A global two component signal transduction system that integrates the control of photosynthesis, carbon dioxide assimilation, and nitrogen fixation

Photosynthesis, biological nitrogen fixation, and carbon dioxide assimilation are three fundamental biological processes catalyzed by photosynthetic bacteria. In the present study, it is shown that mutant strains of the nonsulfur purple photosynthetic bacteria Rhodospirillum rubrum and Rhodobacter sphaeroides, containing a blockage in the primary CO2 assimilatory pathway, derepress the synthesis of components of the nitrogen fixation enzyme complex and abrogate normal control mechanisms. The absence of the Calvin–Benson–Bassham (CBB) reductive pentose phosphate CO2 fixation pathway removes an important route for the dissipation of excess reducing power. Thus, the mutant strains develop alternative means to remove these reducing equivalents, resulting in the synthesis of large amounts of nitrogenase even in the presence of ammonia. This response is under the control of a global two-component signal transduction system previously found to regulate photosystem biosynthesis and the transcription of genes required for CO2 fixation through the CBB pathway and alternative routes. In addition, this two-component system directly controls the ability of these bacteria to grow under nitrogen-fixing conditions. These results indicate that there is a molecular link between the CBB and nitrogen fixation process, allowing the cell to overcome powerful control mechanisms to remove excess reducing power generated by photosynthesis and carbon metabolism. Furthermore, these results suggest that the two-component system integrates the expression of genes required for the three processes of photosynthesis, nitrogen fixation, and carbon dioxide fixation.

Potential responses of soil organic carbon to global environmental change

Recent improvements in our understanding of the dynamics of soil carbon have shown that 20–40% of the approximately 1,500 Pg of C stored as organic matter in the upper meter of soils has turnover times of centuries or less. This fast-cycling organic matter is largely comprised of undecomposed plant material and hydrolyzable components associated with mineral surfaces. Turnover times of fast-cycling carbon vary with climate and vegetation, and range from <20 years at low latitudes to >60 years at high latitudes. The amount and turnover time of C in passive soil carbon pools (organic matter strongly stabilized on mineral surfaces with turnover times of millennia and longer) depend on factors like soil maturity and mineralogy, which, in turn, reflect long-term climate conditions. Transient sources or sinks in terrestrial carbon pools result from the time lag between photosynthetic uptake of CO2 by plants and the subsequent return of C to the atmosphere through plant, heterotrophic, and microbial respiration. Differential responses of primary production and respiration to climate change or ecosystem fertilization have the potential to cause significant interrannual to decadal imbalances in terrestrial C storage and release. Rates of carbon storage and release in recently disturbed ecosystems can be much larger than rates in more mature ecosystems. Changes in disturbance frequency and regime resulting from future climate change may be more important than equilibrium responses in determining the carbon balance of terrestrial ecosystems.

A long marine history of carbon cycle modulation by orbital-climatic changes

Pacing of the marine carbon cycle by orbital forcing during the Pliocene and Pleistocene Ice Ages [past 2.5 million years (Myr)] is well known. As older deep-sea sediment records are being studied at greater temporal resolution, it is becoming clear that similar fluctuations in the marine carbon system have occurred throughout the late Mesozoic and Tertiary, despite the absence of large continental ice sheets over much of this time. Variations in both the organic and the calcium carbonate components of the marine carbon system seem to have varied cyclically in response to climate forcing, and carbon and carbonate time series appear to accurately characterize the frequency spectrum of ancient climatic change. For the past 35 Myr, much of the variance in carbonate content carries the “polar” signal of obliquity [41,000 years (41 kyr)] forcing. Over the past 125 Myr, there is evidence from marine sediments of the continued role of precessional (≈21 kyr) climatic cycles. Repeat patterns of sedimentation at about 100, 400, and 2,400 kyr, the modulation periods of precession, persistently enter into marine carbon cycle records as well. These patterns suggest a nonlinear response of climate and/or the sedimentation of organic carbon and carbonates to precessional orbital perturbations. Nonlinear responses of the carbon system may help to amplify relatively weak orbital insolation anomalies into more significant climatic perturbations through positive feedback effects. Nonlinearities in the carbon cycle may have transformed orbital-climatic cycles into long-wavelength features on time scales comparable to the residence times of carbon and nutrient elements in the ocean.