The operation of photosynthetic energy-dissipating processes is commonly characterized by measuri... more The operation of photosynthetic energy-dissipating processes is commonly characterized by measuring the light response of the nonphotochemical quenching (NPQ) of chlorophyll fluorescence, or NPQ versus E curves. This study proposes a mathematical model for the quantitative description of the generic NPQ versus E curve. The model is an adaptation of the Hill equation and is based on the close dependence of NPQ on the xanthophyll cycle (XC). The model was tested on NPQ versus E curves measured in the plant Arabidopsis thaliana and the diatom Nitzschia palea, representing the two main types of XC, the violaxanthin–antheraxanthin–zeaxanthin (VAZ) type and the diadinoxanthin–diatoxanthin (DD–DT) type, respectively. The model was also fitted to a large number of published light curves, covering the widest possible range of XC types, taxa, growth conditions, and experimental protocol of curve generation. The model provided a very good fit to experimental and published data, coping with the large variability in curve characteristics. The model was further used to quantitatively compare the light responses of NPQ and of PSII electron transport rate, ETR, through the use of indices combining parameters of the models describing the two types of light–response curves. Their application to experimental and published data showed a systematic large delay of the buildup of NPQ relatively to the saturation of photochemistry. It was found that when ETR reaches saturation, NPQ is on average still below one fifth of its maximum attainable level, which is only reached at irradiances about three times higher. It was also found that organisms having the DD–DT type of XC appeared to be able to start operating the XC at lower irradiances than those of the VAZ type.
Community assemblages of diatoms, green algae and cyanobacteria comprise the microphytobenthos (M... more Community assemblages of diatoms, green algae and cyanobacteria comprise the microphytobenthos (MPB), which inhabit benthic sediment ecosystems (Admiraal 1984; Underwood and Kromkamp 1999; Consalvey et al. 2004). Particular attention has been paid to the analysis of intertidal soft sediment systems, e.g. cohesive mudflat and sandy substrata typical of estuarine habitats. Variable chlorophyll fluorescence has been applied to these systems since the 1990s, in an attempt to investigate the primary productivity and photophysiology of the integrated biofilms, when viewed as a “black box system”, and also at the species level (Sections 5, 6 and 7). These transient (i.e. temporary) biofilms are not confined to such soft sediment habitats however, and more recently application of fluorescence methodologies has been applied to biofilms inhabiting rocky shores and stromatolite systems (Kromkamp et al. 2007; Perkins et al. 2007). However the large majority of published work has centred upon benthic soft-sediment biofilms, due to their important ecosystem functions of carbon flow and sediment stability (Underwood and Kromkamp 1999). In the former their high magnitude of productivity fuels carbon flow through invertebrate and bacterial food webs to support important trophic levels of anthropogenically exploited taxa, including coastal fish and shell fisheries and coastal avifauna. In the case of sediment stability, biogenic exopolymers, usually referred to as extracellular polymeric substances (EPS), produced by the MPB in part to facilitate mobility, may contribute significantly to sediment stability, hence increasing the sediment resistance to hydrodynamic stresses and thus resistance to coastal erosion (e.g. Underwood and Kromkamp 1999 and citations there-in). Finally, the photosynthetic production of oxygen can be regarded as an important ecosystem function.
A study on spatio-temporal distribution of microphytobhethos in intertidal zones of Tagus Estuary... more A study on spatio-temporal distribution of microphytobhethos in intertidal zones of Tagus Estuary was carried out from 1990 to 1992. Near Lisbon, Portugal, Tagus Estuary is a shallow mesotidal estuary, covering an area of 320 km2. The intertidal area ranges from 20 to 40% off the total area and it is constituted mainly by mudflats. Intertidal flats are richly populated by microalgae, diatoms being the most important and ubiquitos group. Spatial variation of microphytobethos was studied in spring 1990, 21 different sites were sampled. Microphytobenthos biomass was evaluated as chlorophyll a content of the surface centimeter, ranging from 10 to 240 mg m−2. A Principal Component Analysis showed that 62% of the total variability found in intertidal flats of Tagus estuary could be attributed to two major factors: sediment type and tidal height. A hierarchical grouping defined 3 major groups of similar stations, each one representing a different strata of the ecosystem. One station from each group was chosen for the study of the temporal variation. A sampling, rogram took place from April 1991 to April 1992, with fortnightly sampling, the Chl a ranged from 20–300 mg m−2. No clear seasonal variation was found, and our results indicated that tidal height of sampledsite played an essential role in temporal biomass evolution, thus upper littoral sites were influenced by climatic parameters, whereas in lower sites action of tides mainly controlled microphytic biomass. Une étude sur l'hétérogénéité spatio-temporelle du microphytobenthos dans les sédiments intertidaux de l'Estuaire du Tage a été accompli de 1990 á 1992. L'Estuaire du Tage, prés de Lisbonne (Portugal) est un estuaire peu profond, mesotidal, avec une aire total de 320 km2. L'aire intertidale est comprise entre 20 et 40% du total, et constituéé surtout par des vasiéres. Ces slikkes sont peuplées par une communauté assez riche de microalgues, ou les diatomées sont les plus abundantes. La variation spatialle du microphytobenthos était évalué au Printemps 1990, ou 21 différentes stations étaient échantillonnées. La biomasse était évalué par la concentration enchlorophylle a du premier centimétre de sédiment, qui a varié de 10 á 240 mg Chl a m−2. Une Analyse en Composants Principales a montré que 62% de la variabilité de la biomasse était lié á deux facteurs: le sédiment et l'hauteur vis-á-vis la marée. Une classification hiérarchique des stations par similitude a établi 3 groupes principaux, représantantles différents strates de écecosytéme. Une station de chaque groupement a été choisie pour l'étude de la variation temporelle, qui s'est deroulé d'avril 1991 á avril 1992, avec des prélévements deux fois par mois. Les valeurs de Chl a obtenus vont de 20 á 300 mg m−2. Les variations saisonniéres observées ne sont pas claires: nos résultats indiquent que l'hauteur de la station (m) joue un rôle essentiel dans l'évolution temporel de la biomasse, c'est á dire, la biomasse microalgal des sites du supra-littoral est influencié par les paramétres climatiques, tandis que dans l'infra-littoral c'est l'action des marées le facteur principal.
The communities of benthic microalgae that form dense biofilms at the surface of aquatic sediment... more The communities of benthic microalgae that form dense biofilms at the surface of aquatic sediments, or microphytobenthos, are important primary producers in estuarine intertidal flats and shallow coastal waters.
Rapid light-response curves (RLC) of variable chlorophyll fluorescence were measured on estuarine... more Rapid light-response curves (RLC) of variable chlorophyll fluorescence were measured on estuarine benthic microalgae with the purpose of characterising its response to changes in ambient light, and of investigating the relationship to steady-state light-response curves (LC). The response of RLCs to changes in ambient light (E, defined as the irradiance level to which a sample is acclimated to prior to the start of the RLC) was characterised by constructing light-response curves for the RLC parameters α RLC, the initial slope, ETRm,RLC, the maximum relative electron transport rate, and E k,RLC, the light-saturation parameter. Measurements were carried out on diatom-dominated suspensions of benthic microalgae and RLC and LC parameters were compared for a wide range of ambient light conditions, time of day, season and sample taxonomic composition. The photoresponse of RLC parameters was typically bi-phasic, consisting of an initial increase of all parameters under low ambient light (E < 21–181 μmol m−2 s−1), and of a phase during which α RLC decreased significantly with E, and the increase of ETRm,RLC and E k,RLC was attenuated. The relationship between RLC and LC parameters was dependent on ambient irradiance, with significant correlations being found between α RLC and α, and between ETRm,RLC and ETRm, for samples acclimated to low and to high ambient irradiances, respectively. The decline of α RLC under high light (Δα RLC) was strongly correlated (P < 0.001 in all cases) with the level of non-photochemical quenching (NPQ) measured before each RLC. These results indicate the possibility of using RLCs to characterise the steady-state photoacclimation status of a sample, by estimating the LC parameter E k, and to trace short-term changes in NPQ levels without dark incubation.
Abstract The effects of light exposure on the photosynthetic activity of kleptoplasts were studie... more Abstract The effects of light exposure on the photosynthetic activity of kleptoplasts were studied in the sacoglossan mollusc Elysia viridis. The photosynthetic activity of ingested chloroplasts was assessed in vivo by non-destructively measuring photophysiological parameters using pulse amplitude modulation (PAM) fluorometry.
Coastal and estuarine intertidal sediments are commonly colonized by dense populations of microph... more Coastal and estuarine intertidal sediments are commonly colonized by dense populations of microphytobenthos. Due to wind and tides, important fractions of microphytobenthic populations may be buried. A mathematical model describing the depth variation of chlorophyll a in intertidal sediments was developed and experimentally tested. The model assumed first-order chlorophylla degradation and a constant mean burial velocity which resulted in a negative exponential variationC Z =C O exp{-k/vz} (C Z andC O =chlorophylla concentration at depth zand at the surface;k=specific degradation rate of chlorophyll a to pheopigments;V=mean burial velocity). Chlorophylla concentration depth profiles in different sediment types measured at the Tagus estuary and Ria Formosa (Portugal) were used to validate the model. The model was adjusted to field data. The chlorophyll a degradation rate was measured in a microcosm experiment under total darkness and no tidal action, and sampled during three months. This rate was shown to be independent of time and depth for the upper 0–15 mm depth interval. This result allowed the estimation ofV for each sampling site. Comparison of predicted and observed temporal data further confirmed the validity of the model andk andV values. Despite its simplicity, the proposed model adequately described the depth distribution of chlorophylla in different types of intertidal sediments. The model allowed the quantitative characterization of the buried microphytobenthic biomass (depth-integrated biomass) and the assessment of its importance as potentially productive stock of cells.
ABSTRACT In vivo chlorophyll (Chl) a fluorescence was measured in undisturbed intertidal sediment... more ABSTRACT In vivo chlorophyll (Chl) a fluorescence was measured in undisturbed intertidal sediments with the purpose of tracing the vertical migratory rhythms of benthic microalgae. A pulse amplitude fluorometer, an instrument which does not require physical contact with the sample, was used, thus allowing successive measurements to be taken on the same sample without causing any type of disturbance to the sediment structure. The basis of the method is the possibility to detect changes in the Chl a concentration near the sediment surface ...
Photosynthesis-light response curves of Gelidium sesquipedale from the west coast of Portugal (Ca... more Photosynthesis-light response curves of Gelidium sesquipedale from the west coast of Portugal (Cape Espichel) were determined at four different depths, 3, 10, 15 and 22 m. Data acquisition using chlorophyll a fluorescence methodology and oxygen electrode measurements were compared. Response curves were determined over an increasing range of irradiance values (I), from darkness to 900 μmol photon m-2 s-1 PAR. In general, light response curves obtained for G. sesquipedale showed a similar pattern whether determined by the chlorophyll fluorescence method or by oxygen evolution. The photosynthetic capacity of G. sesquipedale decreased with depth, as expected, revealing a ‘sun’ and ‘shade’ acclimation pattern, between shallow and deeper waters.
The operation of photosynthetic energy-dissipating processes is commonly characterized by measuri... more The operation of photosynthetic energy-dissipating processes is commonly characterized by measuring the light response of the nonphotochemical quenching (NPQ) of chlorophyll fluorescence, or NPQ versus E curves. This study proposes a mathematical model for the quantitative description of the generic NPQ versus E curve. The model is an adaptation of the Hill equation and is based on the close dependence of NPQ on the xanthophyll cycle (XC). The model was tested on NPQ versus E curves measured in the plant Arabidopsis thaliana and the diatom Nitzschia palea, representing the two main types of XC, the violaxanthin–antheraxanthin–zeaxanthin (VAZ) type and the diadinoxanthin–diatoxanthin (DD–DT) type, respectively. The model was also fitted to a large number of published light curves, covering the widest possible range of XC types, taxa, growth conditions, and experimental protocol of curve generation. The model provided a very good fit to experimental and published data, coping with the large variability in curve characteristics. The model was further used to quantitatively compare the light responses of NPQ and of PSII electron transport rate, ETR, through the use of indices combining parameters of the models describing the two types of light–response curves. Their application to experimental and published data showed a systematic large delay of the buildup of NPQ relatively to the saturation of photochemistry. It was found that when ETR reaches saturation, NPQ is on average still below one fifth of its maximum attainable level, which is only reached at irradiances about three times higher. It was also found that organisms having the DD–DT type of XC appeared to be able to start operating the XC at lower irradiances than those of the VAZ type.
Community assemblages of diatoms, green algae and cyanobacteria comprise the microphytobenthos (M... more Community assemblages of diatoms, green algae and cyanobacteria comprise the microphytobenthos (MPB), which inhabit benthic sediment ecosystems (Admiraal 1984; Underwood and Kromkamp 1999; Consalvey et al. 2004). Particular attention has been paid to the analysis of intertidal soft sediment systems, e.g. cohesive mudflat and sandy substrata typical of estuarine habitats. Variable chlorophyll fluorescence has been applied to these systems since the 1990s, in an attempt to investigate the primary productivity and photophysiology of the integrated biofilms, when viewed as a “black box system”, and also at the species level (Sections 5, 6 and 7). These transient (i.e. temporary) biofilms are not confined to such soft sediment habitats however, and more recently application of fluorescence methodologies has been applied to biofilms inhabiting rocky shores and stromatolite systems (Kromkamp et al. 2007; Perkins et al. 2007). However the large majority of published work has centred upon benthic soft-sediment biofilms, due to their important ecosystem functions of carbon flow and sediment stability (Underwood and Kromkamp 1999). In the former their high magnitude of productivity fuels carbon flow through invertebrate and bacterial food webs to support important trophic levels of anthropogenically exploited taxa, including coastal fish and shell fisheries and coastal avifauna. In the case of sediment stability, biogenic exopolymers, usually referred to as extracellular polymeric substances (EPS), produced by the MPB in part to facilitate mobility, may contribute significantly to sediment stability, hence increasing the sediment resistance to hydrodynamic stresses and thus resistance to coastal erosion (e.g. Underwood and Kromkamp 1999 and citations there-in). Finally, the photosynthetic production of oxygen can be regarded as an important ecosystem function.
A study on spatio-temporal distribution of microphytobhethos in intertidal zones of Tagus Estuary... more A study on spatio-temporal distribution of microphytobhethos in intertidal zones of Tagus Estuary was carried out from 1990 to 1992. Near Lisbon, Portugal, Tagus Estuary is a shallow mesotidal estuary, covering an area of 320 km2. The intertidal area ranges from 20 to 40% off the total area and it is constituted mainly by mudflats. Intertidal flats are richly populated by microalgae, diatoms being the most important and ubiquitos group. Spatial variation of microphytobethos was studied in spring 1990, 21 different sites were sampled. Microphytobenthos biomass was evaluated as chlorophyll a content of the surface centimeter, ranging from 10 to 240 mg m−2. A Principal Component Analysis showed that 62% of the total variability found in intertidal flats of Tagus estuary could be attributed to two major factors: sediment type and tidal height. A hierarchical grouping defined 3 major groups of similar stations, each one representing a different strata of the ecosystem. One station from each group was chosen for the study of the temporal variation. A sampling, rogram took place from April 1991 to April 1992, with fortnightly sampling, the Chl a ranged from 20–300 mg m−2. No clear seasonal variation was found, and our results indicated that tidal height of sampledsite played an essential role in temporal biomass evolution, thus upper littoral sites were influenced by climatic parameters, whereas in lower sites action of tides mainly controlled microphytic biomass. Une étude sur l'hétérogénéité spatio-temporelle du microphytobenthos dans les sédiments intertidaux de l'Estuaire du Tage a été accompli de 1990 á 1992. L'Estuaire du Tage, prés de Lisbonne (Portugal) est un estuaire peu profond, mesotidal, avec une aire total de 320 km2. L'aire intertidale est comprise entre 20 et 40% du total, et constituéé surtout par des vasiéres. Ces slikkes sont peuplées par une communauté assez riche de microalgues, ou les diatomées sont les plus abundantes. La variation spatialle du microphytobenthos était évalué au Printemps 1990, ou 21 différentes stations étaient échantillonnées. La biomasse était évalué par la concentration enchlorophylle a du premier centimétre de sédiment, qui a varié de 10 á 240 mg Chl a m−2. Une Analyse en Composants Principales a montré que 62% de la variabilité de la biomasse était lié á deux facteurs: le sédiment et l'hauteur vis-á-vis la marée. Une classification hiérarchique des stations par similitude a établi 3 groupes principaux, représantantles différents strates de écecosytéme. Une station de chaque groupement a été choisie pour l'étude de la variation temporelle, qui s'est deroulé d'avril 1991 á avril 1992, avec des prélévements deux fois par mois. Les valeurs de Chl a obtenus vont de 20 á 300 mg m−2. Les variations saisonniéres observées ne sont pas claires: nos résultats indiquent que l'hauteur de la station (m) joue un rôle essentiel dans l'évolution temporel de la biomasse, c'est á dire, la biomasse microalgal des sites du supra-littoral est influencié par les paramétres climatiques, tandis que dans l'infra-littoral c'est l'action des marées le facteur principal.
The communities of benthic microalgae that form dense biofilms at the surface of aquatic sediment... more The communities of benthic microalgae that form dense biofilms at the surface of aquatic sediments, or microphytobenthos, are important primary producers in estuarine intertidal flats and shallow coastal waters.
Rapid light-response curves (RLC) of variable chlorophyll fluorescence were measured on estuarine... more Rapid light-response curves (RLC) of variable chlorophyll fluorescence were measured on estuarine benthic microalgae with the purpose of characterising its response to changes in ambient light, and of investigating the relationship to steady-state light-response curves (LC). The response of RLCs to changes in ambient light (E, defined as the irradiance level to which a sample is acclimated to prior to the start of the RLC) was characterised by constructing light-response curves for the RLC parameters α RLC, the initial slope, ETRm,RLC, the maximum relative electron transport rate, and E k,RLC, the light-saturation parameter. Measurements were carried out on diatom-dominated suspensions of benthic microalgae and RLC and LC parameters were compared for a wide range of ambient light conditions, time of day, season and sample taxonomic composition. The photoresponse of RLC parameters was typically bi-phasic, consisting of an initial increase of all parameters under low ambient light (E < 21–181 μmol m−2 s−1), and of a phase during which α RLC decreased significantly with E, and the increase of ETRm,RLC and E k,RLC was attenuated. The relationship between RLC and LC parameters was dependent on ambient irradiance, with significant correlations being found between α RLC and α, and between ETRm,RLC and ETRm, for samples acclimated to low and to high ambient irradiances, respectively. The decline of α RLC under high light (Δα RLC) was strongly correlated (P < 0.001 in all cases) with the level of non-photochemical quenching (NPQ) measured before each RLC. These results indicate the possibility of using RLCs to characterise the steady-state photoacclimation status of a sample, by estimating the LC parameter E k, and to trace short-term changes in NPQ levels without dark incubation.
Abstract The effects of light exposure on the photosynthetic activity of kleptoplasts were studie... more Abstract The effects of light exposure on the photosynthetic activity of kleptoplasts were studied in the sacoglossan mollusc Elysia viridis. The photosynthetic activity of ingested chloroplasts was assessed in vivo by non-destructively measuring photophysiological parameters using pulse amplitude modulation (PAM) fluorometry.
Coastal and estuarine intertidal sediments are commonly colonized by dense populations of microph... more Coastal and estuarine intertidal sediments are commonly colonized by dense populations of microphytobenthos. Due to wind and tides, important fractions of microphytobenthic populations may be buried. A mathematical model describing the depth variation of chlorophyll a in intertidal sediments was developed and experimentally tested. The model assumed first-order chlorophylla degradation and a constant mean burial velocity which resulted in a negative exponential variationC Z =C O exp{-k/vz} (C Z andC O =chlorophylla concentration at depth zand at the surface;k=specific degradation rate of chlorophyll a to pheopigments;V=mean burial velocity). Chlorophylla concentration depth profiles in different sediment types measured at the Tagus estuary and Ria Formosa (Portugal) were used to validate the model. The model was adjusted to field data. The chlorophyll a degradation rate was measured in a microcosm experiment under total darkness and no tidal action, and sampled during three months. This rate was shown to be independent of time and depth for the upper 0–15 mm depth interval. This result allowed the estimation ofV for each sampling site. Comparison of predicted and observed temporal data further confirmed the validity of the model andk andV values. Despite its simplicity, the proposed model adequately described the depth distribution of chlorophylla in different types of intertidal sediments. The model allowed the quantitative characterization of the buried microphytobenthic biomass (depth-integrated biomass) and the assessment of its importance as potentially productive stock of cells.
ABSTRACT In vivo chlorophyll (Chl) a fluorescence was measured in undisturbed intertidal sediment... more ABSTRACT In vivo chlorophyll (Chl) a fluorescence was measured in undisturbed intertidal sediments with the purpose of tracing the vertical migratory rhythms of benthic microalgae. A pulse amplitude fluorometer, an instrument which does not require physical contact with the sample, was used, thus allowing successive measurements to be taken on the same sample without causing any type of disturbance to the sediment structure. The basis of the method is the possibility to detect changes in the Chl a concentration near the sediment surface ...
Photosynthesis-light response curves of Gelidium sesquipedale from the west coast of Portugal (Ca... more Photosynthesis-light response curves of Gelidium sesquipedale from the west coast of Portugal (Cape Espichel) were determined at four different depths, 3, 10, 15 and 22 m. Data acquisition using chlorophyll a fluorescence methodology and oxygen electrode measurements were compared. Response curves were determined over an increasing range of irradiance values (I), from darkness to 900 μmol photon m-2 s-1 PAR. In general, light response curves obtained for G. sesquipedale showed a similar pattern whether determined by the chlorophyll fluorescence method or by oxygen evolution. The photosynthetic capacity of G. sesquipedale decreased with depth, as expected, revealing a ‘sun’ and ‘shade’ acclimation pattern, between shallow and deeper waters.
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