A PERSPECTIVE ON PHOTOSYNTHESIS IN THE OLIGOTROPHIC OCEANS: HYPOTHESES CONCERNING ALTERNATE ROUTES OF ELECTRON FLOW1

Many regions of the open, oligotrophic oceans are depleted of nutrients, especially nitrogen and iron. The biogenesis and the functioning of the photosynthetic apparatus may be specialized and tailored to the various marine habitats. In this minireview, we discuss some new findings with respect to photosynthetic processes in the oceans. We focus on findings that suggest that some cyanobacteria may route electrons derived from the splitting of H₂O to the reduction of O₂ and H⁺ in a water‐to‐water cycle, and that other cyanobacteria that fix nitrogen during the day are likely missing PSII and enzymes involved in the fixation of inorganic carbon. Both of these proposed “variant” forms of photosynthetic electron flow provide new insights into ways in which marine phytoplankton satisfy their energetic and nutritive requirements.     

STATUS, RELATIONSHIPS, AND DISTRIBUTION OF MESOPLODON BOWDOINI ANDREWS, 1908 (CETACEA: ZIPHIIDAE)

The specific status of Mesoplodon bowdoini Andrews is reviewed and new information on its morphology, reproduction, and distribution is presented. This species of beaked whale, known only from 35 specimens, has a southern, circumpolar distribution north of the Antarctic convergence, between 32° and 54°30'S. It shares with M. bahamondi Reyes, Van Waerebeek, Cárdenas and Yáñez from the south Pacific Ocean including New Zealand (this paper) and M. carlhubbsi Moore from the north Pacific, a number of morphological features such as prominential notches in the maxillary bones in the skull. It is less similar to M. stejnegeri True from the north Pacific and M. ginkgodens Nishiwaki and Kamiya from the tropical Indo-Pacific. Mesoplodon bowdoini can be distinguished from all other species of Mesoplodon by the shape of its teeth (male and female), and differences in the morphology of its skull, especially the proportions of the rostrum, separation of the nasals, the shape of the prominential notches, and the nature of the antorbital processes. The species' distinguishing external characteristics are: a robust body up to about 4.50 m long; a low melon and short, thick beak; an elevated jawline posteriorly; and a low, blunt-tipped, triangular dorsal fin. The occurrence of fetuses of M. bowdoini in May and September, and perinatal juveniles in May and June, indicates a summer-autumn breeding season in the New Zealand region; the length at birth is estimated at about 2.20 m.     

American Security and Law of the Sea

With the election of President Barack Obama and renewed interest in the Law of the Sea Convention inside the United States, this article pulls from obscurity the key U.S. declarations, understandings, and conditions of ratification to the Treaty, considers their context and meaning for U.S. security interests, and helps to place them into the lexicon of oceans law and policy that informs the greater dialogue of international security and strategy.     

Europa as an Abode of Life

Life as we know it on Earth depends on liquid water, a suite of `biogenic' elements (most famously carbon) and a useful source of free energy. Here we review Europa's suitability for life from the perspective of these three requirements. It is likely, though not yet certain, that Europa harbors a subsurface ocean of liquid water whose volume is about twice that of Earth's oceans. Little is known about Europa's inventory of carbon, nitrogen, and other biogenic elements, but lower bounds on these can be placed by considering the role of cometary delivery over Europa's history. Sources of free energy are challenging for a world covered with an ice layer kilometers thick, but it is possible that hydrothermal activity and/or organics and oxidants provided by the action of radiation chemistry at Europa's surface and subsequent mixing into Europa's ocean could provide the electron donors and acceptors needed to power a Europan ecosystem. It is not premature to draw lessons from the search for life on Mars with the Viking spacecraft for planning exobiological missions to Europa.     

Seasonal adaptations and the role of lipids in oceanic zooplankton

Oceanic zooplankton species exhibit quite diverse life history traits. A major driving force determining their life strategies is the seasonal variability in food supply, which is most pronounced in polar oceans where fluctuations in primary production are extreme. Seasonal adaptations are closely related to the trophic level of zooplankters, with strongest pressures occurring on herbivorous organisms. The dominant grazers, calanoid copepods and krill (Euphausiacea), have developed fascinating solutions for successful overwintering at higher latitudes. They usually exhibit a very efficient storage and utilization of energy reserves to reduce the effect of a highly seasonal primary production. The predominant larger Calanus species from the Arctic and Calanoides acutus from the Antarctic biosynthesize large amounts of high-energy wax esters with long-chain monounsaturated fatty acids and alcohols (20:1 and 22:1 isomers) as major components. They survive the dark season at depth in a stage of dormancy called diapause. In contrast, the Antarctic Calanus propinquus, a winter-active species, synthesizes primarily triacylglycerols, which are dominated by long-chain monounsaturated fatty acids with 22 carbon atoms (2 isomers) and yield even higher calorific contents. The omnivorous and carnivorous species, which are less subjected to seasonal food shortage, usually do not exhibit such an elaborate lipid biosynthesis. Herbivores usually do not utilize much of their enormous lipid reserves for overwintering, but channel this energy towards reproductive processes in late winter/early spring. Timing of reproduction is critical especially at high latitudes due to the short production period, and lipid reserves ensure early spawning independent of external resources. These energetic adaptations (dormancy, lipid storage) are supplemented by other life strategies such as extensive vertical migrations, change in the mode of life, and trophic flexibility.     

The role of phytoplankton photosynthesis in global biogeochemical cycles

Phytoplankton biomass in the world's oceans amounts to only 1–2% of the total global plant carbon, yet these organisms fix between 30 and 50 billion metric tons of carbon annually, which is about 40% of the total. On geological time scales there is profound evidence of the importance of phytoplankton photosynthesis in biogeochemical cycles. It is generally assumed that present phytoplankton productivity is in a quasi steady-state (on the time scale of decades). However, in a global context, the stability of oceanic photosynthetic processes is dependent on the physical circulation of the upper ocean and is therefore strongly influenced by the atmosphere. The net flux of atmospheric radiation is critical to determining the depth of the upper mixed layer and the vertical fluxes of nutrients. These latter two parameters are keys to determining the intensity, and spatial and temporal distributions of phytoplankton blooms. Atmospheric radiation budgets are not in steady-state. Driven largely by anthropogenic activities in the 20th century, increased levels of IR- absorbing gases such as CO₂, CH₄ and CFC's and NO_x will potentially increase atmospheric temperatures on a global scale. The atmospheric radiation budget can affect phytoplankton photosynthesis directly and indirectly. Increased temperature differences between the continents and oceans have been implicated in higher wind stresses at the ocean margins. Increased wind speeds can lead to higher nutrient fluxes. Throughout most of the central oceans, nitrate concentrations are sub-micromolar and there is strong evidence that the quantum efficiency of Photosystem II is impaired by nutrient stress. Higher nutrient fluxes would lead to both an increase in phytoplankton biomass and higher biomass-specific rates of carbon fixation. However, in the center of the ocean gyres, increased radiative heating could reduce the vertical flux of nutrients to the euphotic zone, and hence lead to a reduction in phytoplankton carbon fixation. Increased desertification in terrestrial ecosystems can lead to increased aeolean loadings of essential micronutrients, such as iron. An increased flux of aeolean micronutrients could fertilize nutrient-replete areas of the open ocean with limiting trace elements, thereby stimulating photosynthetic rates. The factors which limit phytoplankton biomass and photosynthesis are discussed and examined with regard to potential changes in the Earth climate system which can lead the oceans away from steady-state. While it is difficult to confidently deduce changes in either phytoplankton biomass or photosynthetic rates on decadal time scales, time-series analysis of ocean transparency data suggest long-term trends have occurred in the North Pacific Ocean in the 20th century. However, calculations of net carbon uptake by the oceans resulting from phytoplankton photosynthesis suggest that without a supply of nutrients external to the ocean, carbon fixation in the open ocean is not presently a significant sink for excess atmospheric CO₂.The submitted paper has been authored under Contract No. DE-AC02-76H00016 with the US Department of Energy. Accordingly, the US Government retains a non-exclusive, royalty-free license to publish or reproduce the published form of this contribution, or allow others to do so, for US Government purposes.     

Common evolutionary origin of planktonic and benthic nitrogen-fixing oscillatoriacean cyanobacteria from tropical oceans

The filamentous cyanobacteria belonging to the genus Hydrocoleum (Blennothrix) are among the most common mat-forming cyanobacteria in tropical oceans. We present here the evidence that these benthic cyanobacteria are morphologically and phylogenetically very close to the planktonic species of Trichodesmium. Genetic relationship was established independently with regard to sequences of the 16S rRNA gene, nifH gene, and phycocyanin and phycoerythrin intergenic spacers. The species of both genera formed a common distinct branch in phylogenetically reconstructed cyanobacterial trees, suggesting that the main constituents of cyanobacterial benthos and plankton have an early common origin and both represent major contributors to nitrogen budget of tropical oceans today as in the distant geological past.     

United States legislation and the polar oceans

The United States has important national interests vested in both the Arctic and Antarctic Oceans. Thus, in recent decades the United States has progressively codified its national commitment to conserve and manage both these marine regions and their resources and to protect them from activities that might produce adverse impacts. The U.S. legislation passed since the 1960s selectively affects both polar regions and supports major American policy objectives there: to maintain the Arctic and Antarctic as areas of international cooperation for peaceful purposes; to satisfy economic needs, especially hydrocarbon, mineral, and living resources; to protect opportunities for scientific research; to protect the marine environment; and to conserve living resources in the circumpolar seas. Future trends suggest a shift in U.S. policy attention to the Arctic. Accordingly, new U.S. legislation will be needed to regulate increased activities in the polar north.