Oceanic protists with different forms of acquired phototrophy display contrasting biogeographies and abundance.

This first comprehensive analysis of the global biogeography of marine protistan plankton with acquired phototrophy shows these mixotrophic organisms to be ubiquitous and abundant; however, their biogeography differs markedly between different functional groups. These mixotrophs, lacking a constitutive capacity for photosynthesis (i.e. non-constitutive mixotrophs, NCMs), acquire their phototrophic potential through either integration of prey-plastids or through endosymbiotic associations with photosynthetic microbes. Analysis of field data reveals that 40-60% of plankton traditionally labelled as (non-phototrophic) microzooplankton are actually NCMs, employing acquired phototrophy in addition to phagotrophy. Specialist NCMs acquire chloroplasts or endosymbionts from specific prey, while generalist NCMs obtain chloroplasts from a variety of prey. These contrasting functional types of NCMs exhibit distinct seasonal and spatial global distribution patterns. Mixotrophs reliant on 'stolen' chloroplasts, controlled by prey diversity and abundance, dominate in high-biomass areas. Mixotrophs harbouring intact symbionts are present in all waters and dominate particularly in oligotrophic open ocean systems. The contrasting temporal and spatial patterns of distribution of different mixotroph functional types across the oceanic provinces, as revealed in this study, challenges traditional interpretations of marine food web structures. Mixotrophs with acquired phototrophy (NCMs) warrant greater recognition in marine research.

Life and light: exotic photosynthesis in binary and multiple-star systems.

The potential for Earth-like planets within binary/multiple-star systems to host photosynthetic life was evaluated by modeling the levels of photosynthetically active radiation (PAR) such planets receive. Combinations of M and G stars in (i) close-binary systems; (ii) wide-binary systems, and (iii) three-star systems were investigated, and a range of stable radiation environments were found to be possible. These environmental conditions allow for the possibility of familiar, but also more exotic, forms of photosynthetic life, such as IR photosynthesizers and organisms that are specialized for specific spectral niches.

Stomatal vs. genome size in angiosperms: the somatic tail wagging the genomic dog?

BACKGROUND AND AIMS: Genome size is a function, and the product, of cell volume. As such it is contingent on ecological circumstance. The nature of 'this ecological circumstance' is, however, hotly debated. Here, we investigate for angiosperms whether stomatal size may be this 'missing link': the primary determinant of genome size. Stomata are crucial for photosynthesis and their size affects functional efficiency. METHODS: Stomatal and leaf characteristics were measured for 1442 species from Argentina, Iran, Spain and the UK and, using PCA, some emergent ecological and taxonomic patterns identified. Subsequently, an assessment of the relationship between genome-size values obtained from the Plant DNA C-values database and measurements of stomatal size was carried out. KEY RESULTS: Stomatal size is an ecologically important attribute. It varies with life-history (woody species < herbaceous species < vernal geophytes) and contributes to ecologically and physiologically important axes of leaf specialization. Moreover, it is positively correlated with genome size across a wide range of major taxa. CONCLUSIONS: Stomatal size predicts genome size within angiosperms. Correlation is not, however, proof of causality and here our interpretation is hampered by unexpected deficiencies in the scientific literature. Firstly, there are discrepancies between our own observations and established ideas about the ecological significance of stomatal size; very large stomata, theoretically facilitating photosynthesis in deep shade, were, in this study (and in other studies), primarily associated with vernal geophytes of unshaded habitats. Secondly, the lower size limit at which stomata can function efficiently, and the ecological circumstances under which these minute stomata might occur, have not been satisfactorally resolved. Thus, our hypothesis, that the optimization of stomatal size for functional efficiency is a major ecological determinant of genome size, remains unproven.

Darwin--a mission to detect and search for life on extrasolar planets.

The discovery of extrasolar planets is one of the greatest achievements of modern astronomy. The detection of planets that vary widely in mass demonstrates that extrasolar planets of low mass exist. In this paper, we describe a mission, called Darwin, whose primary goal is the search for, and characterization of, terrestrial extrasolar planets and the search for life. Accomplishing the mission objectives will require collaborative science across disciplines, including astrophysics, planetary sciences, chemistry, and microbiology. Darwin is designed to detect rocky planets similar to Earth and perform spectroscopic analysis at mid-infrared wavelengths (6-20 mum), where an advantageous contrast ratio between star and planet occurs. The baseline mission is projected to last 5 years and consists of approximately 200 individual target stars. Among these, 25-50 planetary systems can be studied spectroscopically, which will include the search for gases such as CO(2), H(2)O, CH(4), and O(3). Many of the key technologies required for the construction of Darwin have already been demonstrated, and the remainder are estimated to be mature in the near future. Darwin is a mission that will ignite intense interest in both the research community and the wider public.

Biomineralization by photosynthetic organisms: evidence of coevolution of the organisms and their environment?

Biomineralization is widespread among photosynthetic organisms in the ocean, in inland waters and on land. The most quantitatively important biogeochemical role of land plants today in biomineralization is silica deposition in vascular plants, especially grasses. Terrestrial plants also increase the rate of weathering, providing the soluble substrates for biomineralization on land and in water bodies, a role that has had global biogeochemical impacts since the Devonian. The dominant photosynthetic biomineralizers in today's ocean are diatoms and radiolarians depositing silica and coccolithophores and foraminifera depositing calcium carbonate. Abiotic precipitation of silica from supersaturated seawater in the Precambrian preceded intracellular silicification dominated by sponges, then radiolarians and finally diatoms, with successive declines in the silicic acid concentration in the surface ocean, resulting in some decreases in the extent of silicification and, probably, increases in the silicic acid affinity of the active influx mechanisms. Calcium and bicarbonate concentrations in the surface ocean have generally been supersaturating with respect to the three common calcium carbonate biominerals through geological time, allowing external calcification as well as calcification in compartments within cells or organisms. The forms of calcium carbonate in biominerals, and presumably the evolution of the organisms that produce them, have been influenced by abiotic variations in calcium and magnesium concentrations in seawater, and calcium carbonate deposition has probably also been influenced by carbon dioxide concentration whose variations are in part biologically determined. Overall, there has been less biological feedback on the availability of substrates for calcification than is the case for silicification.

Are there ecological implications for the proposed energetic restrictions on photosynthetic oxygen evolution at high oxygen concentrations?

It has recently been shown that, in subthylakoid particles prepared using detergent, there is inhibition of oxygen production reactions in photosynthesis by thermodynamic feedback from oxygen build-up, with 50% inhibition at 230 kPa partial pressure of oxygen. This article presents a comprehensive analysis of laboratory data on the effects of high oxygen partial pressures on photosynthesis, and on photo-lithotrophic and chemo-organotrophic growth, of oxygen-producing organisms. The article also contains an analysis of the extent to which high oxygen concentrations occur at the site of photosystem II (PSII) activity under natural conditions today and in the past. The conclusion is that the oxygen concentrations found in nature are very unlikely to reach that needed to cause 50% inhibition of the photosynthetic oxygen production reaction in subthylakoid particles, but that it is just possible that a small part of the inhibition of photosynthesis and of photo-lithotrophic growth by oxygen can be attributed to inhibition of oxygen production by PSII.

Influence on photosynthesis of starlight, moonlight, planetlight, and light pollution (reflections on photosynthetically active radiation in the universe).

Photosynthesis on Earth can occur in a diversity of organisms in the photosynthetically active radiation (PAR) range of 10 nmol of photons m(-2) s(-1) to 8 mmol of photons m(-2) s(-1). Similar considerations would probably apply to photosynthetic organisms on Earth-like planets (ELPs) in the continuously habitable zone of other stars. On Earth, starlight PAR is inadequate for photosynthetically supported growth. An increase in starlight even to reach the minimum theoretical levels to allow for photosynthesis would require a universe that was approximately ten million times older, or with a ten million times greater density of stars, than is the case for the present universe. Photosynthesis on an ELP using PAR reflected from a natural satellite with the same size as our Moon, but at the Roche limit, could support a low rate of photosynthesis at full Moon. Photosynthesis on an ELP-like satellite of a Jupiter-sized planet using light reflected from the planet could be almost 1% of the rate in full sunlight on Earth when the planet was full. These potential contributions to photosynthesis require that the contribution is compared with the rate of photosynthesis driven by direct radiation from the star. Light pollution on Earth only energizes photosynthesis by organisms that are very close to the light source. However, effects of light pollution on photosynthesis can be more widespread if the photosynthetic canopy is retained for more of the year, caused by effects on photoperiodism, with implications for the influence of civilizations on photosynthesis.

A role for shoot protein in shoot-root dry matter allocation in higher plants.

BACKGROUND AND AIMS: It is stated in many recent publications that nitrate (NO3-) acts as a signal to regulate dry matter partitioning between the shoot and root of higher plants. Here we challenge this hypothesis and present evidence for the viewpoint that NO3- and other environmental effects on the shoot:root dry weight ratio (S:R) of higher plants are often related mechanistically to changes in shoot protein concentration. METHODS: The literature on environmental effects on S:R is reviewed, focusing on relationships between S:R, growth and leaf NO3- and protein concentrations. A series of experiments carried out to test the proposal that S:R is dependent on shoot protein concentration is highlighted and new data are presented for tobacco (Nicotiana tabacum). KEY RESULTS/EVIDENCE: Results from the literature and new data for tobacco show that S:R and leaf NO3- concentration are not significantly correlated over a range of environmental conditions. A mechanism involving the relative availability of C and N substrates for growth in shoots can explain how shoot protein concentration can influence shoot growth and hence root growth and S:R. Generally, results in the literature are compatible with the hypothesis that macronutrients, water, irradiance and CO2 affect S:R through changes in shoot protein concentration. In detailed studies on several species, including tobacco, a linear regression model incorporating leaf soluble protein concentration and plant dry weight could explain the greater proportion of the variation in S:R within and between treatments over a wide range of conditions. CONCLUSIONS: It is concluded that if NO3- can influence the S:R of higher plants, it does so only over a narrow range of conditions. Evidence is strong that environmental effects on S:R are often related mechanistically to their effects on shoot protein concentration.

Extension growth of Impatiens glandulifera at low irradiance: importance of nitrate and potassium accumulation.

BACKGROUND AND AIMS: The summer annual Impatiens glandulifera can reach 3 m in height within deciduous woodland. The primary objective was to determine if NO(3)(-) accumulation, and hence its osmotic effect, is an important physiological mechanism allowing Impatiens to achieve substantial height under low irradiance. METHODS: Stem extension, concentrations of K(+) and NO(3)(-) in leaves and concentrations of K(+), NO(3)(-) and other inorganic anions, malate, sugars, total N and total osmoticum in stem were measured in I. glandulifera sampled at different irradiance levels in deciduous woodland and in a glasshouse. Also, the energetic costs, as absorbed photons, of generating osmolarity in stem cell vacuoles with KNO(3), K(2)malate or hexose sugar were determined. KEY RESULTS: Results were similar in the woodland and glasshouse. At 50-100 % relative irradiance (Ir; open ground PAR = 100 % Ir) and 2-10 % Ir, plant height increased from 7-14 cm to 130-154 cm in 64-67 d. Leaf and stem NO(3)(-) concentrations were negligible at 50-100 % Ir while K(+), malate(2-) and sugars, respectively, accounted for 33.2-50.1 %, 19.3-20.8 % and 2.0-2.6 % of total osmoticum in stems. At 2-10 % Ir, NO(3)(-) concentrations were four to eight times greater in stems than leaves. Here, NO(3)(-) constituted 26.7-34.3 % of the total osmotic concentration in the stem and NO(3)(-)-N constituted 69-81 % of total N in stem tissue. Also at 2-10 % Ir, K(+) comprised 44.9-45.9 % and malate plus sugars 2.2-3.1 % of total osmotic concentration. The energy cost of osmoticum as KNO(3) was calculated as less than half that of malate and less than one-seventh that for hexose. Further calculations suggest that use of KNO(3), K(2)malate or glucose as osmoticum at low irradiance would, respectively, cost approx. 7 %, 16 % and 50 % of the total construction cost of the stem. CONCLUSIONS: It is concluded that accumulation of NO(3)(-) in place of organic molecules in stems is an important mechanism allowing I. glandulifera to achieve substantial height at low irradiance.

Leaf age-related differences in apoplastic NH(4)(+) concentration, pH and the NH(3) compensation point for a wild perennial.

Extracts of the foliar apoplast of leaves of different ages of Luzula sylvatica (Huds.) Gaud. were prepared by vacuum infiltration and centrifugation. Measurements of pH and concentration were performed on extracts. From these bioassay measurements the relative magnitude of NH(3) compensation points for leaves of different ages were inferred. Young leaves were found to have much higher apoplast pH than old leaves, leading to the calculation of 4-10-fold higher NH(3) compensation points. Such age-related differences in the NH(3) compensation point are considerably larger than those previously reported. Apoplast pH and concentration were found to increase during leaf expansion before declining prior to senescence. Bulk foliar tissue pH, and total N concentrations were also found to be generally higher in young leaves than in old leaves. Where a significant correlation was found, total foliar N, bulk tissue foliar and the calculated NH(3) compensation point were all found to increase with N supplied to roots, whilst apoplast and bulk tissue H(+) concentrations were found to decline. The potential of bulk foliar tissue measurements to act as simple predictors of the NH(3) compensation point is discussed.

Altritol synthesis by Notheia anomala.

The linear hexitol altritol has only been identified in six genera, all of them in the order Fucales of the brown algae. Five of these genera are closely related according to molecular phylogenetic and other data, while the sixth (Notheia) is an obligate epiphyte on two other altritol-containing genera with which it is symphanic. The possibility that Notheia obtains altritol from the algae on which it is epiphytic rather than by synthesizing altritol independently was investigated by supplying 13C-inorganic carbon in the light followed by mass spectrometric and nuclear magnetic resonance analysis. Notheia separated from the phorophyte Hormosira during exposure to 13C showed 13C enrichment in both altritol and mannitol, while the Hormosira only showed significant labelling of mannitol. These data show that altritol can be synthesized by Notheia, with implications for the number of gains and losses of the capacity to synthesize altritol in the evolution of the Fucales.

Phosphorus limitation of nitrogen fixation by Trichodesmium in the central Atlantic Ocean.

Marine fixation of atmospheric nitrogen is believed to be an important source of biologically useful nitrogen to ocean surface waters, stimulating productivity of phytoplankton and so influencing the global carbon cycle. The majority of nitrogen fixation in tropical waters is carried out by the marine cyanobacterium Trichodesmium, which supplies more than half of the new nitrogen used for primary production. Although the factors controlling marine nitrogen fixation remain poorly understood, it has been thought that nitrogen fixation is limited by iron availability in the ocean. This was inferred from the high iron requirement estimated for growth of nitrogen fixing organisms and the higher apparent densities of Trichodesmium where aeolian iron inputs are plentiful. Here we report that nitrogen fixation rates in the central Atlantic appear to be independent of both dissolved iron levels in sea water and iron content in Trichodesmium colonies. Nitrogen fixation was, instead, highly correlated to the phosphorus content of Trichodesmium and was enhanced at higher irradiance. Furthermore, our calculations suggest that the structural iron requirement for the growth of nitrogen-fixing organisms is much lower than previously calculated. Although iron deficiency could still potentially limit growth of nitrogen-fixing organisms in regions of low iron availability-for example, in the subtropical North Pacific Ocean-our observations suggest that marine nitrogen fixation is not solely regulated by iron supply.

Roots: evolutionary origins and biogeochemical significance.

Roots, as organs distinguishable developmentally and anatomically from shoots (other than by occurrence of stomata and sporangia on above-ground organs), evolved in the sporophytes of at least two distinct lineages of early vascular plants during their initial major radiation on land in Early Devonian times (c. 410-395 million years ago). This was some 15 million years after the appearance of tracheophytes and c. 50 million years after the earliest embryophytes of presumed bryophyte affinity. Both groups are known initially only from spores, but from comparative anatomy of extant bryophytes and later Lower Devonian fossils it is assumed that, during these times, below-ground structures (if any) other than true roots fulfilled the functions of anchorage and of water and nutrient acquisition, despite lacking an endodermis (as do the roots of extant Lycopodium spp.). By 375 million years ago root-like structures penetrated almost a metre into the substratum, greatly increasing the volume of mineral matter subject to weathering by the higher than atmospheric CO(2) levels generated by plant and microbial respiration in material with restricted diffusive contact with the atmosphere. Chemical weathering consumes CO(2) in converting silicates into bicarbonate and Si(OH)(4). The CO(2) consumed in weathering ultimately came from atmospheric CO(2) via photosynthesis and respiration; this use of CO(2) probably accounts for most of the postulated 10-fold decrease in atmospheric CO(2) from 400-350 million years ago, with significant effects on shoot evolution. Subsequent evolution of roots has yielded much-branched axes down to 40 microm diameter, a lower limit set by long-distance transport constraints. Finer structures involved in the uptake of nutrients of low diffusivity in soil evolved at least 400 million years ago as arbuscular mycorrhizas or as evaginations of "roots" ("root hairs").

Significance of epidermal fusion and intercalary growth for angiosperm evolution.

The ancestral angiosperm flower probably had many separate elements in each floral whorl (sepals, petals, stamens and carpels). Derived character states include "fusion" of elements within a whorl (cohesion) and fusion between whorls (adhesion), as well as epigyny and the emergence of the other floral elements from the apex of the fused carpels. This article considers the roles of epidermal fusion and intercalary growth in the phylogeny and ontogeny of fused floral elements, and the importance of fusion for angiosperm evolution.

Comparison of gas exchange and bioassay determinations of the ammonia compensation point in Luzula sylvatica (Huds.) Gaud.

Determinations of the NH(3) compensation point for the understory plant of semi-natural woodlands Luzula sylvatica (Huds.) Gaud. were carried out by measurements of gas exchange and by calculation from the NH(4)(+) concentration and pH of extracts of the foliar apoplast. Compensation points determined by gas exchange measurements were among the lowest yet reported (0.51-1.10 microg NH(3) m(-3)) and those calculated from apoplast extracts were lower than any yet reported (0.017-0.54 microg NH(3) m(-3)). Those determined by gas exchange were consistently found to be between 2 and 30 times higher than those determined from apoplast extracts. Consideration of possible causes of this discrepancy, which is not confined to this investigation, showed that all likely errors would result in an increase in the discrepancy, or were insufficient to account for observed differences. It is suggested that spatial variability of pH and NH(4)(+) concentration within the foliar apoplast represents the most promising line for further investigation. It is also shown that the foliar apoplast of L. sylvatica is sufficiently buffered to eliminate the need for correction of H(+) concentration for dilution during extraction, but that it is necessary to correct the NH(4)(+) concentration of apoplast extracts for dilution.

Iron stress restricts photosynthetic intersystem electron transport in Synechococcus sp. PCC 7942.

Although exposure of Synechococcus sp. PCC 7942 to iron stress induced the accumulation of the isiA gene product (CP43') compared with non-stressed controls, immunodetection of the N-terminus of cytochrome (Cyt) f indicated that iron stress not only reduced the content of the 40 kDa, heme-binding, Cyt f polypeptide by 32% but it also specifically induced the accumulation of a new, 23 kDa, non-heme-binding, putative Cyt f polypeptide. Concomitantly, iron stress restricted intersystem electron transport based on the in vivo reduction of P700(+), monitored as delta A(820)/A(820) in the presence and absence of electron transport inhibitors, as well as the inhibition of the Emerson enhancement effect on O(2) evolution. However, iron stress appeared to be associated with enhanced rates of PS I cyclic electron transport, low rates of PS I-driven photoreduction of NADP(+) but comparable rates for PS II+PS I photoreduction of NADP(+) relative to controls. We hypothesize that Synechococcus sp. PCC 7942 exhibits a dynamic capacity to uncouple PS II and PS I electron transport, which may allow for the higher than expected growth rates observed during iron stress.

Land plant biochemistry.

Biochemical studies have complemented ultrastructural and, subsequently molecular genetic evidence consistent with the Charophyceae being the closest extant algal relatives of the embryophytes. Among the genes used in such molecular phylogenetic studies is that rbcL) for the large subunit of ribulose bisphosphate carboxylase-oxygenase (RUBISCO). The RUBISCO of the embryophytes is derived, via the Chlorophyta. from that of the cyanobacteria. This clade of the molecular phylogeny of RUBISCO shows a range of kinetic characteristics, especially of CO2 affinities and of CO2/O2 selectivities. The range of these kinetic values within the bryophytes is no greater than in the rest of the embryophytes; this has implications for the evolution of the embryophytes in the high atmospheric CO2 environment of the late Lower Palaeozoic. The differences in biochemistry between charophycean algae and embryophytes can to some extent be related functionally to the structure and physiology of embryophytes. Examples of components of embryophytes, which are qualitatively or quantitatively different from those of charophytes, are the water repellent/water resistant extracellular lipids, the rigid phenolic polymers functional in water-conducting elements and mechanical support in air, and in UV-B absorption, flavonoid phenolics involved in UV-B absorption and in interactions with other organisms, and the greater emphasis on low Mr organic acids. retained in the plant as free acids or salts, or secreted to the rhizosphere. The roles of these components are discussed in relation to the environmental conditions at the time of evolution of the terrestrial embryophytes. A significant point about embryophytes is the predominance of nitrogen-free extracellular structural material (a trait shared by most algae) and UV-B screening components, by contrast with analogous components in many other organisms. An important question, which has thus far been incompletely addressed, is the extent to which the absence from bryophytes of the biochemical pathways which produce components found only in tracheophytes is the result of evolutionary loss of these functions.

Effect of elevated CO2 on the stomatal distribution and leaf physiology of Alnus glutinosa.

Variation in stomatal development and physiology of mature leaves from Alnus glutinosa plants grown under reference (current ambient, 360 µmol mol-1 CO2 ) and double ambient (720 µmol mol-1 CO2 ) carbon dioxide (CO2 ) mole fractions is assessed in terms of relative plant growth, stomatal characters (i.e. stomatal index and density) and leaf photosynthetic characters. This is the first study to consider the effects of elevated CO2 concentration on the distribution of stomata and epidermal cells across the whole leaf and to try to ascertain the cause of intraleaf variation. In general, a doubling of the atmospheric CO2 concentration enhanced plant growth and significantly increased stomatal index. However, there was no significant change in relative stomatal density. Under elevated CO2 concentration there was a significant decrease in stomatal conductance and an increase in assimilation rate. However, no significant differences were found for the maximum rate of carboxylation (Vcmax ) and the light saturated rate of electron transport (Jmax ) between the control and elevated CO2 treatment.