Short-term variations in the physiological state of phytoplankton [2pt] in a shallow temperate estuary

Short-term changes in the photosynthetic carbon metabolism and physiological state of phytoplankton were studied over a summer fortnight-long period in the Urdaibai estuary (Bay of Biscay) and related to observed environmental patterns. Day-to-day variability in the hydrographical and biological features of the estuary during the study period was due to changes in meteorological and tidal conditions. Phytoplankton biomass and primary production increased with the improvement of weather, i.e., light conditions, during neap tides. Thus a mixed bloom of cryptophyceans, Euglena sp., and the dinoflagellate Peridinium foliaceum developed in the middle and upper estuary. Photosynthetic responses of phytoplankton were related to the time-scale of changes in light regime. Allocation of photosynthate to major macromolecular classes (LMWM, lipid, polysaccharide, and protein), like phytoplankton biomass and primary production, showed strong spatio-temporal variability. High carbon fixation into low molecular weight metabolites was associated with growth limitation by low light. The relative incorporation of photosynthetic carbon into proteins increased at the beginning of the phytoplankton bloom but overall, it was rather constant. However, carbon allocation into storage products such us lipid or polysaccharide increased when carbon and energy produced under optimal growth conditions exceeded what could be assimilated into protein. These patterns are explained by both spatio-temporal changes in the environmental conditions and species-specific differences. In general, daily variability appeared to be more important than diurnal periodicity in the physiological responses of phytoplankton. Results from this study show that phytoplankton photosynthesis and carbon metabolism are simultaneously affected by biotic and abiotic factors, although short-term light fluctuations may have a major influence on the physiological state of phytoplankton in the Urdaibai estuary.     

Diversity patterns of trait-based phytoplankton functional groups in two basins of a large, shallow lake (Lake Balaton, Hungary) with different trophic state

The application of functional approaches in understanding phytoplankton community-level responses to changes in the environment has become increasingly widespread in recent years. Eutrophication is known to cause profound modifications in ecosystem processes; however, the underlying relationships between environmental drivers and phytoplankton diversity and functioning are complex and largely unknown. Therefore, in the present study, we investigated and compared the temporal diversity patterns of phytoplankton functional groups in the mesotrophic eastern and eutrophic western basin of the shallow Lake Balaton situated in Hungary. Diversity data were derived from taxonomic composition and biomass data corresponding to the years 2005–2006 and 2008–2009. With the use of cluster analysis, phytoplankton species were classified into eight distinct groups representing different combinations of functionally relevant traits including greatest axial linear dimension; surface-to-volume ratio; photosynthetic pigment composition; N₂ fixation; phagotrophic potential; growth form/complexity (unicell, filamentous, colony-, or coenobium-forming); and motility. Our results have revealed that there is a significant inverse relationship between the functional group diversity used in our study and trophic state (total phytoplankton biomass) as opposed to species diversity, where no correlation was observed. In addition, group evenness showed an even stronger negative correlation with trophic state, while species evenness yielded only a weak relationship. The observed variability in functional group diversity suggests that such an approach could provide an efficient means of revealing structural changes in phytoplankton communities, establishing new hypotheses and highlighting fundamental points in ecosystem functioning.     

The role of phytoplankton photosynthesis in global biogeochemical cycles

Phytoplankton biomass in the world's oceans amounts to only 1–2% of the total global plant carbon, yet these organisms fix between 30 and 50 billion metric tons of carbon annually, which is about 40% of the total. On geological time scales there is profound evidence of the importance of phytoplankton photosynthesis in biogeochemical cycles. It is generally assumed that present phytoplankton productivity is in a quasi steady-state (on the time scale of decades). However, in a global context, the stability of oceanic photosynthetic processes is dependent on the physical circulation of the upper ocean and is therefore strongly influenced by the atmosphere. The net flux of atmospheric radiation is critical to determining the depth of the upper mixed layer and the vertical fluxes of nutrients. These latter two parameters are keys to determining the intensity, and spatial and temporal distributions of phytoplankton blooms. Atmospheric radiation budgets are not in steady-state. Driven largely by anthropogenic activities in the 20th century, increased levels of IR- absorbing gases such as CO₂, CH₄ and CFC's and NO_x will potentially increase atmospheric temperatures on a global scale. The atmospheric radiation budget can affect phytoplankton photosynthesis directly and indirectly. Increased temperature differences between the continents and oceans have been implicated in higher wind stresses at the ocean margins. Increased wind speeds can lead to higher nutrient fluxes. Throughout most of the central oceans, nitrate concentrations are sub-micromolar and there is strong evidence that the quantum efficiency of Photosystem II is impaired by nutrient stress. Higher nutrient fluxes would lead to both an increase in phytoplankton biomass and higher biomass-specific rates of carbon fixation. However, in the center of the ocean gyres, increased radiative heating could reduce the vertical flux of nutrients to the euphotic zone, and hence lead to a reduction in phytoplankton carbon fixation. Increased desertification in terrestrial ecosystems can lead to increased aeolean loadings of essential micronutrients, such as iron. An increased flux of aeolean micronutrients could fertilize nutrient-replete areas of the open ocean with limiting trace elements, thereby stimulating photosynthetic rates. The factors which limit phytoplankton biomass and photosynthesis are discussed and examined with regard to potential changes in the Earth climate system which can lead the oceans away from steady-state. While it is difficult to confidently deduce changes in either phytoplankton biomass or photosynthetic rates on decadal time scales, time-series analysis of ocean transparency data suggest long-term trends have occurred in the North Pacific Ocean in the 20th century. However, calculations of net carbon uptake by the oceans resulting from phytoplankton photosynthesis suggest that without a supply of nutrients external to the ocean, carbon fixation in the open ocean is not presently a significant sink for excess atmospheric CO₂.The submitted paper has been authored under Contract No. DE-AC02-76H00016 with the US Department of Energy. Accordingly, the US Government retains a non-exclusive, royalty-free license to publish or reproduce the published form of this contribution, or allow others to do so, for US Government purposes.     

Fatty Acids in Lipids from Particulate Organic Matter in a Eutrophic Lake,Lake Nakanuma,Japan

The depth profiles and seasonal abundance of fatty acids in lipids from the particulate organic matter in the eutrophic Lake Nakanuma, Japan, were closely related to both photosynthetic activity and the distribution of phytoplankton. The changes in the relative proportions of the fatty acids suggest that their synthesis is directly related to the photosynthetic activity of phytoplankton.     

Functional Groups in the Protozoa: Roles in Differing Ecosystems1,2

Feeding habits of freshwater protozoa were used to group species into functional, trophic groups. Community structure in differing ecosystems was examined in relation to the number of species occurring in the functional group categories. Six wetland ecosystems and a large river ecosystem were studied. Changes in community structure during the colonization of artificial substrates were also examined. Changes during colonization were studied in a mesotrophic lake, in low-order streams, and in laboratory microecosystems. In the latter case, the response of colonizing communities to a heavy metal toxicant was studied. All communities studied were dominated by bactivorous-detritivorous species and, to a lesser extent, by photosynthetic species. The chief functional role of substrate-associated protozoans appears to be the processing of dead organic matter and its associated bacterial flora. Functional groups utilizing resources other than detrital or mineral nutrients (saprotrophs, algivores, omnivores, and predators) were always minor community components. Colonizing communities were often dominated by photosynthetic species during early colonization stages but were again dominated by bactivorous-detritivorous species at species equilibrium. Low levels of toxicant (Cd) reduced numbers of both photosynthetic and bactivorous-detritivorous species. Higher toxicant levels virtually eliminated photosynthetic species and reduced bacterial detritivores by over one-half. Roles of protozoan species in ecosystems are closely tied to the processing of detritus and the recycling of mineral nutrients. Enumeration of individuals in functional categories is proposed as a simplified method for studying the abundance and activity of protozoa in ecosystems. Examination of changes in functional group composition and the relationship of functional group abundances to rates of carbon processing are suggested for studies of the importance of protozoa to the flow of energy and materials in ecosystems.     

A dynamic model of darkness tolerance for phytoplankton: model description

To analyze various effects of prolonged darkness on phytoplankton population dynamics, we developed a dynamic model of darkness tolerance for phytoplankton and investigated its characteristics. To construct the basic concepts of the model, we categorized various changes in abundance of phytoplankton both during prolonged darkness and after reillumination into several patterns, and then considered the physiological processes producing these patterns. The nature of darkness tolerance was considered to incorporate previously experienced light conditions, including darkness, as a physiological activity, and members of the same phytoplankton species exhibit different dynamics even in identical light conditions due to such career effects. Taking this into consideration, we developed a cell quota model in relation to darkness tolerance. State variables for abundance were indicated by cell numbers, and physiological condition by three intracellular carbon pools with different physiological functions. Using our model, we analyzed the various changes in abundance of phytoplankton in relation to exposure to prolonged darkness. Various responses in terms of phytoplankton abundance to prolonged darkness and after reillumination were successfully reproduced by the model that simply assumed that deterioration of physiological mechanics, such as photosynthetic functions, was due to a prolonged dark condition. On the basis of the results of calculation and assumptions for the model, we discuss the limitations, problems, and effectiveness of the model. Handling editor: Luigi Naselli-Flores     

Changes in phytoplankton ecophysiology across a coastal upwelling front

The abundance, taxonomic composition and patterns of macromolecularsynthesis of phytoplankton were determined across an upwelling-inducedthermal front in the central Cantabrian Sea (southern Bay ofBiscay) during July 1993. Enhanced levels of phytoplankton biomass,diatom abundance and photosynthetic rate were measured on thecoastal side of the front. Relative carbon (C) incorporationinto proteins increased noticeably on the oceanic side, takingvalues of up to 64%, whereas changes in the relative C incorporationinto lipids and low-molecular-weight metabolites followed anopposite trend. Phytoplankton cells on the oceanic side of thefront were adapted to the prevailing growth-limiting conditionsby maintaining the synthesis of functionally essential molecules—proteins—ratherthan the synthesis of storage compounds. As a result, the carbonto nitrogen uptake ratio varied from {small tilde}5.7 in offshorewaters to 8.0 in the nearshore region. Our results suggest thatthe taxonomic and physiological changes in phytoplankton assemblagesas a response to upwelling may result in an increase in thesynthesis of organic C relative to the upward flux of nitrate.     

The hidden function of photosynthesis: a sensing system for environmental conditions that regulates plant acclimation responses

Plants convert light energy from the sun into chemical energy by photosynthesis. Since they are sessile, they have to deal with a wide range of conditions in their immediate environment. Many abiotic and biotic parameters exhibit considerable fluctuations which can have detrimental effects especially on the efficiency of photosynthetic light harvesting. During evolution, plants, therefore, evolved a number of acclimation processes which help them to adapt photosynthesis to such environmental changes. This includes protective mechanisms such as excess energy dissipation and processes supporting energy redistribution, e.g. state transitions or photosystem stoichiometry adjustment. Intriguingly, all these responses are triggered by photosynthesis itself via the interplay of its light reaction and the Calvin–Benson cycle with the residing environmental condition. Thus, besides its primary function in harnessing and converting light energy, photosynthesis acts as a sensing system for environmental changes that controls molecular acclimation responses which adapt the photosynthetic function to the environmental change. Important signalling parameters directly or indirectly affected by the environment are the pH gradient across the thylakoid membrane and the redox states of components of the photosynthetic electron transport chain and/or electron end acceptors coupled to it. Recent advances demonstrate that these signals control post-translational modifications of the photosynthetic protein complexes and also affect plastid and nuclear gene expression machineries as well as metabolic pathways providing a regulatory framework for an integrated response of the plant to the environment at all cellular levels.     

The hidden function of photosynthesis: a sensing system for environmental conditions that regulates plant acclimation responses

Plants convert light energy from the sun into chemical energy by photosynthesis. Since they are sessile, they have to deal with a wide range of conditions in their immediate environment. Many abiotic and biotic parameters exhibit considerable fluctuations which can have detrimental effects especially on the efficiency of photosynthetic light harvesting. During evolution, plants, therefore, evolved a number of acclimation processes which help them to adapt photosynthesis to such environmental changes. This includes protective mechanisms such as excess energy dissipation and processes supporting energy redistribution, e.g. state transitions or photosystem stoichiometry adjustment. Intriguingly, all these responses are triggered by photosynthesis itself via the interplay of its light reaction and the Calvin-Benson cycle with the residing environmental condition. Thus, besides its primary function in harnessing and converting light energy, photosynthesis acts as a sensing system for environmental changes that controls molecular acclimation responses which adapt the photosynthetic function to the environmental change. Important signalling parameters directly or indirectly affected by the environment are the pH gradient across the thylakoid membrane and the redox states of components of the photosynthetic electron transport chain and/or electron end acceptors coupled to it. Recent advances demonstrate that these signals control post-translational modifications of the photosynthetic protein complexes and also affect plastid and nuclear gene expression machineries as well as metabolic pathways providing a regulatory framework for an integrated response of the plant to the environment at all cellular levels.     

Light control on phytoplankton production in a shallow and turbid estuarine system

Tagus estuary is one of the largest estuaries of Western Europe. With the aim of unravelling the drivers of primary production in this shallow and turbid nutrient replete estuary, we tested the hypothesis that light availability is a major factor controlling phytoplankton production. Environmental parameters, phytoplankton biomass, community composition, and photosynthetic parameters were monitored at two sites in the estuary during a complete annual cycle. Despite the fact that nutrient concentrations were always above growth-limiting values, Chl a concentrations were relatively low throughout the study period. High water column turbidity, due to riverine inputs, promoted a rapid attenuation of light and created a compressed profile with optimal photosynthetic conditions. Therefore, the phytoplankton community, dominated by small cells, such as diatoms and cryptophycean flagellates, displayed highly photosynthetic efficiency and low light-saturated photosynthetic rates as a photo-acclimation response to low light conditions year-round. Primary production rate was unimodal, peaking in the summer months. In such estuarine system, gross primary production could thus be predicted by an existing robust empirical model based on pigment standing crop (Chl a), surface irradiance (E ₀) and optical depth (Z _eup). Compared to other shallow estuaries, the Tagus can be classified as a low- to moderately productive estuary, being the turbidity-induced low light conditions the principal factor limiting phytoplankton growth.     

The development and persistence of alternative ecosystem states in a large,shallow lake

1. Palaeolimnological data were used to investigate drivers of the community of primary producers in Lake Mattamuskeet, North Carolina, U.S.A. This is a large, shallow lake with two basins currently dominated by phytoplankton and macrophytes. The two basins were divided in 1940 by the building of a roadway across the lake, which also corresponded with the divergence in their ecosystem state. 2. Photosynthetic pigments, organic matter and nutrients (P, N, C, S) were analysed in sediment cores from each basin to reconstruct the primary producer community over the past c. 100 years. We sought to answer two questions. First, what changes to the ecosystem resulting from the building of the roadway caused the development of different primary producer communities in the two basins? Second, why have the alternative ecosystem states persisted despite a variety of human perturbations since 1940? 3. K‐means cluster analysis and principal component analysis were applied to identify three sediment types based on photosynthetic pigment data: sediments indicating low productivity (low pigment concentrations), sediments associated with macrophytes (chlorophyll a and b) and with phytoplankton (alloxanthin and aphanizophyll). In addition, other palaeolimnological proxies measured, such as loss on ignition, total phosphorus, total organic carbon/total nitrogen and other nutrients, were different in post‐1940 sediments within the two basins. 4. These differences suggest characteristics, such as nutrient cycling, water depth and other physical changes resulting from roadway construction, combined to establish and maintain the differing communities of primary producers in the two basins. Furthermore, Fe/S dynamics and waterfowl herbivory probably contributed to the development of the two ecosystem states.     

Mixing, stratification and the fate of primary production in an oligotrophic multibasin lake system (Quebec, Canada)

Impacts of mixing and stratification on the fate of primaryproduction were studied in an oligotrophic lake by comparingthe size-distributions of phytoplankton standing stock and productionin two basins, only one of which experiences seasonal thermalstratification. In both basins, the phytoplankton was dominatedby small cells (pico- and nanoplankton). The contribution ofpicoplankton to both biomass and production remained relativelyconstant throughout the season in both basins. Seasonal variationsin the size structure of phytoplankton communities do not agreewith the paradigm of dominance by small cells during summerstratification and dominance of larger cells during spring andfall mixing events. Nutrient control of productivity throughmixing and stratification is unlikely to affect the structureof phytoplankton communities when nutrients (allochthonous)derived from the catchment basin or sediments are in short supply.In such environments, nutrients (autochthonous) are largelyderived in the lake through heterotrophic food web processessuch as grazing, excretion and decomposition. Maximum ratesof production and losses in July and August in both basins areconsistent with increased regeneration and may represent a responseof larger-sized cells to higher nutrient availability resultingfrom enhanced grazing on picoplankton. The high correlationbetween the rates of loss and of potential growth for the phytoplanktoncommunity during all sampling periods, and the relative constancyof the picoplankton biomass, leads us to propose a long-term,steady-state equilibrium in the phytoplankton community underthe control of grazing by herbivores and/or other loss processes.     

Sorption of hydrophobic organic contaminants and trace metals on phytoplankton and implications for toxicity assessment

The partitioning of trace metals and hydrophobic organic contaminants to phytoplankton determines their toxicity as well as their fate and transport in aquatic ecosystems. Accurate impact assessments, therefore, depend on a good understanding of the factors regulating the sorption of these compounds to biotic particles. The accumulation of chlorinated organic compounds in phytoplankton is generally considered as being due solely to physical sorption, described by reversible equilibrium models based on Langmuir or Freundlich isotherms. On the other hand, the uptake of trace metals is a two phase process: a fast sorption component viewed as an ionexchange or a covalent bonding process with cell surface ligands, followed by an intracellular transport phase that is dependent on cellular metabolic activity. The uptake of inorganic and hydrophobic organic pollutants and their bioaccumulation are influenced in a complex manner by duration of exposure and cell density, by environmental factors such as pH, the concentration of cations and of dissolved and colloidal organic matter, as well as by phytoplankton physiological condition. High concentrations of H⁺, Ca²⁺, and Mg²⁺ ions will reduce trace metal sorption by directly competing for uptake sites on the cell's surface, whereas the presence of dissolved organic carbon such as natural and synthetic chelators and phytoplankton exudates will reduce the bioavailability of both trace metals and hydrophobic organic contaminants. Thus, the impact of toxic contaminants on phytoplankton may be determined as much by the factors influencing uptake and partitioning as by the potency of the toxicants and interspecies differences in sensitivity. Recommendations for improving toxicity assessments are presented.     

Light-Shade Adaptation : TWO STRATEGIES IN MARINE PHYTOPLANKTON

Using chlorophyll/P700 ratios, the size and number of photosynthetic units were estimated, as a function of light-shade adaptation in two species of marine phytoplankton: Skeletonema costatum, a diatom, and Dunaliella tertiolecta, a chlorophyte. In the diatom, light-shade adaptation is characterized primarily by changes in the size and not the number of P700 units, whereas in the chlorophyte, overall changes in chlorophyll content are related to changes in the number and not the size of P700 units. A correlation between the characteristics of P700 units and photosynthetic responses was not established. Both strategies of light-shade adaptation effectively harvest and transfer light energy to reaction centers, however, the Skeletonema strategy is more effective at subsaturating intensities. The two strategies may represent an evolutionary divergence in photosynthetic adaptation to variations in light intensity.     

A mechanistic model of the adaptation of phytoplankton photosynthesis

The mechanistic model of the phytoplankton photosynthesis-light intensity relationship by Eilers and Peeters (1988.Ecol. Modelling 42, 199–215) is investigated mathematically. The model is based on the physiological idealization of transition probabilities between states of the photosynthetic factories,PSF. The model was found to have under constant light condition a globally stable unique positive equilibrium, while under periodically varying light (e.g. daily periodicity) there exists a unique globally asymptotically stable periodic solution. Based on this, the adaptation to a change of light intensity is defined as a process by which the state ofPSF converges to an equilibrium. Assuming that phytoplankton regulates its photosynthetic production rate with a certain strategy which maximizes production, two such possible strategies were examined. Both the instantaneous and the integral maximal photosynthetic production were shown to have the same result. With realistic qualitative assumptions of the shape of the dependence of the four model parameters on the light intensity to which phytoplankton is adapted, the numerical values of parameters under both constant and periodically varying conditions are determined by applying Pontryagin's maximum principle.     

Optimizing photosynthesis under fluctuating light: The role of the Arabidopsis STN7 kinase

Optimal photosynthetic performance requires that equal amounts of light are absorbed by photosystem ii (PSii) and photosystem i (PSi), which are functionally linked through the photosynthetic electron transport chain. However, photosynthetic organisms must cope with light conditions that lead to the preferential stimulation of one or the other of the photosystems. Plants react to such imbalances by mounting acclimation responses that redistribute excitation energy between photosystems and restore the photosynthetic redox poise. in the short term, this involves the so-called state transition process, which, over periods of minutes, alters the antennal crosssections of the photosystems through the reversible association of a mobile fraction of light-harvesting complex ii (LHCii) with PSi or PSii. Longer-lasting changes in light quality initiate a long-term response (LTr), occurring on a timescale of hours to days, that redresses imbalances in excitation energy by changing the relative amounts of the two photosystems. Despite the differences in their timescales of action, state transitions and LTr are both triggered by the redox state of the plastoquinone (PQ) pool, via the activation of the thylakoid kinase STN7, which appears to act as a common redox sensor and/or signal transducer for both responses. This review highlights recent findings concerning the role of STN7 in coordinating short- and long-term photosynthetic acclimation responses.Key words: state transitions, long-term acclimation, photosynthesis, STN7, Arabidopsis     

Physiological response of 10 phytoplankton species exposed to macondo oil and the dispersant,Corexit

Culture experiments were conducted on ten phytoplankton species to examine their biological and physiological responses during exposure to oil and a combination of oil and dispersant. The species tested included a range of taxa typically found in the Gulf of Mexico such as cyanobacteria, chlorophytes, and diatoms. Cultures were exposed to Macondo surrogate oil using the water accommodated fraction (WAF), and dispersed oil using a chemically enhanced WAF (CEWAF) and diluted CEWAF, to replicate conditions following the Deepwater Horizon spill in the Gulf of Mexico. A range of responses were observed, that could broadly class the algae as either “robust” or “sensitive” to oil and/or dispersant exposure. Robust algae were identified as Synechococcus elongatus, Dunaliella tertiolecta, two pennate diatoms Phaeodactylum tricornutum and Navicula sp., and Skeletonema grethae CCMP775, and were largely unaffected by any of the treatments (no changes to growth rate or time spent in lag phase relative to controls). The rest of the phytoplankton, all centric diatoms, exhibited at least some combination of reduced growth rates or increased lag time in response to oil and/or dispersant exposure. Photophysiology did not have a strong treatment effect, with significant inhibition of photosynthetic efficiency (F_v/F_m) only observed in the CEWAF, if at all. We found that the effects of oil and dispersants on phytoplankton physiology were species‐dependent, and not always detrimental. This has significant implications on how oil spills might impact phytoplankton community structure and bloom dynamics in the Gulf of Mexico, which in turn impacts higher trophic levels.     

A 3-D model of ligand transport in a deforming extracellular space

Cells communicate through shed or secreted ligands that traffic through the interstitium. Force-induced changes in interstitial geometry can initiate mechanotransduction responses through changes in local ligand concentrations. To gain insight into the temporal and spatial evolution of such mechanotransduction responses, we developed a 3-D computational model that couples geometric changes observed in the lateral intercellular space (LIS) of mechanically loaded airway epithelial cells to the diffusion-convection equations that govern ligand transport. By solving the 3-D fluid field under changing boundary geometries, and then coupling the fluid velocities to the ligand transport equations, we calculated the temporal changes in the 3-D ligand concentration field. Our results illustrate the steady-state heterogeneities in ligand distribution that arise from local variations in interstitial geometry, and demonstrate that highly localized changes in ligand concentration can be induced by mechanical loading, depending on both local deformations and ligand convection effects. The occurrence of inhomogeneities at steady state and in response to mechanical loading suggest that local variations in ligand concentration may have important effects on cell-to-cell variations in basal signaling state and localized mechanotransduction responses.     

Photosynthetic sunfleck utilization potential of understory saplings growing under elevated CO<Subscript>2</Subscript> in FACE

Few studies have evaluated elevated CO₂ responses of trees in variable light despite its prevalence in forest understories and its potential importance for sapling survival. We studied two shade-tolerant species (Acer rubrum, Cornus florida) and two shade-intolerant species (Liquidambar styraciflua, Liriodendron tulipifera) growing in the understory of a Pinus taeda plantation under ambient and ambient+200 ppm CO₂ in a free air carbon enrichment (FACE) experiment. Photosynthetic and stomatal responses to artificial changes in light intensity were measured on saplings to determine rates of induction gain under saturating light and induction loss under shade. We expected that growth in elevated CO₂ would alter photosynthetic responses to variable light in these understory saplings. The results showed that elevated CO₂ caused the expected enhancement in steady-state photosynthesis in both high and low light, but did not affect overall stomatal conductance or rates of induction gain in the four species. Induction loss after relatively short shade periods (<6 min) was slower in trees grown in elevated CO₂ than in trees grown in ambient CO₂ despite similar decreases in stomatal conductance. As a result leaves grown in elevated CO₂ that maintained induction well in shade had higher carbon gain during subsequent light flecks than was expected from steady-state light response measurements. Thus, when frequent sunflecks maintain stomatal conductance and photosynthetic induction during the day, enhancements of long-term carbon gain by elevated CO₂ could be underestimated by steady-state photosynthetic measures. With respect to species differences, both a tolerant, A. rubrum, and an intolerant species, L. tulipifera, showed rapid induction gain, but A. rubrum also lost induction rapidly (c. 12 min) in shade. These results, as well as those from independent studies in the literature, show that induction dynamics are not closely related to species shade tolerance. Therefore, it cannot be concluded that shade-tolerant species necessarily induce faster in the variable light conditions common in understories. Although our study is the first to examine dynamic photosynthetic responses to variable light in contrasting species in elevated CO₂, studies on ecologically diverse species will be required to establish whether shade-tolerant and -intolerant species show different photosynthetic responses in elevated CO₂ during sunflecks. We conclude that elevated CO₂ affects dynamic gas exchange most strongly via photosynthetic enhancement during induction as well as in the steady state. Received: 1 April 1999 / Accepted: 16 August 1999     

Photoacoustics: a novel tool for the determination of photosynthetic energy storage efficiency in phytoplankton

Photosynthetic energy storage efficiency controls the development and decline of phytoplankton biomass. All abiotic environmental factors such as light intensity; temperature, nutrient availability and pollutants will exert detectable changes in the photosynthetic energy storage efficiency of phytoplankton, and subsequently affect total biomass and composition of phytoplankton assemblages. Since this efficiency is a sensitive amplifier of ambient conditions, it thereby is an excellent reporter of water quality parameter. We demonstrate the applicability of the novel photoacoustic method in easily and directly estimating the energy storage efficiency of phytoplankton in a drinking water reservoir of different nutrient status. Electronic Supplementary Material Supplementary material is available in the online version of this article at and accessible for authorised users Handling editor: J. Padisak