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Biomedical subjects

John A Raven

Publications and source records attributed to John A Raven.

17 recordsLinked to original sources

Sensing inorganic carbon: CO2 and HCO3-.

Enzymes and transporters that catalyse reactions involving inorganic carbon are well characterized with respect to the species of inorganic carbon (CO2 or HCO3-) with which they interact. There is less information on the species recognized by proteins that sense inorganic carbon. In this issue of the Biochemical Journal, Hammer and colleagues show conclusively that cyanobacterial adenylyl cyclases are activated by CO2 and not HCO3-, as was believed previously. While in some circumstances a similar in vivo regulatory outcome is achieved from sensing HCO3- as from sensing CO2, there are cases in which the outcomes are significantly different. The most striking example is where a compartment lacks carbonic anhydrase yet supports large metabolic fluxes of inorganic carbon species so that CO2 and HCO3- are not at equilibrium. Other examples involve changes in pH, or temperature, of a compartment containing an equilibrium mixture of CO2 and HCO3-.

Acid-Base Equilibrium↗

Limitations on microalgal growth at very low photon fluence rates: the role of energy slippage.

The lower limits of photosynthetically useable radiation at which growth and photosynthesis can occur establish the lower boundaries for the extent of photolithotrophy in the biosphere. Photolithotrophic growth denotes the capacity to grow with photons as the sole energy input. Slippage in terms of photosynthetic energy conversion implies a less than theoretical stoichiometry of energy-transduction process(es) such as the dissipation of intermediates of O(2) evolution and of ATP synthesis (H(+)/e(-) and H(+)/ATP ratios). Slippage is particularly important in limiting the growth of photolithotrophic organisms at very low photon fluence rates. We found that Dunaliella tertiolecta and Phaeodactylum tricornutum avoid such reductions in photon use efficiency by increasing the size and number of their photosynthetic units, respectively, and by altering Q(A) reduction kinetics on the reducing side of PS II. P. tricornutum is also less susceptible to slippage in terms of the breakdown of intermediates in its O(2) evolution pathway than D. tertiolecta. Minimizing H(+) leakage through the CF(0)-CF(1) ATP synthetase (and other H(+ )porters) is also discussed briefly. In combination, strategies employed by P. tricornutum effectively allow it to grow and photosynthesize at lower rates of energy input than D. tertiolecta, consistent with our observations. Differences in the responses of the photosynthetic apparatus of these two marine microalgae are mechanistic and probably representative of evolutionary divergences associated with strategies for dealing with environmental perturbations.

Energy Transfer↗

Homeostasis of cell composition during prolonged darkness.

The chemical composition of organisms in relation to their environmental resource availability is an area of intense research activity. We studied the changes in cell composition of the cyanobacterium Phormidium autumnale in response to prolonged darkness. Cells allocated their internal resources in a homeostatic manner, oxidizing all the three major cellular constituents in a proportional way. This resulted in constant C/N and carbohydrates, lipids and proteins ratios that remained unaltered throughout the whole incubation period. We propose the maintenance of balanced cell composition (homeostasis) as an evolutionary strategy related to environmental transitory changes.

Bacterial Proteins↗

CO2 concentrating mechanisms in algae: mechanisms, environmental modulation, and evolution.

The evolution of organisms capable of oxygenic photosynthesis paralleled a long-term reduction in atmospheric CO2 and the increase in O2. Consequently, the competition between O2 and CO2 for the active sites of RUBISCO became more and more restrictive to the rate of photosynthesis. In coping with this situation, many algae and some higher plants acquired mechanisms that use energy to increase the CO2 concentrations (CO2 concentrating mechanisms, CCMs) in the proximity of RUBISCO. A number of CCM variants are now found among the different groups of algae. Modulating the CCMs may be crucial in the energetic and nutritional budgets of a cell, and a multitude of environmental factors can exert regulatory effects on the expression of the CCM components. We discuss the diversity of CCMs, their evolutionary origins, and the role of the environment in CCM modulation.

Biological Evolution↗

The evolution of modern eukaryotic phytoplankton.

The community structure and ecological function of contemporary marine ecosystems are critically dependent on eukaryotic phytoplankton. Although numerically inferior to cyanobacteria, these organisms are responsible for the majority of the flux of organic matter to higher trophic levels and the ocean interior. Photosynthetic eukaryotes evolved more than 1.5 billion years ago in the Proterozoic oceans. However, it was not until the Mesozoic Era (251 to 65 million years ago) that the three principal phytoplankton clades that would come to dominate the modern seas rose to ecological prominence. In contrast to their pioneering predecessors, the dinoflagellates, coccolithophores, and diatoms all contain plastids derived from an ancestral red alga by secondary symbiosis. Here we examine the geological, geochemical, and biological processes that contributed to the rise of these three, distantly related, phytoplankton groups.

Biodiversity↗

Global aspects of C/N interactions determining plant-environment interactions.

The atomic C:N ratio in photolithotrophs is a function of their content of nucleic acids, proteins, lipids, polysaccharides, and other organic materials, and varies from about 5 in some protein-rich microalgae to much higher values in macroalgae and in higher plants with relatively more structural and energy storage materials. These differences in C:N ratios among organisms means that there is more N assimilation by photosynthetic organisms in the oceans than on land despite the near equality of global photosynthetic C assimilation rates in the two environments. Aquatic organisms obtain inorganic C and inorganic N from the surrounding water. Terrestrial photolithotrophs obtain inorganic C, dinitrogen (by diazotrophy) and some combined N from the atmosphere, with the remaining combined N coming from the soil. The nitrogen cost of growth (biomass production rate per unit plant N) varies with the C:N ratio and specific growth rate of the organism. The C:N ratio of plants can be increased with no, or minimal, decrease in growth rate by switching from N-containing to N-free solutes involved in, for example, UV-B screening or by reducing the content of particular proteins. The water cost of growth (water lost per unit biomass gain) in terrestrial plants is a function of N supply and of C supply; water cost is lower with higher N and C availability. Water supply is also important in determining denitrification rates on land and on N (and C) fluxes from terrestrial to aquatic systems.

Carbon↗

Genomics and chloroplast evolution: what did cyanobacteria do for plants?

The complete genome sequences of cyanobacteria and of the higher plant Arabidopsis thaliana leave no doubt that the plant chloroplast originated, through endosymbiosis, from a cyanobacterium. But the genomic legacy of cyanobacterial ancestry extends far beyond the chloroplast itself, and persists in organisms that have lost chloroplasts completely.

Chloroplasts↗

Genomes at the interface between bacteria and organelles.

The topic of the transition of the genome of a free-living bacterial organism to that of an organelle is addressed by considering three cases. Two of these are relatively clear-cut as involving respectively organisms (cyanobacteria) and organelles (plastids). Cyanobacteria are usually free-living but some are involved in symbioses with a range of eukaryotes in which the cyanobacterial partner contributes photosynthesis, nitrogen fixation, or both of these. In several of these symbioses the cyanobacterium is vertically transmitted, and in a few instances, sufficient unsuccessful attempts have been made to culture the cyanobiont independently for the association to be considered obligate for the cyanobacterium. Plastids clearly had a cyanobacterial ancestor but cannot grow independently of the host eukaryote. Plastid genomes have at most 15% of the number of genes encoded by the cyanobacterium with the smallest number of genes; more genes than are retained in the plastid genome have been transferred to the eukaryote nuclear genome, while the rest of the cyanobacterial genes have been lost. Even the most cyanobacteria-like plastids, for example the "cyanelles" of glaucocystophyte algae, are functionally and genetically very similar to other plastids and give little help in indicating intermediates in the evolution of plastids. The third case considered is the vertically transmitted intracellular bacterial symbionts of insects where the symbiosis is usually obligate for both partners. The number of genes encoded by the genomes of these obligate symbionts is intermediate between that of organelles and that of free-living bacteria, and the genomes of the insect symbionts also show rapid rates of sequence evolution and AT (adenine, thymine) bias. Genetically and functionally, these insect symbionts show considerable similarity to organelles.

Animals↗

Inorganic carbon concentrating mechanisms in relation to the biology of algae.

Significant advances have recently been made in our understanding of the mechanism of inorganic carbon transport in algae and, especially, cyanobacteria with inorganic carbon concentrating mechanisms (CCMs). Furthermore, the role of CCMs in increasing the rate of photosynthesis in air-equilibrated solutions is also quite well understood. However, less often considered is how the presence (or absence) of a CCM relates to the biology of algae. This mini-review relates the occurrence of algal CCMs to phylogeny, life form, life history, and interactions with other organisms. While some patterns can be seen, the occurrence of CCMs in relation to the overall biology of the algae needs more investigation.

Journal Article↗

An anaplerotic role for mitochondrial carbonic anhydrase in Chlamydomonas reinhardtii.

Previous studies of the mitochondrial carbonic anhydrase (mtCA) of Chlamydomonas reinhardtii showed that expression of the two genes encoding this enzyme activity required photosynthetically active radiation and a low CO(2) concentration. These studies suggested that the mtCA was involved in the inorganic carbon-concentrating mechanism. We have now shown that the expression of the mtCA at low CO(2) concentrations decreases when the external NH(4)(+) concentration decreases, to the point of being undetectable when NH(4)(+) supply restricts the rate of photoautotrophic growth. The expression of mtCA can also be induced at supra-atmospheric partial pressure of CO(2) by increasing the NH(4)(+) concentration in the growth medium. Conditions that favor mtCA expression usually also stimulate anaplerosis. We therefore propose that the mtCA is involved in supplying HCO(3)(-) for anaplerotic assimilation catalyzed by phosphoenolpyruvate carboxylase, which provides C skeletons for N assimilation under some circumstances.

Animals↗

An in situ study of photosynthetic oxygen exchange and electron transport rate in the marine macroalga Ulva lactuca (Chlorophyta).

Direct comparisons between photosynthetic O(2) evolution rate and electron transport rate (ETR) were made in situ over 24 h using the benthic macroalga Ulva lactuca (Chlorophyta), growing and measured at a depth of 1.8 m, where the midday irradiance rose to 400-600 mumol photons m(-2) s(-1). O(2) exchange was measured with a 5-chamber data-logging apparatus and ETR with a submersible pulse amplitude modulated (PAM) fluorometer (Diving-PAM). Steady-state quantum yield ((F(m)'-F(t))/F(m)') decreased from 0.7 during the morning to 0.45 at midday, followed by some recovery in the late afternoon. At low to medium irradiances (0-300 mumol photons m(-2) s(-1)), there was a significant correlation between O(2) evolution and ETR, but at higher irradiances, ETR continued to increase steadily, while O(2) evolution tended towards an asymptote. However at high irradiance levels (600-1200 mumol photons m(-2) s(-1)) ETR was significantly lowered. Two methods of measuring ETR, based on either diel ambient light levels and fluorescence yields or rapid light curves, gave similar results at low to moderate irradiance levels. Nutrient enrichment (increases in [NO(3) (-)], [NH(4) (+)] and [HPO(4) (2-)] of 5- to 15-fold over ambient concentrations) resulted in an increase, within hours, in photosynthetic rates measured by both ETR and O(2) evolution techniques. At low irradiances, approximately 6.5 to 8.2 electrons passed through PS II during the evolution of one molecule of O(2), i.e., up to twice the theoretical minimum number of four. However, in nutrient-enriched treatments this ratio dropped to 5.1. The results indicate that PAM fluorescence can be used as a good indication of the photosynthetic rate only at low to medium irradiances.

Journal Article↗

Seaweeds in cold seas: evolution and carbon acquisition.

Much evidence suggests that life originated in hydrothermal habitats, and for much of the time since the origin of cyanobacteria (at least 2.5 Ga ago) and of eukaryotic algae (at least 2.1 Ga ago) the average sea surface and land surface temperatures were higher than they are today. However, there have been at least four significant glacial episodes prior to the Pleistocene glaciations. Two of these (approx. 2.1 and 0.7 Ga ago) may have involved a 'Snowball Earth' with a very great impact on the algae (sensu lato) of the time (cyanobacteria, Chlorophyta and Rhodophyta) and especially those that were adapted to warm habitats. By contrast, it is possible that heterokont, dinophyte and haptophyte phototrophs only evolved after the Carboniferous-Permian ice age (approx. 250 Ma ago) and so did not encounter low (</=5 degrees C) sea surface temperatures until the Antarctic cooled some 15 Ma ago. Despite this, many of the dominant macroalgae in cooler seas today are (heterokont) brown algae, and many laminarians cannot reproduce at temperatures above 18-25 degrees C. By contrast to plants in the aerial environment, photosynthetic structures in water are at essentially the same temperature as the fluid medium. The impact of low temperatures on photosynthesis by marine macrophytes is predicted to favour diffusive CO(2) entry rather than a CO(2)-concentrating mechanism. Some evidence favours this suggestion, but more data are needed.

Antarctic Regions↗

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.

Biological Transport↗