The maximum growth rate hypothesis is correct for eukaryotic photosynthetic organisms, but not cyanobacteria.

The (maximum) growth rate (µmax ) hypothesis predicts that cellular and tissue phosphorus (P) concentrations should increase with increasing growth rate, and RNA should also increase as most of the P is required to make ribosomes. Using published data, we show that though there is a strong positive relationship between the µmax of all photosynthetic organisms and their P content (% dry weight), leading to a relatively constant P productivity, the relationship with RNA content is more complex. In eukaryotes there is a strong positive relationship between µmax and RNA content expressed as % dry weight, and RNA constitutes a relatively constant 25% of total P. In prokaryotes the rRNA operon copy number is the important determinant of the amount of RNA present in the cell. The amount of phospholipid expressed as % dry weight increases with increasing µmax in microalgae. The relative proportions of each of the five major P-containing constituents is remarkably constant, except that the proportion of RNA is greater and phospholipids smaller in prokaryotic than eukaryotic photosynthetic organisms. The effect of temperature differences between studies was minor. The evidence for and against P-containing constituents other than RNA being involved with ribosome synthesis and functioning is discussed.

Effect of carbon limitation on photosynthetic electron transport in Nannochloropsis oculata.

This study describes the impacts of inorganic carbon limitation on the photosynthetic efficiency and operation of photosynthetic electron transport pathways in the biofuel-candidate microalga Nannochloropsis oculata. Using a combination of highly-controlled cultivation setup (photobioreactor), variable chlorophyll a fluorescence and transient spectroscopy methods (electrochromic shift (ECS) and P700 redox kinetics), we showed that net photosynthesis and effective quantum yield of Photosystem II (PSII) decreased in N. oculata under carbon limitation. This was accompanied by a transient increase in total proton motive force and energy-dependent non-photochemical quenching as well as slightly elevated respiration. On the other hand, under carbon limitation the rapid increase in proton motive force (PMF, estimated from the total ECS signal) was also accompanied by reduced conductivity of ATP synthase to protons (estimated from the rate of ECS decay in dark after actinic illumination). This indicates that the slow operation of ATP synthase results in the transient build-up of PMF, which leads to the activation of fast energy dissipation mechanisms such as energy-dependent non-photochemical quenching. N. oculata also increased content of lipids under carbon limitation, which compensated for reduced NAPDH consumption during decreased CO2 fixation. The integrated knowledge of the underlying energetic regulation of photosynthetic processes attained with a combination of biophysical methods may be used to identify photo-physiological signatures of the onset of carbon limitation in microalgal cultivation systems, as well as to potentially identify microalgal strains that can better acclimate to carbon limitation.

Chloride: essential micronutrient and multifunctional beneficial ion.

Cl− is an essential micronutrient for oxygenic photolithotrophs. About half of global primary productivity is carried out by oxygenic photolithotrophs exposed to saline waters with Cl− concentrations orders of magnitude higher than that needed to satisfy the micronutrient requirement. The other half of primary productivity involves terrestrial and freshwater glycophytes sometimes in environments containing significantly more Cl− than is needed for the micronutrient requirement, but less than the toxic Cl­ concentration for glycophytes. Intracellular Cl− acts in regulation of cell turgor and volume, including that of stomatal and pulvinar nastic movements, is a major ion in streptophyte and ulvophycean action potentials, and is involved in ion currents flowing around apices of pollen tubes and Acetabularia cells. More work is needed on the essentiality of Cl− in these processes, as well as the recent finding that Cl− at 1­5 mol m−3 increases water use efficiency of growth and leaf area in Nicotiana tabacum.

Energy cost of intracellular metal and metalloid detoxification in wild-type eukaryotic phytoplankton.

Microalgae use various cellular mechanisms to detoxify both non-essential and excess essential metals or metalloids. There exists however, a threshold in intracellular metal(loid) concentrations beyond which detoxification mechanisms are no longer effective and inhibition of cell division inevitably occurs. It is therefore important to determine whether the availability of energy in the cell could constrain metal(loid) detoxification capacity and to better define the thresholds beyond which a metal(loid) becomes toxic. To do this we performed the first extensive bioenergetics analysis of intracellular metal(loid) detoxification mechanisms (e.g., metal-binding peptides, polyphosphate granules, metal efflux, metal and metalloid reduction, metalloid methylation, enzymatic and non-enzymatic antioxidants) in wild-type eukaryotic phytoplankton based on the biochemical mechanisms of each detoxification strategy and on experimental measurements of detoxifying biomolecules in the literature. The results show that at the onset of metal(loid) toxicity to growth, all the detoxification strategies considered required only a small fraction of the total cellular energy available for growth indicating that intracellular detoxification ability in wild-type eukaryotic phytoplankton species is not constrained by the availability of cellular energy. The present study brings new insights into metal(loid) toxicity mechanisms and detoxification strategies in wild-type eukaryotic phytoplankton.

Gas Transfer Controls Carbon Limitation During Biomass Production by Marine Microalgae.

This study presents the first in-depth analysis of CO2 limitation on the biomass productivity of the biofuel candidate marine microalga Nannochloropsis oculata. Net photosynthesis decreased by 60% from 125 to 50 µmol O2 L(-1)h(-1) over a 12 h light cycle as a direct result of carbon limitation. Continuous dissolved O2 and pH measurements were used to develop a detailed diurnal mechanism for the interaction between photosynthesis, gas exchange and carbonate chemistry in the photo-bioreactor. Gas exchange determined the degree of carbon limitation experienced by the algae. Carbon limitation was confirmed by delivering more CO2 , which increased net photosynthesis back to its steady-state maximum. This study highlights the importance of maintaining replete carbon concentrations in photo-bioreactors and other culturing facilities, either by constant pH operation or preferably by designing a feedback loop based on the dissolved O2 concentration.

Iron acquisition and allocation in stramenopile algae.

The essential element iron has a low biological availability in the surface ocean where photosynthetic organisms live. Recent advances in our understanding of iron acquisition mechanisms in brown algae and diatoms (stramenopile algae) show the importance of the reduction of ferric to ferrous iron prior to, or during, transport in the uptake process. The uses of iron in photosynthetic stramenopiles resembles that in other oxygenic organisms, although (with the exception of the diatom Thalassiosira oceanica from an iron-deficient part of the ocean) they lack plastocyanin, instead using cytochrome c 6, This same diatom further economizes genotypically on the use of iron in photosynthesis by decreasing the expression of photosystem I, cytochrome c 6, and the cytochrome b 6 f complex per cell and per photosystem II relative to the coastal Thalassiosira pseudonana; similar changes occur phenotypically in response to iron deficiency in other diatoms such as Phaeodactylum tricornutum. In some diatoms grown under iron-limiting conditions, essentially all of the iron in the cells can be accounted for by the iron occurring in catalytic proteins. However, stramenopiles can store iron. Genomic studies show that pennate, but not centric, diatoms have the iron storage protein ferritin. While Mössbauer and X-ray analysis of (57)Fe-labelled Ectocarpus siliculosus shows iron in an amorphous mineral phase resembling the core of ferritin, the genome shows no protein with significant sequence similarity to ferritin.

Evolution of tree nutrition.

Using a broad definition of trees, the evolutionary origins of trees in a nutritional context is considered using data from the fossil record and molecular phylogeny. Trees are first known from the Late Devonian about 380 million years ago, originated polyphyletically at the pteridophyte grade of organization; the earliest gymnosperms were trees, and trees are polyphyletic in the angiosperms. Nutrient transporters, assimilatory pathways, homoiohydry (cuticle, intercellular gas spaces, stomata, endohydric water transport systems including xylem and phloem-like tissue) and arbuscular mycorrhizas preceded the origin of trees. Nutritional innovations that began uniquely in trees were the seed habit and, certainly (but not necessarily uniquely) in trees, ectomycorrhizas, cyanobacterial, actinorhizal and rhizobial (Parasponia, some legumes) diazotrophic symbioses and cluster roots.

Inorganic carbon acquisition by eukaryotic algae: four current questions.

The phylogenetically and morphologically diverse eukaryotic algae are typically oxygenic photolithotrophs. They have a diversity of incompletely understood mechanisms of inorganic carbon acquisition: this article reviews four areas where investigations continue. The first topic is diffusive CO(2) entry. Most eukaryotic algae, like all cyanobacteria, have inorganic carbon concentrating mechanisms (CCMs). The ancestral condition was presumably the absence of a CCM, i.e. diffusive CO(2) entry, as found in a small minority of eukaryotic algae today; however, it is likely that, as is found in several cases, this condition is due to a loss of a CCM. There are a number of algae which are in various respects intermediate between diffusive CO(2) entry and occurrence of a CCM: further study is needed on this aspect. A second topic is the nature of cyanelles and their role in inorganic carbon assimilation. The cyanelles (plastids) of the euglyphid amoeba Paulinella have been acquired relatively recently by endosymbiosis with genetic integration of an α-cyanobacterium with a Form 1A Rubisco. The α-carboxysomes in the cyanelles are presumably involved in a CCM, but further investigation is needed.Also called cyanelles are the plastids of glaucocystophycean algae, but is it now clear that these were derived from the ß-cyanobacterial ancestor of all plastids other than that of Paulinella. The resemblances of the central body of the cyanelles of glaucocystophycean algae to carboxysomes may not reflect derivation from cyanobacterial ß-carboxysomes; although it is clear that these algae have CCMs but these are now well characterized. The other two topics concern CCMs in other eukaryotic algae; these CCMs arose polyphyletically and independently of the cyanobacterial CCMs. It is generally believed that eukaryotic algal, like cyanobacterial, CCMs are based on active transport of an inorganic carbon species and/or protons, and they have C(3) biochemistry. This is the case for the organism considered as the third topic, i.e. Chlamydomonas reinhardtii, the eukaryotic alga with the best understood CCM. This CCM involves HCO(3)(-) conversion to CO(2) in the thylakoid lumen so the external inorganic carbon must cross four membranes in series with a final CO(2) effux from the thylakoid. More remains to be investigated about this CCM. The final topic is that of the occurrence of C(4)-like metabolism in the CCMs of marine diatoms. Different conclusions have been reached depending on the organism investigated and the techniques used, and several aspects require further study.

The evolution of inorganic carbon concentrating mechanisms in photosynthesis.

Inorganic carbon concentrating mechanisms (CCMs) catalyse the accumulation of CO(2) around rubisco in all cyanobacteria, most algae and aquatic plants and in C(4) and crassulacean acid metabolism (CAM) vascular plants. CCMs are polyphyletic (more than one evolutionary origin) and involve active transport of HCO(3)(-), CO(2) and/or H(+), or an energized biochemical mechanism as in C(4) and CAM plants. While the CCM in almost all C(4) plants and many CAM plants is constitutive, many CCMs show acclimatory responses to variations in the supply of not only CO(2) but also photosynthetically active radiation, nitrogen, phosphorus and iron. The evolution of CCMs is generally considered in the context of decreased CO(2) availability, with only a secondary role for increasing O(2). However, the earliest CCMs may have evolved in oxygenic cyanobacteria before the atmosphere became oxygenated in stromatolites with diffusion barriers around the cells related to UV screening. This would decrease CO(2) availability to cells and increase the O(2) concentration within them, inhibiting rubisco and generating reactive oxygen species, including O(3).

Critical issues in trace gas biogeochemistry and global change.

The atmospheric composition of trace gases and aerosols is determined by the emission of compounds from the marine and terrestrial biospheres, anthropogenic sources and their chemistry and deposition processes. Biogenic emissions depend upon physiological processes and climate, and the atmospheric chemistry is governed by climate and feedbacks involving greenhouse gases themselves. Understanding and predicting the biogeochemistry of trace gases in past, present and future climates therefore demands an interdisciplinary approach integrating across physiology, atmospheric chemistry, physics and meteorology. Here, we highlight critical issues raised by recent findings in all of these key areas to provide a framework for better understanding the past and possible future evolution of the atmosphere. Incorporating recent experimental and observational findings, especially the influence of CO2 on trace gas emissions from marine algae and terrestrial plants, into earth system models remains a major research priority. As we move towards this goal, archives of the concentration and isotopes of N2O and CH4 from polar ice cores extending back over 650,000 years will provide a valuable benchmark for evaluating such models. In the Pre-Quaternary, synthesis of theoretical modelling with geochemical and palaeontological evidence is also uncovering the roles played by trace gases in episodes of abrupt climatic warming and ozone depletion. Finally, observations and palaeorecords across a range of timescales allow assessment of the Earth's climate sensitivity, a metric influencing our ability to decide what constitutes 'dangerous' climate change.

A large population of small chloroplasts in tobacco leaf cells allows more effective chloroplast movement than a few enlarged chloroplasts.

We generated transgenic tobacco (Nicotiana tabacum cv Xanthi) plants that contained only one to three enlarged chloroplasts per leaf mesophyll cell by introducing NtFtsZ1-2, a cDNA for plastid division. These plants were used to investigate the advantages of having a large population of small chloroplasts rather than a few enlarged chloroplasts in a leaf mesophyll cell. Despite the similarities in photosynthetic components and ultrastructure of photosynthetic machinery between wild-type and transgenic plants, the overall growth of transgenic plants under low- and high-light conditions was retarded. In wild-type plants, the chloroplasts moved toward the face position under low light and toward the profile position under high-light conditions. However, chloroplast rearrangement in transgenic plants in response to light conditions was not evident. In addition, transgenic plant leaves showed greatly diminished changes in leaf transmittance values under both light conditions, indicating that chloroplast rearrangement was severely retarded. Therefore, under low-light conditions the incomplete face position of the enlarged chloroplasts results in decreased absorbance of light energy. This, in turn, reduces plant growth. Under high-light conditions, the amount of absorbed light exceeds the photosynthetic utilization capacity due to the incomplete profile position of the enlarged chloroplasts, resulting in photodamage to the photosynthetic machinery, and decreased growth. The presence of a large number of small and/or rapidly moving chloroplasts in the cells of higher land plants permits more effective chloroplast phototaxis and, hence, allows more efficient utilization of low-incident photon flux densities. The photosynthetic apparatus is, consequently, protected from damage under high-incident photon flux densities.

Photosynthetic gas exchange under emersed conditions in eulittoral and normally submersed members of the Fucales and the Laminariales: interpretation in relation to C isotope ratio and N and water use efficiency.

Ten species of brown macroalgae (five eulittoral and one submersed species of the Fucales; four submersed species of the Laminariales) from a rocky shore at Arbroath, Scotland, were examined for characteristics of emersed photosynthesis in relation to the partial pressure of CO2 and O2. The five eulittoral species of the Fucaceae were approaching CO2 saturation for light-saturated photosynthesis at normal air levels of CO2 (35 Pa) in 21 kPa O2. The normally submersed algae are further from CO2 saturation under these conditions, especially in the case of the four members of the Laminariales. The rate of net photosynthesis in the Fucaceae is O2-independent in the range 2-21 kPa O2 over the entire range of CO2 partial pressure tested (compensation up to 95 Pa). For the other five algae tested, net photosynthesis is slightly inhibited by O2 at 21 kPa relative to 2 kPa over the entire range of CO2 partial pressures tested (compensation up to 95 Pa). CO2 compensation partial pressures are low (<0.5 Pa) for the Fucaceae and independent of O2 in the range 2-42 kPa. For the other five algae, the CO2 compensation partial pressure are higher, and increased with O2 partial pressure in the range 2-42 kPa. These gas exchange data show that the Fucaceae exhibit more C4-like characteristics of their photosynthetic physiology than do the other five species tested, although even the Laminariales and Halidrys siliquosa are not classic C3 plants in their photosynthetic physiology. These data suggest that, in emersed conditions as well as in the previously reported work on submersed photosynthesis, a "CO2 concentrating mechanism" is operating which, by energized transmembrane transport of inorganic C, accumulates CO2 at the site of RUBISCO and, at least in part, suppresses the oxygenase activity. Work with added extracellular carbonic anhydrase (CA), and with a relatively membrane-impermeant inhibitor of the native extracellular CA activity (acetazolamide), suggests that, in emersed conditions as well as in the previously reported work on algae submersed in seawater at pH 8, HCO inf3sup- is the major inorganic C species entering the cell. At optimal hydration, the rate of emersed photosynthesis in air is not less than the rate of photosynthesis when submersed in seawater, at least for the Fucaceae. δ13C ratios of organic C for the Fucaceae are slightly more negative than is the case for the other five algae; these data are consitent with substantial (half or more of the entering inorganic C) leakage of CO2 from the accumulated pool, and with some contribution of atmospheric CO2 to the organic C gain by the eulittoral algae. The predicted increase in N use efficiency of photosynthesis in the Fucaceae, with their more strongly developed CO2 concentrating mechanism, is consistent with data on emersed, but not submersed, photosynthesis for the algae collected from the wild and thus at a poorly defined N status. The more C4-like gas exchange charateristics of photosynthesis in the eulittoral Fucaceae may be important in increasing the water use efficiency of emersed photosynthesis from the limited capital of water available for transpiration by a haptophyte.

Exogenous inorganic carbon sources for photosynthesis in seawater by members of the Fucales and the Laminariales (Phaeophyta): ecological and taxonomic implications.

Characteristics of inorganic carbon assimilation by photosynthesis in seawater were investigated in six species of the Fucales (five Fucaceae, one Cystoseiraceae) and four species of the Laminariales (three Laminariaceae, one Alariaceae) from Arbroath, Scotland. All of the algae tested could photosynthesise faster at high external pH values than the uncatalysed conversion of HCO 3- to CO2 can occur, i.e. can "use" external HCO 3- . They all had detectable extracellular carbonic anhydrase activity, suggesting that HCO 3- use could involve catalysis of external CO2 production, a view supported to some extent by experiments with an inhibitor of carbonic anhydrase. All of the algae tested had CO2 compensation concentrations at pH 8 which were lower than would be expected from diffusive entry of CO2 supplying RUBISCO as the initial carboxylase, consistent with the operation of energized entry of HCO 3- and / or CO2 acting as a "CO2 concentrating mechanism". Quantitative differences among the algae examined were noted with respect to characteristics of inorganic C assimilation. The most obvious distinction was between the eulittoral Fucaceae, which are emersed for part of, or most of, the tidal cycle, and the other three families (Cystoseiraceae, Laminariaceae, Alariaceae) whose representatives are essentially continually submersed. The Fucaceae examined are able to photosynthesise at high pH values, and have lower CO2 compensation concentrations, and lower K1/2 values for inorganic C use in photosynthesis, at pH 8, than the other algae tested. Furthermore, the Fucaceae are essentially saturated with inorganic C for photosynthesis at the normal seawater concentration at pH 8 and 10°C. These characteristics are consistent with the dominant role of a "CO2 concentrating mechanism" in CO2 acquisition by these plants. Other species tested have characteristcs which suggest a less effective HCO 3- use and "CO2 concentrating mechanism", with the Laminariaceae being the least effective; unlike the Fucaceae, photosynthesis by these algae is not saturated with inorganic C in normal seawater. Taxonomic and ecological implications of these results are considered in relation to related data in the literature.