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- Population-level effects of clam harvesting on the seagrass Zostera noltiiPublication . Cabaço, Susana; Alexandre, Ana; Santos, RuiSeagrass declines have been reported worldwide, mostly as a consequence of anthropogenic disturbance. In Ria Formosa lagoon, southern Portugal, the intertidal meadows of Zostera noltii are highly disturbed by clam harvesters. The most common technique used to collect the clams consists of digging and tilling the sediment with a modified knife with a large blade. Here we present both descriptive and experimental evidence of the negative effects of clam harvest on the Z. noltii populations of Ria Formosa. A comparison between disturbed and undisturbed meadows suggests that clam harvesting activities change the species population structure by significantly reducing shoot density and total biomass, particularly during August, when the harvest effort is higher. Experimental harvest revealed a short-term impact on shoot density, which rapidly recovered to control levels during the following month. An experimental manipulation of rhizome fragmentation revealed that plant survival is reduced only when fragmented rhizomes are left with 1 intact internode. Shoot production and rhizome elongation and production of fragmented rhizomes having 2 to 5 internodes were not significantly affected, even though growth and production were lower when only 2 internodes were left. Experimental shoot damage at different positions along the rhizome had a significant effect on plant survival, rhizome elongation, and production only when the apical shoot was removed. Our results show that clam harvest can adversely affect Z. noltii meadows of Ria Formosa while revealing a low modular integration that allows the species to rapidly recover from physical damage.
- Nutrition of the seagrass Cymodocea nodosa: pulses of ammonium but not of phosphate are crucial to sustain the species growthPublication . Alexandre, Ana; Santos, RuiWe investigated the nutrient acquisition of the seagrass Cymodocea nodosa and the importance of sediment to water column nutrient pulses, which regularly occur in coastal lagoons forced by incoming flood tides. Ammonium was preferentially taken up and accounted for 60-90% of the total daily N acquisition, whereas amino acid acquisition through belowground plant parts represented the second most important source of N, accounting 8-34%. The uptake of ammonium pulses increased dramatically the daily N acquisition from 9.5 mmol m-2 day-1 to 1.33 mmol m-2 day-1 at ambient nutrient concentrations, enabling the species to meet its N demand. Roots accounted for 96% of the total phosphate acquisition, with no evidence for P limitation. The ability of seagrasses to adapt to nutrient pulses may be an overlooked nutrition strategy common to many ecosystems where nutrient availability in the water column is low but nutrient pulses occur.
- Light is more important than nutrient ratios of fertilization for cymodocea nodosa seedling developmentPublication . Alexandre, Ana; Silva, João; Santos, RuiRestoration of seagrass beds through seedlings is an alternative to the transplantation of adult plants that reduces the impact over donor areas and increases the genetic variability of restored meadows. To improve the use of Cymodocea nodosa seedlings, obtained from seeds germinated in vitro, in restoration programs, we investigated the ammonium and phosphate uptake rates of seedlings, and the synergistic effects of light levels (20 and 200 mu mol quanta m(-2) s(-1)) and different nitrogen to phosphorus molar ratios (40 mu M N:10 mu M P, 25 mu M N:25 mu M P, and 10 mu N:40 mu M P) on the photosynthetic activity and growth of seedlings. The nutrient content of seedlings was also compared to the seed nutrient reserves to assess the relative importance of external nutrient uptake for seedling development. Eighty two percent of the seeds germinated after 48 days at a mean rate of 1.5 seeds per day. All seedlings under all treatments survived and grew during the 4 weeks of the experiment. Seedlings of C. nodosa acquired ammonium and phosphate from the incubation media while still attached to the seed, at rates of about twice of adult plants. The relevance of external nutrient uptake was further highlighted by the observation that seedlings' tissues were richer in nitrogen and phosphorus than non-germinated seeds. The uptake of ammonium followed saturation kinetics with a half saturation constant of 32 mu M whereas the uptake of phosphate increased linearly with nutrient concentration within the range tested (5 - 100 mu M). Light was more important than the nutrient ratio of fertilization for the successful development of the young seedlings. The seedlings' photosynthetic and growth rates were about 20% higher in the high light treatment, whereas different nitrogen to phosphorus ratios did not significantly affect growth. The photosynthetic responses of the seedlings to changes in the light level and their capacity to use external nutrient sources showed that seedlings of C. nodosa have the ability to rapidly acclimate to the surrounding light and nutrient environment while still attached to the seeds. C. nodosa seedlings experiencing fertilization under low light levels showed slightly enhanced growth if nourished with a balanced formulation, whereas a slight increase in growth was also observed with unbalanced formulations under a higher light level. Our results highlight the importance of high light availability at the seedling restoration sites.
- Nitrogen uptake in light versus darkness of the seagrass Zostera noltei: integration with carbon metabolismPublication . Alexandre, Ana; Silva, João; Santos, RuiWe conducted a study that shows that light and dark conditions do not affect the uptake rates of ammonium and nitrate by the seagrass Zostera noltei. This is an important advantage over some seaweed species in which these rates are severely reduced at night. In the light, the ammonium uptake rates were initially higher (15 and 20molg(-1)h(-1)) and stabilized at a rate of 5molg(-1)h(-1) after 1h, whereas in the dark the rates remained constant at a rate of 10molg(-1)h(-1) over the first 180min of incubation. The rates of nitrate uptake in the light were high within the first 120min of incubation(7.2-11.1molg(-1)h(-1)) and decreased afterwards to lower values (0.8-3.9molg(-1)h(-1)), whereas in the dark the rates fluctuated around 0.0-11.1molg(-1)h(-1) throughout the whole incubation time (7h). The soluble sugar content of Z.noltei leaves increased significantly with both ammonium and nitrate incubations in the light, indicating the metabolic outcome of photosynthesis. In the dark, there was no significant variation in either the soluble sugar or in the starch content of leaves, rhizomes or roots in either the ammonium or nitrate incubations. However, the total starch content of plants decreased at night whereas the total soluble sugars increased, suggesting a process of starch catabolism to generate energy with the consequent production of smaller monosaccharide products. The starch content of rhizomes decreased significantly during the light incubations with nitrate but not with ammonium. These results suggest that carbohydrate mobilization is necessary for Z.noltei to account for extra energetic costs needed for the uptake and assimilation of nitrate. Furthermore, our results suggest that nitrate uptake, at least during the day, requires the mobilization of starch whereas the uptake of ammonium does not.
- Effects of elevated CO2 and nutrients on the community metabolism of a Cymodocea nodosa bedPublication . Khan, Faisal; Alexandre, Ana; Ullah, Hadayet; Santos, RuiWe assessed the combined effects of elevated CO2 and nutrients on the metabolism of a benthic community dominated by the seagrass Cymodocea nodosa (Ucria) Ascherson in a mesocosm experiment. C. nodosa plants and their associated community were exposed to two CO2 levels simulating future (700 ppm, pH 7.84) and current (360 ppm, pH 8.12) conditions, and two nutrient levels (enriched and ambient concentration) in a total of four treatments (-C-N, -C+N, +C-N, +C+N). Net community production (NCP) was estimated from changes in the concentration of dissolved inorganic carbon in the seawater in light incubations using benthic chambers. The variation pattern of NCP with the ordinance was consistent for all treatments. Although differences among treatments were not statistically significant, average NCP values were lowest under CO2 enrichment conditions. NCP was lower at a high CO2 level and ambient nitrogen concentration compared to when nutrient availability was higher, suggesting that the low nutrient availability may modulate the community response to CO2 enrichment. The results obtained suggest that the stimulation of the net community production of C. nodosa by elevated CO2 concentrations may be curtailed by low nutrient availability.
- The effects of epiphytes on light harvesting and antioxidant responses in the seagrass posidonia oceanicaPublication . Costa, Monya; Silva, João; Olivé, Irene; Barrote, Isabel; Alexandre, Ana; Albano, Sílvia; Santos, RuiPosidonia oceanica (L.) Delile is a subtidal seagrass whose leaves are commonly colonized by epiphytes. Epiphytes pose physical barriers to light penetration within the leaves, with possible significant impacts on photosynthesis. Furthermore, epiphytes can indirectly be responsible for leaf chlorosis, necrosis and senescence which are known to be related with the increase of oxygen reactive species (ROS) levels, potentially leading to oxidative stress. The aim of this work was to investigate in situ (i) the effect of epiphytes on the composition and balance of light harvesting pigments in leaves of the naturally growing seagrass P. oceanica, and (ii) evaluate differences in antioxidant responses. Epiphytized and non-epiphytized plants were analyzed to establish potential photosynthetic pigment roleshift between light harvesting and photoprotection functions. The experiments were carried out in Cabo de Gata Natural Park, southern Spain, where epiphytized and non-epiphytized plants can be found at identical depths and light exposure. The results showed that both O2 evolution rate along the day and chlorophyll a/b ratio were higher in non-epiphytized plants, indicating a negative effect of epiphytes on photosynthesis and light harvesting. Although under high irradiance (at solar noon) the xanthophyll cycle was activated in both epiphytized and non-epiphytized leaves, the de-epoxidation-ratio (AZ/VAZ) was lower in epiphytized leaves, due to light attenuation by epiphytes. The antioxidant capacity (TEAC and ORAC essays) and the activity of the antioxidant enzymes ascorbate peroxidase and dehydroascorbate were higher in epiphytized plants, showing that epiphytes can also be a potential source of oxidative stress to P. oceanica. Our results show that despite the light attenuation effect, leaf colonization by epiphytes can also be potentially stressful and reduces plant productivity.
- Nitrogen uptake kinetics of an enteric methane inhibitor, the red seaweed Asparagopsis armataPublication . Torres, Raquel; Mata, Leonardo; Santos, Rui; Alexandre, AnaThe use of red seaweeds of the genus Asparagopsis as a feed ingredient drastically decreases the enteric methane production by ruminant livestock, thereby reducing the environmental impact of this industry. To address the world demand for Asparagopsis mass production, it is essential to understand the species nutrition. In this study, we evaluated for the first time the uptake kinetics of inorganic and organic nitrogen forms of the tetrasporophyte of Asparagopsis armata, to identify the different uptake phases (surge and internally controlled) and to reveal the species preferential nitrogen sources. The time course of nitrogen uptake rates was followed, and the preferential nitrogen sources were assessed in choice-uptake experiments through the incorporation of (15) N-labelled ammonium, nitrate and amino acids. As expected, the short-term, surge uptake rates were much higher than the stabilized internally controlled uptake rates. Ammonium was the preferred form during the internally controlled phase but surprisingly, the surge uptake rates of amino acids were much higher than those of inorganic forms. The importance of amino acids for A. armata nutrition was further supported by its internally controlled uptake rates that where higher than those of nitrate. Ammonium is, thus, the main nitrogen form for the species nutrition, but pulses of organic nitrogen may be considered in the mass production of Asparagopsis for the enteric methane inhibition of the livestock industry.
- The takeover of Thalassia testudinum by Anadyomene sp. at Biscayne Bay, USA, cannot be simply explained by competition for nitrogen and phosphorousPublication . Alexandre, Ana; Collado-Vides, Ligia; Santos, RuiApart from direct light effects, we tested whether the takeover of the seagrass Thalassia testudinum by the seaweed Anadyomene sp. in high nutrient areas of Biscayne Bay, Florida, USA, is related to a faster nutrient surge uptake capacity of the seaweed and/or a negative effect on the seagrass uptake rates. Anadyomene sp. and T. testudinum showed a similar ammonium surge uptake capacity, but the seagrass performed better than the seaweed in mixed incubations at high ammonium concentrations. T. testudinum was faster than Anadyomene sp. at taking up pulses of phosphate, but the uptake rates of the seagrass were significantly decreased in the presence of the seaweed. The takeover of T. testudinum by Anadyomene sp. at Biscayne Bay is likely dominated by light and cannot be simply explained by their single or mixed nutrient surge uptake rates, but the phosphate availability and the seagrass uptake inhibition by the seaweed may also play a key role in the process.
- High nitrogen and phosphorous acquisition by belowground parts of Caulerpa prolifera (Chlorophyta) contribute to the species' rapid spread in Ria Formosa lagoon, Southern PortugalPublication . Alexandre, Ana; Santos, RuiDespite worldwide proliferation of the genus Caulerpa and subsequent effects on benthic communities, little is known about the nutritional physiology of the Caulerpales. Here, we investigated the uptake rates of ammonium, nitrate, amino acids, and phosphate through the fronds and rhizoids + stolon, the internal translocation of nitrogen, and developed a nitrogen budget for the rapidly spreading Caulerpa prolifera in Ria Formosa lagoon, southern Portugal. Caulerpa prolifera acquired nutrients by both aboveground and belowground parts at similar rates, except nitrate, for which fronds showed 2-fold higher uptake rates. Ammonium was the preferential nitrogen source (81% of the total nitrogen acquisition), and amino acids, which accounted for a significant fraction of total N acquisition (19%), were taken up at faster rates than nitrate. Basipetal translocation of 15 N incorporated as ammonium was nearly 3-fold higher than acropetal translocation, whereas 15 N translocation as nitrate and amino acids was smaller but equal in either direction. The estimated total nitrogen acquisition by C. prolifera was 689 μmol · m-2 · h-1 , whereas the total nitrogen requirement for growth was 672 μmol · m-2 · h-1 . The uptake of ammonium and amino acids by belowground parts accounted for the larger fraction of the total nitrogen acquisition of C. prolifera and is sufficient to satisfy the species nitrogen requirements for growth. This may be one reason explaining the fast spreading of the seaweed in the bare sediments of Ria Formosa where it does not have any macrophyte competitors and the concentration of nutrients is high.
- First description of seagrass distribution and abundance in Sao Tome and PrincipePublication . Alexandre, Ana; Silva, João; Ferreira, Rogério; Paulo, Diogo; Serrao, Ester; Santos, RuiSeagrass meadows in Sao Tome and Principe, eastern Atlantic Ocean, are described here for the first time. Specifically, we quantified the biomass and density of seagrasses, characterized the plant morphology and measure their nutrient content as a proxy of the nutrient environmental conditions where the meadows develop. The seagrass Halodule wrightii was found in two locations of the northeastern coast of the island of Sao Tome: 1) developing throughout an estimated area of 1500 ha surrounding Cabras islet, at a depth range of 4-10 m, on sandy bottom; and 2) at Santana bay with an area of 1500 m(2) at 5-10 m depth, on sandy bottom. A highly morphologically different population of Halodule wrightii was found on the northeastern coast of the island of Principe, off Abade beach, covering an area of 135 m2 at 4 m depth. Further research is needed to assess if this is a different species. Shoot biomass and density was 10 and 4-fold higher in Sao Tome than in Principe, respectively. CN ratios of above and belowground tissues of plants collected in Sao Tome were also significantly higher than in Principe. The carbon content of Halodule leaves from Sao Tome and Principe (41%) was much higher than that reported for other Halodule species, suggesting that meadows may have an important ecological role for carbon fixation. The presence of H. wrightii in Sao Tome and Principe raises ecological and evolutionary questions that warrant further research.