Nutrient supply regulates the activity of phytoplankton, but the global biogeography of nutrient limitation and co-limitation is poorly understood. Prochlorococcus adapt to local environments by gene gains and losses, and we used genomic changes as an indicator of adaptation to nutrient stress. We collected metagenomes from all major ocean regions as part of the Global Ocean Ship-based Hydrographic Investigations Program (Bio-GO-SHIP) and quantified shifts in genes involved in nitrogen, phosphorus, and iron assimilation. We found regional transitions in stress type and severity as well as widespread co-stress. Prochlorococcus stress genes, bottle experiments, and Earth system model predictions were correlated. We propose that the biogeography of multinutrient stress is stoichiometrically linked by controls on nitrogen fixation. Our omics-based description of phytoplankton resource use provides a nuanced and highly resolved description of nutrient stress in the global ocean.

Thermal algae in alkaline hot springs of Yellowstone National Park (Wyoming) grow as compact mats in which self‐shading is extensive, as shown by measurement by autoradiography of photosynthetic activity of cells at different levels in the mat. The effect of light intensity on photosynthesis of the algal mats was studied using neutral density filters during incubation with 14CO2. Despite the intense sunlight at the altitude of Yellowstone, light inhibition by full sunlight was observed only occasionally; the rate of photosynthesis fell progressively with decreasing light, although the most efficient use was at 7–14% of full sunlight. Later, the light intensity over portions of the algal mats was reduced to 18% of full sunlight by installing neutral density glass plates, and changes of chlorophyll content, cell number, and response of photosynthesis to light intensity were determined over the next year. Although the chlorophyll content of the algae at the surface of the mat rose quickly, the chlorophyll content of the mat as a whole rose slowly or not at all; the photosynthetic response of the algal mats to full and reduced sunlight also changed slowly or not at all. Although individual algal cells can adapt rapidly to changes in light, the entire population, because of its existence in compact mats, adapts slowly. At the latitude of Yellowstone there is sufficient light throughout the year to enable algal growth to occur even at temperatures near the upper limit at which blue‐green algae can grow; in Iceland, hot spring algae cannot grow during several winter months. Natural ultraviolet radiation neither inhibited nor stimulated photosynthesis.

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SUMMARY The temperature optima of bacteria occurring at various temperatures along the thermal gradient of a hot spring in Yellowstone Park was studied directly in nature by measuring the rate of incorporation in the dark of [14C]glucose or 14CO2. Bacteria found at environmental temperatures over the range 35‐70° were studied. For each temperature, the optimum for glucose and CO2 incorporation was similar to the environmental temperature.

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The physiology of the bacteria living in Boulder Spring (Yellowstone National Park) at 90 to 93 C was studied with radioactive isotope techniques under conditions approximating natural ones. Cover slips were immersed in the spring; after a fairly even, dense coating of bacteria had developed, these cover slips were incubated with radioactive isotopes under various conditions and then counted in a gas flow or liquid scintillation counter. Uptake of labeled compounds was virtually completely inhibited by formaldehyde, hydrochloric acid, and mercuric bichloride, and inhibition was also found with streptomycin and sodium azide. The water of Boulder Spring contains about 3 ,ug of sulfide per ml. Uptake of labeled compounds occurs only if sulfide or another reduced sulfur compound is present during incubation. The pH optimum for uptake of radioactive compounds by Boulder Spring bacteria is 9.2, a value near that of the natural spring water (8.9). Many experiments with a variety of compounds were performed to determine the temperature optimum for uptake of labeled compounds. The results with all the compounds were generally similar, with broad temperature optima between 80 and 90 C, and with significant uptake in boiling (93 C) but not in superheated water (97 C). The results show that the bacteria of Boulder Spring are able to function at the temperature of their environment, although they function better at temperatures somewhat lower. The fine structure of these bacteria has been studied by allowing bacteria in the spring to colonize glass slides or Mylar strips which were immediately fixed, and the bacteria were then embedded and sectioned. The cell envelope structure of these bacteria is quite different from that of other mesophilic or thermophilic bacteria. There is a very distinct plasma membrane, but no morphologically distinct peptidoglycan layer was seen outside of the plasma membrane. Instead, a rather thick diffuse layer was seen, within which a subunit structure was often distinctly visible, and connections frequently occurred between this outer layer and the plasma membrane. The thick outer layer usually consisted of two parts, the outer part of which was sometimes missing. Within the cells, structures resembling ribosomes were seen, and regions lacking electron density which probably contained deoxyribonucleic acid were also visible.