may explain fine‐scale within‐shoot ecological processes of leaves at different developmental stages, such as leaf shedding and herbivory consumption in P. oceanica.     

Temporal Dynamics and Biomass Partitioning in Three Adriatic Seagrass Species: Posidonia oceanica,Cymodocea nodosa,Zostera marina

Abstract. The temporal dynamics of three seagrasses, Posidonia oceanica, Cymodocea nodosa and Zostera marina, was studied in different areas of the Adriatic Sea by analysing phenological parameters and biomass trends in different compartments of seagrass systems. For this purpose, samplings were conducted in 1997 once per season at each station, Otranto (southern Adriatic Sea) and Grado (northern Adriatic Sea). Structural parameters and biomass of plant compartments differed among seagrasses both in absolute values and in seasonal variability. P. oceanica was the largest plant, showing the highest number of leaves per shoot, highest leaf surface, Leaf Area Index and shoot weight. Z. marina was intermediate in size and had the longest leaves, whereas C. nodosa was the smallest seagrass. P. oceanica accounted for the highest total biomass (mean ± SE: 1895.9 ± 180.2 g DW · m^–2; CV = coefficient of variation: 19.0 %), considerably more than C. nodosa (mean ± SE: 410.4 ± 88.4 g DW·m^–2; CV: 43.1 %) and Z. marina (mean ± SE: 312.1 ± 75.1 g DW · m^–2; CV: 48.1 %), although the two latter species displayed a higher seasonal variability. Similarly, other features, such as shoot density, leaf surface, LAI, shoot weight and relative contributions of above‐ and below‐ground compartments, were less variable across seasons in P. oceanica than in the two other seagrasses, while leaf length showed the highest seasonal fluctuation in P. oceanica. As for biomass partitioning, C. nodosa showed a higher proportion of the below‐ground relative to above‐ground biomass (up to 90 %), with a distinct seasonality, whereas in P. oceanica the proportion of below‐ground biomass (around 80 %) was fairly constant during the year. We infer that in P. oceanica the seasonal forcing is probably buffered by the availability of internal resources stored permanently during the year in the below‐ground. In C. nodosa and Z. marina, instead, growth processes seem to be amplified by a greater influence of environmental factors.     

Growth Dynamics in Posidonia oceanica (L.) Delile

Abstract. Amounts of photosynthate in the rhizomes, in photosynthetically inactive basal parts of the leaves and different old leaf blades were examined. Winter leaf growth was supported by mobilisation of starch in the rhizome. This winter growth enables Posidonia oceanica to utilize the increased energy influx in early spring via the substantial leaf area already developed and to approach highest productivity in spring. During summer and autumn considerable concentrations of soluble carbohydrates were found in the leaves and rhizomes. Starch was stored in the rhizomes in concentrations of up to 6.8 % of dry weight. Levels of nitrogen and free amino acids were correlated with growth rates. The percentage of total nitrogen present as free amino acid-nitrogen decreased from November (35 %) to summer (less than 1 %), by which time leaf growth had stopped. In contrast to this, amounts of organic anions were low in winter and spring and reached their maximum in summer.
Within a shoot, sites of leaf growth were characterized by high amounts of total nitrogen and free amino acids (innermost leaves), while carbon Fixation was highest in the leaves #2–#4. Particularly high concentrations of soluble carbohydrates were found midway along leaf blades.
It is evident that this unusual growth rhythm of Posidonia oceanica was only possible because of the ability to store considerable amounts of carbon and nitrogen in the rhizomes.
The results of this investigation demonstrate on the one hand that the chemical composition of the plant is strongly correlated with growth and production, and on the other hand that it is dependent on environmental factors, such as energy influx and temperature, which change with season.     

Effect of different substrata on rhizome growth,leaf biometry and shoot density of Posidonia oceanica

The effects of different substratum typologies on Posidonia oceanica growth and morphology were estimated in four Sicilian meadows using Generalized and Linear Mixed Models combined with retrodating and biometric analyses. Substratum exerted a multiple effect, resulting in different biometric features for P. oceanica shoots settled on rock from those growing on sand and matte. On rock, values for growth rate, leaf length and shoot surface were lower than those on other substrata, with 42%, 23% and 32% the highest degree of difference respectively. The present study may have interesting methodological consequences for the comprehensive understanding of the causative variables potentially affecting meadows features and their health status. The importance of substratum in the prediction of likely biometry changes in P. oceanica meadows, means that knowledge of substratum type should receive due attention in the future to derive reliable estimates of meadow status.     

Variability in the structure of epiphytic assemblages of the Mediterranean seagrass Posidonia oceanica in relation to depth

The aim of the present study was to evaluate whether the variability in the structure of the epiphytic assemblages of leaves and rhizomes of the Mediterranean seagrass Posidonia oceanica differed between depths at a large spatial scale. A hierarchical sampling design was used to compare epiphytic assemblages at two different depths (10 and 20 m) in terms of both species composition and abundance and patterns of spatial variability in the Tuscan Archipelago (North Western Mediterranean Sea, Italy). Results showed significant differences in the structure of assemblages on rhizomes and leaves at different depths. These differences were related to species composition and abundance; differences were not significant for total biomass, total percentage cover and percentage cover of animals and algae. Whereas the higher variability was observed among shoots in all the studied systems, patterns of spatial variability at the other spatial scales investigated differed between the two studied depths. Moreover, in the present study, analogous patterns between depths resulted for both the assemblages of leaves and rhizomes, suggesting that factors that change with depth can be responsible for the spatial variability of both the assemblages (leaves and rhizomes), and operate regardless of the microclimatic conditions and the structure of assemblages.     

Fish Assemblages of Posidonia oceanica Beds at Port-Cros (France, NW Mediterranean): Assessment of Composition and Long-Term Fluctuations by Visual Census

Abstract. Observations of the fish fauna of Posidonia oceanica (L.) DELILE seagrass beds were undertaken in two shallow stations and one deeper station in Port-Cros National Park, between November 1984 and September 1987. Demographic structure, density and biomass of the 19 most abundant species or taxa were assessed by several non-destructive sampling methods. The fish assemblages studied were dominated in number and biomass by a small number of species (Labridae: Symphodus ocellatus and S. tinca ; Sparidae: Diplodus annularis and Sarpa salpa ). Large individuals were infrequent and small size class fish were dominant especially in sheltered shallow areas. Recruitment and migrations (shortor long-term) have been identified as the most important causes of abundance variations over the year. The four trophic categories were sampled only in the sheltered shallow zone, where the herbivore Sarpa salpa represents a strong proportion of the fish fauna (40–70%) during summer. The fish assemblages differ between the three seagrass beds, and the depth appears as the most important factor affecting the fish assemblages in P. oceanica . The functional difference between shallow and deeper seagrass beds is therefore discussed.     

Patterns of spatial variation of nutrient content,epiphyte load and shoot size of Posidonia oceanica seagrass meadows (Mediterranean Sea)

Knowledge of patterns of spatial variability of vegetative development, epiphyte load and nutrient availability in seagrass meadows is essential for the adequate design of research and environmental monitoring programmes. Differences in shoot size, epiphyte load and nutrient content of leaves and epiphytes of the Mediterranean endemic seagrass Posidonia oceanica at spatial scales ranging from metres to hundreds of metres are evaluated using a hierarchical nested sampling design. The size and epiphyte load of P. oceanica shoots and the nitrogen and phosphorus content of leaves and epiphytes were different in most of the spatial scales considered. Sampling efforts concentrated at the metre scale incorporated most of the variability in size, epiphyte load and nutrient content of the leaves and epiphytes of P. oceanica shoots. Epiphyte load showed no correlation with nutrient content in the epiphytes or in the leaves. However, epiphyte load and shoot size were negatively correlated, which suggests that light penetration in the canopy may be a main determinant of epiphyte load.     

Seasonal Changes in Nitrogen, Free Amino Acids, and C/N Ratio in Mediterranean Seagrasses

Abstract. Seasonal changes in nitrogen, free amino acids, and carbon were investigated in the three Mediterranean seagrasses Posidonia oceanica, Cymodocea nodosa , and Zostera noltii. Leaves, rhizomes, roots, as well as dead plant material were analysed separately. Highest N-concentrations were obtained in the winter months, regardless of species or plant part. In contrast to the other two species, the N-content in Posidonia was higher in the rhizomes than in the leaves.
In the investigated species, marked differences in the free amino acid (FAA) composition were detected between species: Proline, lacking in Posidonia , was the main component in Cymodocea. In Posidonia , FAA decreased from 320umol g^-1 (dry wt) in leaf sheaths to 1.5 umol g^-1 (dry wt) in the leaf tips. The function of proline as a possible storage and/or stress metabolite is discussed.
High C/N values in dead P. oceanica and C. nodosa rhizomes as well as in P. oceanica wrack seem to be related to considerable resistance to decomposition. Low C/N ratios did not increase in detached C. nodosa and Z. noltii leaves, which began to decompose shortly after detachment from the plants.     

Feeding of the Sparid Fish Sarpa salpa in a Seagrass Ecosystem: Diet and Carbon Flux

Abstract. In Revellata Bay Gulf of Calvi, Corsica, France, the sparid fish Sarpa salpa L. is the main macro-consumer of Posidonia oceanica L. D elile leaf. Stomach contents were analysed and ¹³C/¹²C isotopic ratios were measured in fish muscle and potential food sources algae, P. oceanica leaf and its epiphytes to determine their relative contribution to the fish diet. S. salpa has an age-related mixed diet: juveniles are plankton feeders, young, sub-adults and adults are herbivorous, and, the older the individuals, the higher the relative contribution of P. oceanica to the diet. Our results and former studies of carbon stocks and fluxes in the P. oceanica bed of Revellata Bay have enabled an estimation of the general impact of S. salpa grazing on infralittoral communities: the studied species consumes 24 g C.m^-2 -a^-1 from P. oceanica leaf, 4.8 from epiphytes and 13 from epilithic algae. The fish net production and biomass turnover in that zone have been estimated to be 1.2gC.m^-2.a^-1 and 1.5 a^-1, respectively.     

A comparative study of the photosynthetic activity among three temperate seagrass species in Northern Japan

The photosynthetic activity of Zostera marina, Zostera asiatica and Phyllospadix iwatensis shoots from populations of Hokkaido (Northern Japan) was determined using the pulse amplitude modulated (PAM) fluorometer. Several fluorescence parameters were measured as a function of irradiance and leaf age: electron transport rate (ETR), quantum yield, photochemical quenching (qP) and non‐photochemical quenching (NPQ). The leaf age determined by the leaf position in the shoot bundle strongly influenced the photosynthetic activity of Z. marina, Z. asiatica and P. iwatensis. Young leaves had the maximum electron transport rate (Zm: Leaf 1 = 15.7, Leaf 2 = 16.3; Za: Leaf 1 = 13.0, Leaf 2 = 12.2; and Pi: Leaf 1 = 12.5, Leaf 2 = 11.7) and showed higher photoprotection (NPQ) than old leaves. Among the studied seagrass species, Z. marina had the highest photosynthetic activity (ETR_max = 15.3), in accordance with the highest production in the field in comparison with the other two seagrass species. The PAM fluorometry technique showed to be effective in determining intraspecific (among‐leaves) and interspecific (among‐species) variation in seagrass photosynthetic activity.     

Structure of the Epiphytic Community of Posidonia oceanica Leaves in a Shallow Meadow

Abstract. The structure of the epiphytic community (hydroids, bryozoans, and the most abundant macrophytes) of Posidonia oceanica leaves in a shallow meadow near Lacco Ameno, Ischia was studied. The structure of the community changed along an age gradient that extended from the inner to the outer leaves and from the basis to the apex of each leaf. This structure was also affected by microclimatic factors. The dynamics of community "maturation" is described.     

Ecological Differences in the Distribution of two Tethya (Porifera, Demospongiae) Species Coexisting in a Mediterranean Coastal Lagoon

Abstract. In the Sicilian lagoon Stagnone di Marsala, Tethya aurantium and T. citrina show different ecological distributions, although some overlapping occurs. The first species reaches its highest density where linear currents are maximal, and in this habitat it is particularly frequent on Posidonia oceanica rhizomes. The second species is more abundant in still water locations and is equally common on Posidonia rhizomes and on calcareous concretions. Such distributions can be related to water movement, Posidonia shoot density, and associated factors such as sedimentation and food availability. These observations are consistent with an adaptive interpretation of the morphofunctional features distinguishing the two species.     

Long-term change in the structure of a Posidonia oceanica landscape and its reference for a monitoring plan

Abstract On the basis of a detailed cartographic survey carried out by Side Scan Sonar and a towed underwater video camera during 2005, and from a series of historical maps (1959, 1980, 1990), an extensive regression of Posidonia oceanica (L.) Delile beds was evidenced for a vast area of the central Tyrrhenian Sea (Latium coast, Italy). The total loss of P. oceanica surface was assessed through GIS estimate. In 1959, the Posidonia beds extended over 7290 ha, while in the 2005 survey they had regressed to 2899 ha, a loss of about 60% of their coverage. Also the seagrass lower limit showed a general depth decrease in time. Total seagrass coverage loss and lower limit regression were not uniform along the whole investigated areas and three main sub-areas have been identified with different degrees of regression somehow related with coastal potential human-mediated impacts. From different coverage estimates of the present survey and of the previous maps, minimum sampling areas were calculated through bootstrapping simulation routines from small sampling areas (Landscape Units) to reach the nearest estimate of the observed condition in the different periods.     

Temporal variations in the spatial distribution of shoot density in a Posidonia oceanica meadow and patterns of genetic diversity

The meadows of the Mediterranean endemic seagrass Posidonia oceanica exhibit relatively high variations of structural and biometric features at various spatial scales. An investigation performed in 1992 in the meadow off Lacco Ameno (Island of Ischia, Gulf of Naples, Italy) detected peculiar spatial patterns of plant distribution, characterized by nestlike structures with radially increasing or decreasing shoot densities. Eight years later (2000), geo‐referenced collections at selected points were repeated to trace the temporal variations of shoot density and investigate the recurrence of the density cores previously detected. In addition, shoots for molecular analyses were collected to check the hypothesis that nestlike patterns exhibit highest levels of genetic variability, due to the confluence of several genetically distinct stolons. The 2000 survey confirmed the presence of the main density cores detected in 1992, although their spatial distribution was slightly shifted and a general decrease of spatial anisotropy was observed, probably due to an increased disturbance, mainly due to pleasure boat anchoring. Patterns of genetic diversity showed a more complex picture, well related to the shoot density spatial pattern, especially when compared with the previous 1992 survey. Patterns of genetic diversity confirmed our previous hypotheses on the genesis of shoot density cores, suggesting they are produced over long time, due to a slow stolonization process and a convergence of different genotypes. Regression of the meadow and decrease of density may lead, in short periods, to a homogenization of the density patterns, while genetic diversity cores represent a long‐term ‘memory’ of their previous distribution.     

Seasonal Changes in Soluble Carbohydrates, Starch, and Energy Content in Mediterranean Seagrasses

Abstract. In all scagrass species investigated, rhizomes were the main storage organs for soluble carbohydrate reserves, and the highest levels were found in Cymodocea nodosa (U cria ) A scherson . Sucrose was the primary soluble carbohydrate identified in Posidonia oceanica (L.) D elile , and Zostera noltii H ornem . This disaccharide was dominant in Cymodocea nodosa roots, as well as in its rhizome in winter. Myo-inositol was the main cyclitol in Zostera noltii. In Cymodocea nodosa leaves 1-chiro-inositol was the main sugar component; it was also present to a lesser extent in the roots and rhizomes. The amounts of glucose and fructose were correlated with growth to a certain extent in all species. Higher amounts of starch were stored in the rhizomes of Posidonia oceanica and Zostera noltii. The energy content of Mediterranean scagrasses was significantly higher than in tropical species. No distinct seasonal trend in energy content could be observed.