Fernando G. Brun

Effects of shoot stiffness, shoot size and current velocity on scouring sediment from around seedlings and propagules

Successful management and restoration of coastal vegetation requires a quantitative process-based understanding of thresholds hampering (re-)establishment of pioneer vegetation. We expected scouring to be important in explaining the disappearance of seedlings and/or small propag- ules of intertidal plant species, and therefore quantified the dependence of scouring on plant traits (flexibility, size) and physical forcing by current velocity. Flume studies with unidirectional flow revealed that scouring around seedlings increased exponentially with current velocity and according to a power relationship with plant size. Basal stem diameter rather than shoot length controlled scour- ing volume. Flexible shoots caused far less scouring than stiff shoots, provided that the bending occurred near the sediment surface as was the case for Zostera, and not on top of a solid tussock base as we observed for Puccinellia. Therefore, shoot stiffness is likely to strongly affect the chances for initial establishment in hydrodynamically exposed areas. Plant traits such as shoot stiffness are sub- ject to a trade-off between advantages and disadvantages, the outcome of which depends on the physical settings.

Morphological and physiological differences between two morphotypes of Zostera noltii Hornem. from the south-western Iberian Peninsula

Abstract The morphological and physiological differences between two morphotypes of Z. noltii Hornem. were studied in the intertidal meadows on the south-western Iberian Peninsula (Palmones river estuary and Ria Formosa). A small-leaved morphotype (SM) grows mainly at high intertidal sites, meadow edges or in recently deposited sandbanks, whereas a large-leaved morphotype (LM) generally thrives in well-structured beds or in deeper places. This study deals with the morphological, biochemical and physiological differences between these morphotypes as well as the ecological implications of the occurrence of different morphotypes in the same meadow. Shoot length, leaf width, rhizome internode length, roots per node, root length, leaf nutrient and pigment contents, and photosynthetic rates of both morphotypes were compared. The below-ground architecture (root and rhizome complex) of both morphotypes was more developed in sites characterized by higher hydrodynamics and/or a lower nitrogen content in sediments. Both morphotypes showed similar values for photosynthetic efficiency, dark respiration rate and compensation irradiance. On the other hand, the net photosynthetic capacity was much greater (5-fold) for the SM. This difference could explain the greater growth rate and faster leaf turnover rate of the SM compared with the LM. The occurrence of the SM in newly settled areas (and in the meadow edges) could be explained on the basis of its higher growth rate, which would allow a faster spreading of the meadow and/or better recovery after burial resulting from stormy weathers.

Acclimation Responses of Gracilaria sp. (Rhodophyta) and Enteromorpha intestinalis (Chlorophyta) to Changes in the External Inorganic Carbon Concentration

Abstract The acclimation responses of two intertidal macroalgae, Gracilaria sp. and Enteromorpha intestinalis, to different dissolved inorganic carbon (DIC) levels were investigated under laboratory conditions. The effect of DIC availability on growth rate, biochemical composition (C, N, pigments and Rubisco) and on the degree of inhibition of external and total carbonic anhydrase activities (by acetazolamide and 6-ethoxyzolamide, respectively), and of a putative HCO3 − exchanger protein (sensitive to the inhibitor 4,4′-diisothiocyanatostilbene-2,2′-disulfonate, DIDS) was species-specific. Pigment and Rubisco contents co-varied negatively with DIC availability in Gracilaria sp. However, no such pattern was observed in Enteromorpha intestinalis. The mechanisms of DIC uptake were also modulated by the external DIC concentration. Under limiting DIC conditions, the induction of mechanisms for CO2 acquisition above the diffusive rate was observed in Gracilaria sp., while a repression of the DIDS-sensitive mechanism was obtained for Enteromorpha intestinalis. The results revealed the plasticity of these intertidal macroalgae to acclimate to different ambient DIC levels, and indicate the important role of DIC as a factor controlling biochemical and physiological processes.

Patch distribution and within-patch dynamics of the seagrass Zostera noltii hornem. in los Toruños Salt-Marsh, Cádiz Bay, Natural Park, Spain

Abstract Patch distribution and small-scale (i.e., within-patch) temporal variability in plant morphology, biomass, shoot density, tissue nutrient content and growth were studied in the intertidal seagrass Zostera noltii at Los Toruāos salt-marsh (Cádiz, Spain). The area covered by Z. noltii was 7.3%, and decreased exponentially with depth. The lack of recruitment from seedlings largely explained the normal patch size distribution observed. Overall, lower biomass, shoot density, above-ground to below-ground biomass ratios and higher leaf elongation rates were recorded at the patch edge, where biomass dynamics were primarily controlled by recruitment and mortality processes. In contrast, temporal variation in above-ground biomass in the patch centre depended largely on changes in shoot leaf number and size. There was no correlation between total biomass and shoot density in the patch centre suggesting that below-ground biomass may be space-limited. Low aboveground biomass and shoot elongation rates were recorded in May, regardless of position within the patch, and were coincident with the abundance of Ulva sp. mats. Light reduction by Ulva canopies also resulted in a mobilization of carbohydrate reserves. Such resource reallocation may represent an important short-term survival strategy, maintaining leaf and rhizome elongation capacity.

Interactions of light and organic matter under contrasting resource simulated environments: the importance of clonal traits in the seagrass Zostera noltii

Light reduction in the water column and enhanced organic matter (OM) load into the sediments are two main consequences of eutrophication in marine coastal areas. This study addresses the combined effects of light, OM, and clonal traits in the seagrass Zostera noltii. Large Z. noltii plants were grown in sand with or without the addition of OM and under two light levels (high light and low light). Whereas some complete plant replicates were grown under homogeneous light and/or OM conditions, other replicates were grown under contrasting light and/or OM levels between the apical and the distal parts of the same plant. The three-way factorial design (light, OM load, and apex position) allowed us to determine the harmful effect of light reduction and OM enrichment on the growth, photosynthetic performance, and biochemical composition of Z. noltii. The addition of OM to the sediment promoted a decrease, or even an inhibition, in net plant growth regardless of the light level when the whole plants were grown under homogeneous light conditions. However, the results differed when plants were grown under contrasting light and/or OM conditions between apical and distal parts. In this case, the harmful effect of OM load was alleviated when apical parts were grown under high light conditions. OM loads also negatively affected the photosynthetic performance, evaluated as leaf fluorescence. The results indicate the importance of clonal traits in the response of Z. noltii growth to light conditions and OM enrichment.

Interactions between Seagrass Complexity, Hydrodynamic Flow and Biomixing Alter Food Availability for Associated Filter-Feeding Organisms

Seagrass shoots interact with hydrodynamic forces and thereby a positively or negatively influence the survival of associated species. The modification of these forces indirectly alters the physical transport and flux of edible particles within seagrass meadows, which will influence the growth and survivorship of associated filter-feeding organisms. The present work contributes to gaining insight into the mechanisms controlling the availability of resources for filter feeders inhabiting seagrass canopies, both from physical (influenced by seagrass density and patchiness) and biological (regulated by filter feeder density) perspectives. A factorial experiment was conducted in a large racetrack flume, which combined changes in hydrodynamic conditions, chlorophyll a concentration in the water and food intake rate (FIR) in a model active filter-feeding organism (the cockle). Results showed that seagrass density and patchiness modified both hydrodynamic forces and availability of resources for filter feeders. Chlorophyll a water content decreased to 50% of the initial value when densities of both seagrass shoots and cockles were high. Also, filter feeder density controlled resource availability within seagrass patches, depending on its spatial position within the racetrack flume. Under high density of filter-feeding organisms, chlorophyll a levels were lower between patches. This suggests that the pumping activity of cockles (i.e. biomixing) is an emergent key factor affecting both resource availability and FIR for filter feeders in dense canopies. Applying our results to natural conditions, we suggest the existence of a direct correlation between habitat complexity (i.e. shoot density and degree of patchiness) and filter feeders density. Fragmented and low-density patches seem to offer both greater protection from hydrodynamic forces and higher resource availability. In denser patches, however, resources are allocated mostly within the canopy, which would benefit filter feeders if they occurred at low densities, but would be limiting when filter feeder were at high densities.

Shoot organization in the seagrass zostera noltii: implications for space occupation and plant architecture

The growth pattern of the seagrass Zostera noltii is described through the analysis of the shoot primordium organization within different shoot types using optical and scanning electron microscopy. Both histological approaches showed that Z. noltii shoots are organized by a successive repetition of a unit named “phytomer” (shoot primordium, node, internode, root, sheath and leaf), in resemblance with the shoot structure described for land grasses. This study showed that differences among shoot types are determined by two factors: (1) the presence or absence of some of the fundamental parts (mainly shoot primordium) in the “phytomer”, (2) the evolvement stage of these elements. The branching of Z. noltii was limited by shoot structure and shoot primordium arrangement; in the “natural” branching pattern the first axillary shoot branched opposite to the previous branch. Simulation of the topology of a Z. noltii plant using the “natural” branching pattern, and its opposite one, with two different branching angles for each pattern, showed that the reduction in the branching angle notably decreases the colonizing efficiency (ca. 25% from 90 to 45°). Changes in the timing of shoot primordium development and/or release, and the optimization of the branching angle in response to external forcing (light, nutrients, density, etc.) may elucidate species-specific differences and colonization strategies with respect to abiotic conditions.

The role of hydrodynamics in structuring in situ ammonium uptake within a submerged macrophyte community

In low-nutrient, macrophyte-dominated coastal zones, benthic ammonium (NH4+) uptake may be influenced by the structural properties of plant canopies via their effect on near-bed hydrodynamics. Using a dual-tracer (uranine and 15NH4+) method that does not require enclosures, we examined how this process affects nutrient uptake rates within a tidally dominated, patchy Caulerpa prolifera–Cymodocea nodosa landscape. NH4+ uptake was determined by calculating tissue 15N excesses and correcting for 15N enrichment as derived from uranine concentration. Vertical hydrodynamic profiles were measured in the downstream flow direction from outside to inside of the C. nodosa bed by using an array of acoustic Doppler velocimeters. The transition from a C. prolifera to a C. nodosa bed included a change in both benthic canopy properties (short and dense to tall and sparse) and sediment topography (0.2-m increase in water column depth) that resulted in an increase in longitudinal advection and turbulent diffusivity within the C. nodosa canopy between 0.5 and 1.5 m from the leading edge. Vertical differences in canopy water exchange appeared to explain variations in uptake between biotic functional groups; however, no clear differences in longitudinal uptake were found. Using in situ labeling, this study demonstrated for the first time the role of hydrodynamics in structuring NH4+ uptake within an undisturbed, patchy macrophyte landscape.

Interaction between Ammonium Toxicity and Green Tide Development Over Seagrass Meadows: A Laboratory Study

Eutrophication affects seagrasses negatively by increasing light attenuation through stimulation of biomass of fast-growing, bloom-forming algae and because high concentrations of ammonium in the water can be toxic to higher plants. We hypothesized nevertheless, that moderate amounts of nitrophilic macroalgae that coexists with seagrasses under eutrophic conditions, can alleviate the harmful effects of eutrophication on seagrasses by reducing ammonium concentrations in the seawater to non-toxic levels because such algae have a very large capacity to take up inorganic nutrients. We studied therefore how combinations of different ammonium concentrations (0, 25 and 50 μM) and different standing stocks of macroalgae (i.e. 0, 1 and 6 layers of Ulva sp.) affected survival, growth and net production of the seagrass Zostera noltei. In the absence of Ulva sp., increasing ammonium concentrations had a negative influence on the performance of Z. noltei. The presence of Ulva sp. without ammonium supply had a similar, but slightly smaller, negative effect on seagrass fitness due to light attenuation. When ammonium enrichment was combined with presence of Ulva sp., Ulva sp. ameliorated some of negative effects caused by high ammonium availability although Ulva sp. lowered the availability of light. Benthic microalgae, which increased in biomass during the experiment, seemed to play a similar role as Ulva sp.–they contributed to remove ammonium from the water, and thus, aided to keep the ammonium concentrations experienced by Z. noltei at relatively non-toxic levels. Our findings show that moderate amounts of drift macroalgae, eventually combined with increasing stocks of benthic microalgae, may aid seagrasses to alleviate toxic effects of ammonium under eutrophic conditions, which highlights the importance of high functional diversity for ecosystem resistance to anthropogenic disturbance.

Latitudinal Patterns in European Seagrass Carbon Reserves: Influence of Seasonal Fluctuations versus Short-Term Stress and Disturbance Events

Seagrass meadows form highly productive and valuable ecosystems in the marine environment. Throughout the year, seagrass meadows are exposed to abiotic and biotic variations linked to (i) seasonal fluctuations, (ii) short-term stress events such as, e.g., local nutrient enrichment, and (iii) small-scale disturbances such as, e.g., biomass removal by grazing. We hypothesized that short-term stress events and small-scale disturbances may affect seagrass chance for survival in temperate latitudes. To test this hypothesis we focused on seagrass carbon reserves in the form of starch stored seasonally in rhizomes, as these have been defined as a good indicator for winter survival. Twelve Zostera noltei meadows were monitored along a latitudinal gradient in Western Europe to firstly assess the seasonal change of their rhizomal starch content. Secondly, we tested the effects of nutrient enrichment and/or biomass removal on the corresponding starch content by using a short-term manipulative field experiment at a single latitude in the Netherlands. At the end of the growing season, we observed a weak but significant linear increase of starch content along the latitudinal gradient from south to north. This agrees with the contention that such reserves are essential for regrowth after winter, which is more severe in the north. In addition, we also observed a weak but significant positive relationship between starch content at the beginning of the growing season and past winter temperatures. This implies a lower regrowth potential after severe winters, due to diminished starch content at the beginning of the growing season. Short-term stress and disturbances may intensify these patterns, because our manipulative experiments show that when nutrient enrichment and biomass loss co-occurred at the end of the growing season, Z. noltei starch content declined. In temperate zones, the capacity of seagrasses to accumulate carbon reserves is expected to determine carbon-based regrowth after winter. Therefore, processes affecting those reserves might affect seagrass resilience. With increasing human pressure on coastal systems, short- and small-scale stress events are expected to become more frequent, threatening the resilience of seagrass ecosystems, particularly at higher latitudes, where populations tend to have an annual cycle highly dependent on their storage capacity.