Water status related photosynthesis and carbon isotope discrimination in species of the lichen genusPseudocyphellaria with green or blue-green photobionts and in photosymbiodemes

Summary Green lichens have been shown to attain positive net photosynthesis in the presence of water vapour while blue-green lichens require liquid water (Lange et al. 1986). This behaviour is confirmed not only for species with differing photobionts in the genusPseudocyphellaria but for green and blue-green photobionts in a single joined thallus (photosymbiodeme), with a single mycobiont, and also when adjacent as co-primary photobionts. The different response is therefore a property of the photobiont. The results are consistent with published photosynthesis/water content response curves. The minimum thallus water content for positive net photosynthesis appears to be much lower in green lichens (15% to 30%, related to dry weight) compared to blue-greens (85% to 100%). Since both types of lichen rehydrate to about 50% water content by water vapour uptake only green lichens will show positive net photosynthesis. It is proposed that the presence of sugar alcohols in green algae allow them to retain a liquid pool (concentrated solution) in their chloroplasts at low water potentials and even to reform it by water vapour uptake after being dried. The previously shown difference in δ¹³C values between blue-green and green lichens is also retained in a photosymbiodeme and must be photobiont determined. The wide range of δ¹³C values in lichens can be explained by a C₃ carboxylation system and the various effects of different limiting processes for photosynthetic CO₂ fixation. If carboxylation is rate limiting, there will be a strong discrimination of¹³CO₂, at high internal CO₂ partial pressure. The resulting very low δ¹³C values (-31 to-35‰) have been found only in green lichens which are able to photosynthesize at low thallus water content by equilibraiton with water vapour. When the liquid phase diffusion of CO₂ becomes more and more rate limiting and the internal CO₂ pressure decreases, the¹³C content of the photosynthates increases and less negative δ¹³C values results, as are found for blue-green lichens.     

An investigation of glycolate excretion in two species of blue-green algae

Summary The amount of ¹⁴C-glycolate excreted by Oscillatoria sp. and Anabaena flos-aquae is less than 1% of the ¹⁴C fixed by the algae during photosynthesis. Transfer of cells grown on 5% CO₂ in air to a medium of low bicarbonate concentration or treatment of the cells with isonicotinyl hydrazide (INH) during photosynthesis, caused little increase in glycolate excretion. -Hydroxysulfonates failed to stimulate massive excretion of glycolate. Although these blue-green algae excreted little glycolate, a significant proportion of the photosynthetically fixed carbon was excreted in the form of basic, neutral and acidic compounds, and such excretion was greater in 5% CO₂-grown cells than in air-grown cells.     

Blue-green Dominance in Lakes: The Role and Management Significance of pH and CO2

Field and laboratory experiments show that when nutrients (N and P) are added to a mixed population of algae, blue-greens predominate. However, if CO₂ is added also, or if pH is lowered with HCl, greens predominate. The phenomenon is reproducible, and works with most lakes. Although most successful at pH 5.5, the shift to greens can be made to happen at pH values as high as 8.5. Most blue-greens appear to be susceptible to the shift, and Scenedesmus and Chlorella are the predominant greens resulting. If pH is raised, the shift is reversed. The reason for the shift is not known. It may involve competition by the algae for CO₂, but other evidence suggests that the lowered pH stimulates cyanophage production and lysis of the blue-greens, with release of nutrients which then are used by the greens. Analysis of results of lake circulation data from the literature and from experiments suggests that the algal shifts resulting from circulation may involve the same phenomena. Understanding of these phenomena should lead to predictable use of circulation as a lake-management tool.     

Some ecological effects of artificial circulation on a small eutrophic lake with particular emphasis on phytoplankton II. Kezar lake experiment, 1969

A small eutrophic New Hampshire lake was artificially circulated (mixed) from 28 May to 15 September, 1969, inclusive, to impede the annual bloom of a noxious blue-green alga; yet Aphanizomenon flos-aquae bloomed immediately after mixing was commenced. The bloom collapsed in early July; it was succeeded by heavy growth of predominantly chlorophycean taxa. In an in vitro experiment Aphanizomenon flos-aquae did reattain bloom proportions when the influence of artificial circulation was removed. Other phytoplankton exhibited population pulses only when the dominance of cyanophycean and chlorophycean taxa were in transition. Mixing maintained uniform vertical populations of all phytoplankton. Changes in water transparency attended fluctuations in phytoplankton abundance. An isothermic condition was maintained over the test period, which increased the lake's heat budget, and most chemical nutrients were distributed isometrically in the water column. Increased concentrations were exhibited by Ca, Cl, Cu, K, Mg, SiO₂ and Zn. Sodium was not affected by mixing. Levels of Fe, Mn, Zn, phosphate and ammonia, nitrate, and organic nitrogen were influenced by phytoplankton. Mixing could not maintain orthograde profiles of dissolved O₂ and CO₂ when dense populations of phytoplankton prevailed. Variations in chlorophyll-a followed closely changes in phytoplankton abundance. Its degradation to pheo-pigments appeared to be less for a bloom of Aphanizomenon flos-aquae than during dense growth of chlorophycean taxa. Rates of photosynthesis were considerably greater when the latter algae were predominant. Extracellular release of organic carbon usually increased with depth; it amounted to 19.4 percent of the total carbon fixed in the euphotic zone.     

Blue-green algae and selection in rotifer populations

Summary Observation in natural populations of the rotifer Asplanchna girodi suggested blue-green algae may have toxic effects on rotifers. The blue-greens Anabaena flos-aquae and Lyngbya sp. were isolated from a pond in Central Florida along with three electrophoretically identified genotypes of A. girodi. When tested Anabaena had marked effect on the reproductive rate of A. girodi and the effect was different among genotypes. Likewise, Lyngbya also depressed rotifer reproduction, but required an order of magnitude higher concentration to produce the same effects as Anabaena. The toxicity of these algae was corroborated by mouse bioassays. Results clearly show these blue-greens have the capacity to differentially reduce reproductive rates of rotifer genotypes. Because of this capability blue-greens could play an important role in the ecology of natural selection in rotifer populations.     

Photosynthetic response of two shrubs to rainfall pulses in desert regions of northwestern China

Pulses of rainfall are particularly pivotal in controlling plant physiological processes in ecosystems controlled by limited water, and the response of desert plants to rainfall is a key to understanding the responses of desert ecosystems to global climatic change. We used a portable photosynthesis system to measure the responses of the diurnal course of photosynthesis, light-response curves, and CO₂-response curves of two desert shrubs (Nitraria sphaerocarpa Maxim. and Calligonum mongolicum Turcz) to a rainfall pulse in a desert-oasis ecotone in northwestern China. The photosynthetic parameters, light- and CO₂-response curves differed significantly before and after the rainfall pulse. Their maximum net photosynthetic rate (P _N) values were 23.27 and 32.92 μmol(CO₂) m⁻² s⁻¹ for N. sphaerocarpa and C. mongolicum, respectively, with corresponding maximum stomatal conductance (g _s) values of 0.47 and 0.39 mol(H₂O) m⁻² s⁻¹. The P _N of N. sphaerocarpa after the rainfall was 1.65 to 1.75 times the value before rainfall, whereas those of C. mongolicum increased to approximately 2 times the prerainfall value, demonstrating the importance of the desert plants response by improving their assimilation rate to precipitation patterns under a future climate.     

pH regulation in apoplastic and cytoplasmic cell compartments of leaves

 The regulation of pH in the apoplast, cytosol and chloroplasts of intact leaves was studied by means of fluorescent pH indicators and as a response of photosynthesis to acid stress. The apoplastic pH increased under anaerobiosis. Aeration reversed this effect. Apoplastic responses to CO₂, HCl or NH₃ differed considerably. Whereas HCl and ammonia caused rapid acidification or alkalinization, the return to initial pH values was slow after cessation of fumigation. Addition of CO₂ either did not produce the acidification expected on the basis of known apoplastic buffering or even caused some alkalinization. Removal of CO₂ shifted the apoplastic pH into the alkaline range before the pH returned to initial steady-state levels. In the presence of vanadate, the alkaline shift was absent and the apoplastic pH returned slowly to the initial level when CO₂ was removed from the atmosphere. In contrast to the response of the apoplast, anaerobiosis acidified the cytosol or, in some species, had little effect on its pH. Acidification was rapidly reversed upon re-admission of oxygen. The CO₂-dependent pH changes were very fast in the cytosol. Considerable alkalinization was observed after removal of CO₂ under aerobic, but not under anaerobic conditions. Rates of the re-entry of protons into the cytosol during recovery from CO₂ stress increased in the presence of oxygen with the length of previous exposure to high CO₂. Effective pH regulation in the chloroplasts was indicated by the recovery of photosynthesis after the transient inhibition of photosynthetic electron flow when CO₂ was increased from 0.038% to 16% in air. As photosynthesis became inhibited under high CO₂, reduction of the electron transport chain increased transiently. The time required for recovery of photosynthesis from inhibition during persistent CO₂ stress was similar to the time required for establishing steady-state pH values in the cytosol under acid stress. The high capacity of leaf cells for the rapid re-attainment of pH homeostasis in the apoplast and the cytoplasm under acid or alkaline stress suggested the rapid activation or deactivation of membrane-localised proton-transporting enzymes and corresponding ion channel regulation for co-transport of anions or counter-transport of cations together with proton fluxes. Acidification of the cytoplasm appeared to activate energy-dependent proton export primarily into the vacuoles whereas apoplastic alkalinization resulted in the pumping of protons into the apoplast. Proton export rates from the cytosol into the apoplast after anaerobiosis were about 100 nmol (m² leaf area)⁻¹ s⁻¹ or less. Proton export under acid stress into the vacuole was about 1200 nmol m⁻² s⁻¹. The kinetics of pH responses to the addition or withdrawal of CO₂ indicated the presence of carbonic anhydrase in the cytosol, but not in the apoplast. Received: 19 July 1999 / Accepted: 29 December 1999     

Polymixis of a shallow lake (Groβer Müggelsee,Berlin) and its influence on seasonal phytoplankton dynamics

The polymictic properties of Lake Müggelsee, a eutrophic shallow lake in Berlin, are described by the water column stability (N ²) and gradients in saturation of oxygen at the deepest site of the lake (7.5 m). Mixing and stratification changed irregularly up to 7 times during the vegetation season (April to September), as was indicated by all of the stratification parameters. Thermally stable conditions generally lasted 1–2 weeks. A maximum of 5 weeks stratification was observed in 1982.In order to investigate the response of algal development, the internal rates of change of the dominant algal species in the lake during the vegetation period were estimated from weekly measurements of phytoplankton biomass from 1980 to 1990. The necessity taking a mixed sample in a shallow lake is discussed. The polymictic properties favoured the development of specific blue-green algal species; there dominance was also favoured by the trophic conditions. Among the dominant blue-greens the growth of Limnothrix redekei was independent of polymixis whereas stratification supported the starting conditions for the summer blue-greens Aphanizomenon flos-aquae and Planktothrix agardhii. After these algae reached a distinct level of biomass, they grew under mixing as well as under stratified conditions.For the development of solitary centric diatoms during summer regulation by growth restriction through nutrient limitation, esp. dissolved silicon was more important. However, Melosira sp. developed well under stratified conditions but collapsed due to increased sinking losses when the water column became too stable.An attempt is made to apply Reynolds' possibility matrix of the most likely phytoplankton assemblages as a function of nutrients and mixing in the shallow Lake Müggelsee.     

Field and Laboratory Studies on Nile Phytoplankton in Egypt IV. Phytoplankton of Aswan High Dam Lake (Lake Nasser)

Seasonal, local and monthly vertical variations in chlorophyll a concentration and in qualitative and quantitative counts in Aswan High Dam Lake (AHDL) phytoplankton were followed from March 1982 to February 1984. The chlorophyll a values correlated more or less with the total phytoplankton counts. Total algal counts exhibited considerable local variations in the different seasons and tended to decrease in the most southern part of AHDL, especially during the flood period. The temporal course of algal development showed an almost inverse trend to that of dissolved nitrate-nitrogen. The possibility of nitrogen limitation of algal growth was verified. Diatoms and blue-green algae alternated in the dominance of the phytoplankton community. The genera Cyclotella and Anabaenopsis respectively were the most predominant genera among the diatoms and blue-greens and consequently in the phytoplankton community.     

Do all leaf photosynthesis parameters of rice acclimate to elevated CO2, elevated temperature,and their combination,in FACE environments?

Leaf photosynthesis of crops acclimates to elevated CO₂ and temperature, but studies quantifying responses of leaf photosynthetic parameters to combined CO₂ and temperature increases under field conditions are scarce. We measured leaf photosynthesis of rice cultivars Changyou 5 and Nanjing 9108 grown in two free‐air CO₂ enrichment (FACE) systems, respectively, installed in paddy fields. Each FACE system had four combinations of two levels of CO₂ (ambient and enriched) and two levels of canopy temperature (no warming and warmed by 1.0–2.0°C). Parameters of the C₃ photosynthesis model of Farquhar, von Caemmerer and Berry (the FvCB model), and of a stomatal conductance (g_s) model were estimated for the four conditions. Most photosynthetic parameters acclimated to elevated CO₂, elevated temperature, and their combination. The combination of elevated CO₂ and temperature changed the functional relationships between biochemical parameters and leaf nitrogen content for Changyou 5. The g_s model significantly underestimated g_s under the combination of elevated CO₂ and temperature by 19% for Changyou 5 and by 10% for Nanjing 9108 if no acclimation was assumed. However, our further analysis applying the coupled g_s–FvCB model to an independent, previously published FACE experiment showed that including such an acclimation response of g_s hardly improved prediction of leaf photosynthesis under the four combinations of CO₂ and temperature. Therefore, the typical procedure that crop models using the FvCB and g_s models are parameterized from plants grown under current ambient conditions may not result in critical errors in projecting productivity of paddy rice under future global change.     

Phytoplankton ecology in an unusually stable environment (Montezuma Well,Arizona, U.S.A.)

Relatively minor annual amplitudes of change in certain major nutrients, and especially pH and water temperature were measured in the spring-fed system of Montezuma Well, Arizona during a four year study. phytoplankton diversity was low but for the most part, composition was spatially and temporally constant; total seasonal phytoplankton density was significantly correlated with regional incident light. Phytoplankton species composition changed briefly during and for a short period following the summer monsoon. Ultraplankton (<5 µm diam.) numerically comprised nearly 80% of the phytoplankton community throughout most of the year. The limited residence time of water in the Well may have provided a competitive advantage for cells with high surface area:volume ratios and correspondingly rapid division rates. Nannochloris bacillaris Naum. and Coccomyxa minor Skuja were perennial dominants. Diatom populations did not increase with annual increases in vernal solar radiation. Low pH, high dissolved CO₂, and limited residence time for metabolic inhibitors are considered to be largely responsible for the reduced blue-green populations in the Well. The only flagellated photosynthetic group present in Montezuma Well was the Cryptophyta. Desmid populations were minimal, even though pH was consistently below circumneutral (6.5) and free CO₂ concentrations high. The role of grazing by an amphipod, Hyalella montezuma, on annual phytoplankton abundance is examined.     

Disturbance events affecting phytoplankton biomass,composition and species diversity in a shallow,eutrophic, temperate lake

The seasonal changes in phytoplankton biomass and species diversity in a shallow, eutrophic Danish lake are described and related to different disturbance events acting on the phytoplankton community.Both the spring diatom maximum and the summer bloom of the filamentous blue-green alga, Aphanizomenon flos-aquae (L.) Ralfs, coincided with low values of phytoplankton species diversity and equitability. Diatom collapse was mainly due to internal modifications as nutrient depletion (Si, P) caused by rapid growth of phytoplankton, and increased grazing activity from zooplankton. A large population of Daphnia longispina O.F. Müller in June effectively removed smaller algal competitors, thus favouring the development of a huge summer bloom (140 mm³ l^–1) of Aphanizomenon flos-aquae. Heavy rainfall and storms in late July increased the loss of Apahnizomenon by out-flow and disturbed the stratification of the lake. These events caused a marked decline in phytoplankton biomass but had no effect on species diversity. A second storm period in late August circulated the lake completely and was followed by a rapid increase in phytoplankton diversity, and a change in the phytoplankton community structure from dominance of large, slow-growing K-selected species (Aphanizomenon) to small, fast-growing r-selected species (cryptomonads).     

基于FvCB模型的叶片光合生理对环境因子的响应研究进展

为提高叶片光合速率并更好地理解叶片光合生理对环境因子变化的响应机制,FvCB模型(C_3植物光合生化模型)常用于分析不同环境条件下CO_2响应曲线并预测叶片活体内光合系统的内在变化状况。系统介绍了FvCB模型的建立、发展过程和拟合方法等基本理论,综述了该模型在叶片光合生理对光、CO_2、水、温度和N营养等环境因子变化的响应机制中的应用研究。为进一步完善FvCB模型并更好地理解叶片活体内光合系统对环境因子变化的响应机制,未来拟加强以下研究:1)羧化速率与光合电子传递速率之间的联系;2)叶肉导度的具体组分及其对FvCB模型参数估计的影响;3)叶片气孔导度和叶肉导度对环境因子变化的调控机制。     

The acclimation of leaf photosynthesis of wheat and rice to seasonal temperature changes in T‐FACE environments

Crops show considerable capacity to adjust their photosynthetic characteristics to seasonal changes in temperature. However, how photosynthesis acclimates to changes in seasonal temperature under future climate conditions has not been revealed. We measured leaf photosynthesis (A_n) of wheat (Triticum aestivum L.) and rice (Oryza sativa L.) grown under four combinations of two levels of CO₂ (ambient and enriched up to 500 µmol/mol) and two levels of canopy temperature (ambient and increased by 1.5–2.0°C) in temperature by free‐air CO₂ enrichment (T‐FACE) systems. Parameters of a biochemical C₃‐photosynthesis model and of a stomatal conductance (g_s) model were estimated for the four conditions and for several crop stages. Some biochemical parameters related to electron transport and most g_s parameters showed acclimation to seasonal growth temperature in both crops. The acclimation response did not differ much between wheat and rice, nor among the four treatments of the T‐FACE systems, when the difference in the seasonal growth temperature was accounted for. The relationships between biochemical parameters and leaf nitrogen content were consistent across leaf ranks, developmental stages, and treatment conditions. The acclimation had a strong impact on g_s model parameters: when parameter values of a particular stage were used, the model failed to correctly estimate g_s values of other stages. Further analysis using the coupled g_s–biochemical photosynthesis model showed that ignoring the acclimation effect did not result in critical errors in estimating leaf photosynthesis under future climate, as long as parameter values were measured or derived from data obtained before flowering.     

Photosynthetic and stomatal conductance responses to acid mist of red spruce seedlings

Abstract Two-year-old seedlings of Picea rubens, growing in open-top chambers in Scotland were treated twice weekly from July 1987 to December 1987, with mist containing ammonium sulphate and nitric acid at a pH of either 2.5 or 5.0. The response of photosynthesis and stomatal conductance to light flux density and carbon dioxide concentration were measured in March 1989. Leaf chlorophyll a and b contents were also measured. Acid mist (pH 2.5) resulted in several significant changes. First, both the rate of light saturated photosynthesis (A_max) and CO₂- saturated rate of photosynthesis (J) were substantially increased, when expressed per unit leaf area. Second, the apparent quantum yield and chlorophylls a and b content increased. Third, as a consequence of the greater chlorophyll content of the leaves treated with acid mist, the rate of A_max, and J, expressed per unit chlorophyll, was substantially reduced in pH 2.5 treated branches. Stomatal conductance was enhanced at all but the highest light flux densities, and was independent of the CO₂ concentration, remaining high for all values of CO₂ concentration used. These results show that acid mist caused a number of responses in the gas exchange and photosynthetic properties of red spruce.     

Eutrophication of the salt valley reservoirs, 1968–73 I. The effects of eutrophication on standing crop and composition of phytoplankton

A study was carried out over a period of six years to determine the effects of eutrophication upon standing crop and composition of the phytoplankton in four recently constructed flood control reservoirs in Nebraska. Water samples collected weekly during June, July, and August from 1968–73 were analyzed for chlorophyll a, phytoplankton composition, and phytoplankton abundance. Total volume of phytoplankton was calculated from appropriate dimensions and formulae. Inorganic turbidity in one reservoir was an important factor regulating the size and composition of the phytoplankton standing crop. In that reservoir diatoms were the most important component of the phytoplankton community during those years in which inorganic turbidity was greatest. When inorganic turbidity declined, blue-green algae became dominant.In the clear-water reservoirs chlorophyll a, phytoplankton number, and phytoplankton volume were significantly correlated with reservoir age, with the oldest containing chlorophyll a concentrations up to 247 mg/m³ and mean phytoplankton volumes up to 329 mm³/l, values sufficient to place it in the hypereutrophic category. The reservoirs had 2–6 times more chlorophyll a present at the end of the study than at the beginning, with the increase being greatest in the newest reservoir. Phytoplankton volume was significantly correlated with chlorophyll a in all the reservoirs.Blue-green algae quickly became established as community dominants in the reservoirs, making up over 80 percent of the phytoplankton volume in the newest reservoir by the second year of its existence. In the other clear-water reservoirs, blue-greens usually constituted over 95 percent of the total phytoplankton volume in summer. three genera, Microcystis, Aphanizomenon, and Anabaena, were responsible for virtually the entire standing crop of blue-greens in all the reservoirs.     

Diel, episodic and seasonal changes in pH and concentrations of inorganic carbon in a productive lake

• 1 Two pH electrodes and a thermistor were used to record conditions in the surface of Esthwaite Water every 15 min over a 12-month period. Combined with approximately weekly measurements of alkalinity they allowed inorganic carbon speciation to be calculated.
• 2 Large changes in pH from 7.1 to nearly 10.3, and hence in concentrations of inorganic carbon species, were measured over a year. Carbon speciation and pH varied on a diel, episodic and seasonal basis. Diel variation of up to pH 1.8 was recorded, although typical daily variation was between 0.03 and 1.06 (5 and 95 percentiles). Daily change in concentration of inorganic carbon varied between 4 and 63 mmol m^-3 (5 and 95 percentiles).
• 3 During lake stratification, episodes of high pH, typically of 1–2 weeks' duration were interspersed with episodes of lower pH. These changes appeared to relate to the weather: e.g. low wind velocity, high pressure, low rainfall and high sunshine hours correlated with periods of high pH.
• 4 Seasonal progression of carbon depletion generally followed stratification and the development of high phytoplankton biomass. When the lake was isothermal, the phytoplankton biomass caused relatively small amounts of carbon depletion.
• 5 During autumn, winter and spring, the lake had concentrations of CO₂* (free CO₂) up to 0.12 mol m^-3 which is nearly seven times the calculated atmospheric equilibrium concentration so the lake will accordingly be losing carbon to the atmosphere. In contrast, during periods of elevated pH the concentration of CO₂* was reduced close to zero and the lake will take up atmospheric CO₂. The rates of transfer between water and the atmosphere were estimated using a chemical equilibrium model with three boundary layer thicknesses. The calculations show that over a year the lake loses CO₂ to the atmosphere with the current mean atmospheric level of 360 μmol mol^-1, at between 0.28 and 2.80 mol m^-2 yr^-1. During elevated pH, rates of CO₂-influx increased up to nearly tenfold as a result of chemical-enhancement by parallel flux of HCO^-₃. Input of CO₂* to the lake from the catchment is suggested to be the main source of the carbon lost to the atmosphere.
• 6 The turnover time for CO₂ between the air and water was calculated to be 1 year for the gross influx and 3.3 years for the net flux. These values are less than the average water residence time of 0.25 years, which indicates that over a year inflow from streams is a more important source of inorganic carbon than the atmosphere.
• 7 Influx of CO₂ from the atmosphere was calculated to be roughly equivalent to between 1 and 4% of the rates of production in the water during mid-summer indicating that this source of inorganic carbon is not a major one in this lake.
• 8 Influx of CO₂ from the hypolimnion was estimated on one occasion to be 6.9 10^-9 mol m^-2 s^-1 using transfer values based on mass eddy-diffusion. These rates are equivalent to 23% of the rate of influx of CO₂ from the atmosphere on this occasion which suggests that the hypolimnion is probably a small source of inorganic carbon to the epilimnion. The exception appears to be during windy episodes when pH is depressed. Calculations based on depth-profiles of CO₂* and HCO^-₃ suggest that the measured changes in pH can be accounted for by entrainment of hypolimnetic water into the epilimnion.
• 9 The solubility product for calcite was exceeded by up to about sixfold which, although insufficient to allow homogeneous precipitation, may have allowed heterogeneous precipitation around algal particles.

     

Nitrogen source and pCO2 synergistically affect carbon allocation,growth and morphology of the coccolithophore Emiliania huxleyi: potential implications of ocean acidification for the carbon cycle

Coccolithophores are unicellular phytoplankton that produce calcium carbonate coccoliths as an exoskeleton. Emiliania huxleyi, the most abundant coccolithophore in the world's ocean, plays a major role in the global carbon cycle by regulating the exchange of CO₂ across the ocean‐atmosphere interface through photosynthesis and calcium carbonate precipitation. As CO₂ concentration is rising in the atmosphere, the ocean is acidifying and ammonium (NH₄⁺) concentration of future ocean water is expected to rise. The latter is attributed to increasing anthropogenic nitrogen (N) deposition, increasing rates of cyanobacterial N₂ fixation due to warmer and more stratified oceans, and decreased rates of nitrification due to ocean acidification. Thus, future global climate change will cause oceanic phytoplankton to experience changes in multiple environmental parameters including CO₂, pH, temperature and nitrogen source. This study reports on the combined effect of elevated pCO₂ and increased NH₄⁺ to nitrate (NO₃^?) ratio (NH₄⁺/NO₃^?) on E. huxleyi, maintained in continuous cultures for more than 200 generations under two pCO₂ levels and two different N sources. Herein, we show that NH₄⁺ assimilation under N‐replete conditions depresses calcification at both low and high pCO₂, alters coccolith morphology, and increases primary production. We observed that N source and pCO₂ synergistically drive growth rates, cell size, and the ratio of inorganic to organic carbon. These responses to N source suggest that, compared to increasing CO₂ alone, a greater disruption of the organic carbon pump could be expected in response to the combined effect of increased NH₄⁺/NO₃^? ratio and CO₂ level in the future acidified ocean. Additional experiments conducted under lower nutrient conditions are needed prior to extrapolating our findings to the global oceans. Nonetheless, our results emphasize the need to assess combined effects of multiple environmental parameters on phytoplankton biology to develop accurate predictions of phytoplankton responses to ocean acidification.     

Decreased photosynthesis and growth with reduced respiration in the model diatom Phaeodactylum tricornutum grown under elevated CO2 over 1800 generations

Studies on the long‐term responses of marine phytoplankton to ongoing ocean acidification (OA) are appearing rapidly in the literature. However, only a few of these have investigated diatoms, which is disproportionate to their contribution to global primary production. Here we show that a population of the model diatom Phaeodactylum tricornutum, after growing under elevated CO₂ (1000 μatm, HCL, pH_T: 7.70) for 1860 generations, showed significant differences in photosynthesis and growth from a population maintained in ambient CO₂ and then transferred to elevated CO₂ for 20 generations (HC). The HCL population had lower mitochondrial respiration, than did the control population maintained in ambient CO₂ (400 μatm, LCL, pH_T: 8.02) for 1860 generations. Although the cells had higher respiratory carbon loss within 20 generations under the elevated CO₂, being consistent to previous findings, they downregulated their respiration to sustain their growth in longer duration under the OA condition. Responses of phytoplankton to OA may depend on the timescale for which they are exposed due to fluctuations in physiological traits over time. This study provides the first evidence that populations of the model species, P. tricornutum, differ phenotypically from each other after having been grown for differing spans of time under OA conditions, suggesting that long‐term changes should be measured to understand responses of primary producers to OA, especially in waters with diatom‐dominated phytoplankton assemblages.     

Photosynthesis limitations in three fern species

Maximum photosynthesis rates in ferns are generally lower than those of seed plants, but little is known about the limiting factors, which are crucial to understand the evolution of photosynthesis in land plants. To address this issue, a gas exchange/chlorophyll fluorescence analysis was performed in three fern species spanning high phylogenetic range within Polypodiopsida (Osmunda regalis, Blechnum gibbum and Nephrolepis exaltata) to determine their maximum net photosynthesis (A_N), stomatal (g_s) and mesophyll (g_m) conductances to CO₂, and the maximum velocity of carboxylation (V_c,max). The in vitro Rubisco specificity factor (S_C/O) was also determined. All three species had values for S_C/O similar to those typical of seed plants, but values of A_N, g_s, g_m and V_c,max were within the lowest range of those observed in seed plants. In addition, g_s was unresponsive to light and CO₂, as already described in other fern species. On the contrary, g_m varied with changes CO₂. A quantitative photosynthesis limitation analysis suggested that early land plants (ferns) presented not only stomatal limitations—which were less adjustable to the environment—but also restricted g_m and V_c,max, resulting in limited maximum photosynthesis rates.