Os estuários como sumidouros decarbono: fotossíntese, fotoinibição e resiliência da produtividade primária pelágica e bêntica

Estuaries are recognized amongst the most productive ecosystems on Earth. Their high primary productivity is largely due to the photosynthetic carbon fixation by phytoplankton and microphytobenthos, the communities of microalgae and cyanobacteria that inhabit the water column and subtidal or intertidal sediments, respectively. In comparison with the phytoplankton, the microphytobenthos has been much less studied regarding the photophysiological processes affecting primary productivity, and their relative role as contributors to estuarine-level production. One of these processes is photoinhibition, the high light-induced decrease in photosynthetic activity, considered a major limiting factor of growth and primary productivity in the variable and extreme estuarine environment. The detrimental impact of photoinhibition on photosynthesis depends on the balance between the photoinactivation and repair of photosystem II (PSII). By successfully adapting to microphytobenthos a recently-developed methodology based on multi-actinic imaging of chlorophyll fluorescence, this work evaluated their photoacclimation and photoprotective capacity, as measured by the reduction in PSII photoinactivation. PSII photoinactivation and repair was found to vary between different communities, pointing to a trade-off between cellular motility-based and physiological photoprotective mechanisms. Epipelic (motile) species showed a reduced physiological capacity for preventing photodamage, while epipsammic (non-motile) forms appeared less susceptible to photoinactivation and more dependent on physiological photoprotection. This work further investigated an overlooked aspect of microphytobenthos ecology, related to the presence of substantial amounts of microalgal biomass in subsurface sediments. By studying samples from intertidal areas of the Ria de Aveiro (Portugal), this work found that buried cells can quickly regain photosynthetic activity when exposed to surface conditions. Potential viable subsurface (0.5-10 cm) microalgal biomass was found to represent 2-3 times the amount of biomass present at the surface layers (0.0-0.5 cm). These results support the hypothesis that subsurface biomass may play an important ecological role as a source of photosynthetically competent cells capable of ‘reinoculating’ the surface, contributing to the high productivity of intertidal areas. The relative importance of phytoplankton and microphytobenthos as contributors to ecosystem-level primary productivity was evaluated in the Ria de Aveiro, by comparing the spatio-temporal variability of biomass and productivity of different communities. This study made use of a new type of fluorometer allowing the measurement of absolute rates of PSII electron transport rates and the estimation of carbon fixation rates. Biomass-specific productivity rates for phytoplankton and microphytobenthos were found to reach 68.0 and 19.1 mg C mg Chl a-1 d-1, respectively. Annual areal production rates were higher for the microphytobenthos, reaching 105.2 g C m-2 yr-1, as opposed to 49.9 g C m-2 yr-1 for the phytoplankton. The annual rates upscaled for the whole Ria de Aveiro highlight the importance of the intertidal areas as significant carbon sinks and reservoirs of active ‘blue carbon’, and as main sites of primary productivity, found to contribute with more than 60% of the total ecosystem-level budget 12428.3 t C yr-1.Os estuários são reconhecidos como um dos tipos de ecossistemas mais produtivos na Terra. A sua elevada produtividade primária é devida em larga medida à fixação fotossintética de carbono pelo fitoplâncton e microfitobentos, as comunidades de microalgas e cianobactérias que habitam a coluna de àgua e os sedimentos subtidais e intertidais, respetivamente. Em comparação com o fitoplâncton, o microfitobentos tem sido muito menos estudado relativamente aos processos fotofisiológicos que controlam a sua produtividade, bem como à sua contribuição para a produção primária global do estuário. Um destes processos é a fotoinibição, a diminuição da atividade fotossintética causada pela luz, considerada como um importante fator limitante da produtividade primária no ambiente estuarino. O impacto negativo da fotoinibição na fotossíntese depende do balanço entre a fotoinativação e a reparação do fotossistema II (PSII). Baseado num recente método de imagiologia multi-actínica de fluorescência da clorofila, este trabalho avaliou a fotoaclimatação e capacidade de fotoproteção contra a fotoinibição, medida pela redução da fotoinativação do PSII. A fotoinativação e reparação do PSII variou entre diferentes tipos de comunidades, indicando a existência de um balanço entre a fotoproteção baseada na motilidade celular e em mecanismos fisiológicos. Espécies epipélicas (móveis) mostraram uma menor capacidade fisiológica de prevenir danos, enquanto as formas epipsâmicas (imóveis) aparentaram ser menos suscetíveis à fotoinibição e mais dependentes de fotoproteção fisiológica. Este trabalho investigou ainda um aspeto pouco estudado, relacionado com a presença de quantidades substanciais de biomass de microalgas em sedimentos subsuperficias. Pela análise de sedimentos intertidais da Ria de Aveiro (Portugal), foi descoberto que as células enterradas conseguem recuperar rapidamente a sua atividade fotossintética quando expostas a condições da superfície. Foi também concluído que a biomassa subsuperficial (0.5-10 cm) potencialmente viável representa 2-3 vezes a biomassa presente nas camadas superficiais (0.0-0.5 cm). Estes resultados suportam a hipótese de que a biomassa subsuperficial desempenha um papel ecológico importante enquanto fonte de células fotossinteticamente competentes capazes de ‘reinocular’ a superfície, contribuindo para a elevada produtividade das áreas intertidais. A importância relativa da contribuição do fitoplâncton e do microfitobentos para a produtividade primária ao nível do ecossistema foi avaliada para a Ria de Aveiro, comparando a variabilidade espacio-temporal da biomass e produtividade de diferentes comunidades. Este estudo baseou-se na medição de taxas absolutas de transporte de electrões no PSII e a estimação de taxas de fixação de carbono. Por unidade de biomass, estas atingiram 68,0 e 19,1 mg C mg Chl a-1 d-1, para o fitoplâncton e o microfitobentos, respectivamente. Por unidade de área, a produtividade anual foi mais elevada no caso do microfitobentos, atingindo 105,2 g C m-2 yr-1, por oposição a 49,9 g C m-2 yr-1, para o fitoplâncton. Considerando a totalidade da área da Ria de Aveiro, os resultados salientam a importância das áreas intertidais enquanto sumidouros de carbono e reservatórios de “carbono azul”, e locais de elevada produtividade primária, contribuindo com mais de 60% do total anual de 12428,3 t C yr-1.Programa Doutoral em Biologi

Arctic sea ice algae differ markedly from phytoplankton in their ecophysiological characteristics

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Modelling the effect of vertical mixing on bottle incubations for determining in situ phytoplankton dynamics. I. Growth rates

Reliable estimates of in situ phytoplankton growth rates are central to understanding the dynamics of aquatic ecosystems. A common approach for estimating in situ growth rates is to incubate natural phytoplankton assemblages in clear bottles at fixed depths or irradiance levels and measure the change in chlorophyll a (Chl) over the incubation period (typically 24 h). Using a modelling approach, we investigate the accuracy of these Chl-based methods focussing on 2 aspects: (1) in a freely mixing surface layer, the cells are typically not in balanced growth, and with photoacclimation, changes in Chl may yield different growth rates than changes in carbon; and (2) the in vitro methods neglect any vertical movement due to turbulence and its effect on the cells' light history. The growth rates thus strongly depend on the incubation depth and are not necessarily representative of the depth-integrated in situ growth rate in the freely mixing surface layer. We employ an individual based turbulence and photosynthesis model, which also accounts for photoacclimation and photo - inhibition, to show that the in vitro Chl-based growth rate can differ both from its carbon-based in vitro equivalent and from the in situ value by up to 100%, depending on turbulence intensity, optical depth of the mixing layer, and incubation depth within the layer. We make recommendations for choosing the best depth for single-depth incubations. Furthermore we demonstrate that, if incubation bottles are being oscillated up and down through the water column, these systematic errors can be significantly reduced. In the present study, we focus on Chl-based methods only, while productivity measurements using carbon-based techniques (e.g. 14C) are discussed in Ross et al. (2011; Mar Ecol Prog Ser 435:33-45). © Inter-Research 2011

Photosynthetic responses in Phaeocystis antarctica towards varying light and iron conditions

The effects of iron limitation on photoacclimation to a dynamic light regime were studied in Phaeocystis antarctica. Batch cultures were grown under a sinusoidal light regime, mimicking vertical mixing, under both iron-sufficient and -limiting conditions. Iron-replete cells responded to changes in light intensity by rapid xanthophyll cycling. Maximum irradiance coincided with maximum ratios of diatoxanthin/diadinoxanthin (dt/dd). The maximum quantum yield of photosynthesis (F-v /F-m) was negatively related to both irradiance and dt/dd. Full recovery of F-v /F-m by the end of the light period suggested successful photoacclimation. Iron-limited cells displayed characteristics of high light acclimation. The ratio of xanthophyll pigments to chlorophyll a was three times higher compared to iron-replete cells. Down-regulation of photosynthetic activity was moderated. It is argued that under iron limitation cells maintain a permanent state of high energy quenching to avoid photoinhibition during exposure to high irradiance. Iron-limited cells could maintain a high growth potential due to an increased absorption capacity as recorded by in vivo absorption, which balanced a decrease in F-v/F-m . The increase in the chlorophyll a-specific absorption cross section was related to an increase in carotenoid pigments and a reduction in the package effect. These experiments show that P. antarctica can acclimate successfully to conditions as they prevail in the Antarctic ocean, which may explain the success of this species