Browsing by Author "Alexandre, Ana"
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- Competition for nitrogen between the seaweed Caulerpa prolifera and the seagrass Cymodocea nodosaPublication . Alexandre, Ana; Santos, RuiThe rhizophytic seaweed Caulerpa prolifera has been expanding rapidly in the Ria Formosa lagoon, southern Portugal, taking over deeper unvegetated areas and mixing with the native seagrass Cymodocea nodosa in shallower areas. In the Ria Formosa lagoon, belowground ammonium uptake from the sediment represents the main source of nitrogen for the 2 macrophytes, except during the ammonium pulses from the sediment to the water column that are incorporated through aboveground plant parts. We examined the competition for inorganic and organic nitrogen between C. prolifera and C. nodosa through a series of N-15-ammonium and N-15-amino acid surge uptake experiments combining single-species and mixed incubations at a range of nutrient concentrations. Our results showed that C. prolifera is generally faster than C. nodosa in the acquisition of ammonium and amino acids by both above- and belowground parts, and that the uptake rates of ammonium and amino acids of one species were not affected by the presence of the other species. The exception was the amino acid uptake through the rhizoids of C. prolifera, which was slightly enhanced in the presence of C. nodosa. In this situation, the aboveground ammonium uptake becomes the main contributor to the nitrogen budget of C. nodosa but not to that of C. prolifera. When ammonium pulses are considered, C. nodosa is more competitive for nitrogen than C. prolifera. In this case, the leaf uptake of ammonium is the largest contributor to the total nitrogen (ammonium plus amino acids) budget of the seagrass. Our results showed that the different nutritional strategies of the 2 macrophytes allow their coexistence in the Ria Formosa lagoon.
- Effects of clam harvesting on sexual reproduction of the seagrass Zostera noltiiPublication . Alexandre, Ana; Santos, Rui; Serrão, EsterThe sexual reproduction of Zostera noltii in the Ria Formosa lagoon (southern Portugal) was assessed by quantifying the variation in both flowering effort and evolution of maturation stages during the fertile season. The effects of clam harvesting on the sexual reproduction of Z. noltii were investigated using both descriptive and experimental approaches. Meadows disturbed by clam harvesting were compared with undisturbed meadows in 2 sectors of the lagoon. In both sectors, disturbed meadows showed significantly lower vegetative shoot density and significantly higher reproductive effort. The reproductive effort in the western disturbed meadows was 2-fold higher than in undisturbed meadows, whereas in eastern disturbed meadows it was 4-fold higher. In disturbed meadows, early maturation stages were found until the end of the flowering season and higher seed production was observed. Both the negative effect of clam harvesting on Z. noltii density and the positive effect on its reproductive effort were confirmed by manipulative field experiments. Plots that were experimentally harvested during the fertile season showed significantly lower vegetative shoot density and higher reproductive effort than unharvested plots. Results suggest that Z. noltii responds to clam harvesting disturbance by both increasing its reproductive effort and extending its fertile season.
- Effects of CO2 enrichment on photosynthesis, growth, and nitrogen metabolism of the seagrass Zostera noltiiPublication . Alexandre, Ana; Silva, Joao; Buapet, Pimchanok; Bjork, Mats; Santos, RuiSeagrass ecosystems are expected to benefit from the global increase in CO2 in the ocean because the photosynthetic rate of these plants may be C-i-limited at the current CO2 level. As well, it is expected that lower external pH will facilitate the nitrate uptake of seagrasses if nitrate is cotransported with H+ across the membrane as in terrestrial plants. Here, we investigate the effects of CO2 enrichment on both carbon and nitrogen metabolism of the seagrass Zostera noltii in a mesocosm experiment where plants were exposed for 5 months to two experimental CO2 concentrations (360 and 700 ppm). Both the maximum photosynthetic rate (Pm) and photosynthetic efficiency (alpha) were higher (1.3- and 4.1-fold, respectively) in plants exposed to CO2-enriched conditions. On the other hand, no significant effects of CO2 enrichment on leaf growth rates were observed, probably due to nitrogen limitation as revealed by the low nitrogen content of leaves. The leaf ammonium uptake rate and glutamine synthetase activity were not significantly affected by increased CO2 concentrations. On the other hand, the leaf nitrate uptake rate of plants exposed to CO2-enriched conditions was fourfold lower than the uptake of plants exposed to current CO2 level, suggesting that in the seagrass Z. noltii nitrate is not cotransported with H+ as in terrestrial plants. In contrast, the activity of nitrate reductase was threefold higher in plant leaves grown at high-CO2 concentrations. Our results suggest that the global effects of CO2 on seagrass production may be spatially heterogeneous and depend on the specific nitrogen availability of each system. Under a CO2 increase scenario, the natural levels of nutrients will probably become limiting for Z. noltii. This potential limitation becomes more relevant because the expected positive effect of CO2 increase on nitrate uptake rate was not confirmed.
- 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.
- 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.
- 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.
- Impacto da actividade de marisqueio na floração e no esforço de reprodução sexual em Zostera noltii Hornemann na Ria FormosaPublication . Alexandre, Ana; Santos, RuiA angiospérmica marinha Zostera noltii cresce e reproduz-se sexualmente nas zonas intertidais da Ria Formosa. A época de floração ocorre entre Março e Novembro. O desenvolvimento das flores ao longo da época de floração, desde a formação da inflorescência até à produção da semente, foi observado e organizado numa escala de maturação. A razão entre o número de flores masculinas e femininas determinado para a Z. noltii foi de 1:1 (4±0.14 S.E. flores masculinas para 4±0.13 S.E. flores femininas). O impacto da actividade de marisqueio na reprodução sexual da espécie foi avaliado. Valores mais elevados de esforço reprodutivo (6.38±3.91%), bem como da densidade de rebentos reprodutores (2200 rebentos por m2), foram observados nos campos mariscados em relação aos campos de controlo (1.85±1.14% de esforço reprodutivo; 1062 rebentos por m2). Os resultados obtidos mostram que a Z. noltii aumenta o seu esforço na reprodução sexual como resposta ao distúrbio provocado pela actividade de marisqueio. No entanto, devido ao rápido crescimento clonal da Z. noltii e aos constrangimentos existentes no seu ciclo reprodutor, a recuperação dos campos mariscados na Ria Formosa é provavelmente efectuada através do desenvolvimento vegetativo da espécie.
- Interactions between seagrasses and seaweeds during surge nitrogen acquisition determine interspecific competitionPublication . Alexandre, Ana; Baeta, Alexandra; Engelen, Aschwin H.; Santos, RuiSeagrasses dominate shallow coastal environments where nitrogen (N) availability in the water column is often sporadic and mainly in the form of pulses. We investigated the N uptake competition between seagrasses and seaweeds through a series of N-15 surge uptake experiments combining single-species and mixed incubations across ammonium concentrations. N surge uptake rates of seagrasses were 2 to 14-fold higher than those of seaweeds in the majority of combinations, showing that seagrasses are generally in a competitive advantage over seaweeds in N-poor environments with N-pulses. No threshold concentration of ammonium was found beyond which seaweeds performed better than seagrasses. Mixed incubations revealed interspecific interactions that affected rates positively and negatively. Uptake rates obtained in single-species incubations, therefore, cannot always be used to predict the outcome of uptake competition. Only two (Zostera marina vs. Ulva rotundata and Zostera marina vs. Codium decorticatum) of the nine combinations tested (Z. marina, Z. noltei and Cymodocea nodosa vs. U. rotundata, C. decorticatum and Dictyota dichotoma) were found to enhance macroalgal uptake. Our results showed that the surge uptake capacity of seagrasses represents an important mechanism in their N acquisition strategy that justifies their dominance in shallow oligotrophic environments.
- 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.
- Marine heatwaves can limit the role of surfgrasses as biofilters for wastewatersPublication . Bonet-Melià, Paula; Sandoval-Gil, Jose Miguel; Samperio-Ramos, Guillermo; Vivanco-Bercovich, Manuel; Canino-Herrera, Sergio R.; Durazo, Reginaldo; Camacho-Ibar, Víctor F.; Alexandre, AnaMarine heatwaves (MHWs) can have detrimental effects on seagrasses, but knowledge about the impacts on their ecosystem services remains scarce. This work evaluated Phyllospadix scouleri (surgrass) as a biofilter for wastewater discharges, and how warming associated with MHW may affect this ecological function. The nitrogen uptake kinetics and assimilation abilities for ammonium, nitrate, and urea were examined under two different warming scenarios (single and repeated events) simulated in a mesocosm. N-uptake kinetics were related to urban sewage discharges close to surfgrass meadows. Our results revealed that surfgrasses can serve as effective biofilters because of their high nitrogen uptake rates and above-average canopy biomass. Nonetheless, exposure to both experimental warmings resulted in a significant decline in their ability to incorporate and assimilate nitrogen. Consequently, MHWs may reduce the capacity of surfgrasses to function as nitrogen sinks and green filters for sewage waters, jeopardizing their role as Blue Nitrogen systems.
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