Geological history of oxygen

Before photosynthesis evolved, Earth's atmosphere had no free oxygen (O2).[2] Photosynthetic prokaryotic organisms that produced O2 as a waste product lived long before the first build-up of free oxygen in the atmosphere,[3] perhaps as early as 3.5 billion years ago. The oxygen they produced would have been rapidly removed from the oceans by weathering of reducing minerals, most notably iron.[1] This rusting led to the deposition of iron oxide on the ocean floor, forming banded iron formations. Thus, the oceans rusted and turned red. Oxygen only began to persist in the atmosphere in small quantities about 50 million years before the start of the Great Oxygenation Event.[4] This mass oxygenation of the atmosphere resulted in rapid buildup of free oxygen. At current rates of primary production, today's concentration of oxygen could be produced by photosynthetic organisms in 2,000 years.[5] In the absence of plants, the rate of oxygen production by photosynthesis was slower in the Precambrian, and the concentrations of O2 attained were less than 10% of today's and probably fluctuated greatly; oxygen may even have disappeared from the atmosphere again around 1.9 billion years ago.[6] These fluctuations in oxygen concentration had little direct effect on life, with mass extinctions not observed until the appearance of complex life around the start of the Cambrian period, 541 million years ago.[7] The presence of O
2 provided life with new opportunities. Aerobic metabolism is more efficient than anaerobic pathways, and the presence of oxygen created new possibilities for life to explore.[8][9] Since the start of the Cambrian period, atmospheric oxygen concentrations have fluctuated between 15% and 35% of atmospheric volume.[10] The maximum of 35% was reached towards the end of the Carboniferous period (about 300 million years ago), a peak which may have contributed to the large size of insects and amphibians at that time.[9] Whilst human activities, such as the burning of fossil fuels, affect relative carbon dioxide concentrations, their effect on the much larger concentration of oxygen is less significant.[11]

O2 build-up in the Earth's atmosphere. Red and green lines represent the range of the estimates while time is measured in billions of years ago (Ga).
Stage 1 (3.85–2.45 Ga): Practically no O2 in the atmosphere.
Stage 2 (2.45–1.85 Ga): O2 produced, but absorbed in oceans and seabed rock.
Stage 3 (1.85–0.85 Ga): O2 starts to gas out of the oceans, but is absorbed by land surfaces and formation of ozone layer.
Stages 4 and 5 (0.85 Ga–present): O2 sinks filled, the gas accumulates.[1]

Effects on life
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The Great Oxygenation Event had the first major effect on the course of evolution. Due to the rapid buildup of oxygen in the atmosphere, many organisms that didn't rely on oxygen to live died.[9] The concentration of oxygen in the atmosphere is often cited as a possible contributor to large-scale evolutionary phenomena, such as the origin of the multicellular Ediacara biota, the Cambrian explosion, trends in animal body size, and other extinction and diversification events.[9]

The large size of insects and amphibians in the Carboniferous period, when the oxygen concentration in the atmosphere reached 35%, has been attributed to the limiting role of diffusion in these organisms' metabolism.[12] But Haldane's essay[13] points out that it would only apply to insects. However, the biological basis for this correlation is not firm, and many lines of evidence show that oxygen concentration is not size-limiting in modern insects.[9] There is no significant correlation between atmospheric oxygen and maximum body size elsewhere in the geological record.[9] Ecological constraints can better explain the diminutive size of post-Carboniferous dragonflies - for instance, the appearance of flying competitors such as pterosaurs, birds and bats.[9]

Rising oxygen concentrations have been cited as one of several drivers for evolutionary diversification, although the physiological arguments behind such arguments are questionable, and a consistent pattern between oxygen concentrations and the rate of evolution is not clearly evident.[9] The most celebrated link between oxygen and evolution occurs at the end of the last of the Snowball glaciations, where complex multicellular life is first found in the fossil record. Under low oxygen concentrations and before the evolution of nitrogen fixation, biologically-available nitrogen compounds were in limited supply [14] and periodic "nitrogen crises" could render the ocean inhospitable to life.[9] Significant concentrations of oxygen were just one of the prerequisites for the evolution of complex life.[9] Models based on uniformitarian principles (i.e. extrapolating present-day ocean dynamics into deep time) suggest that such a concentration was only reached immediately before metazoa first appeared in the fossil record.[9] Further, anoxic or otherwise chemically "nasty" oceanic conditions that resemble those supposed to inhibit macroscopic life re-occur at intervals through the early Cambrian, and also in the late Cretaceous – with no apparent effect on lifeforms at these times.[9] This might suggest that the geochemical signatures found in ocean sediments reflect the atmosphere in a different way before the Cambrian - perhaps as a result of the fundamentally different mode of nutrient cycling in the absence of planktivory.[7][9]

An oxygen-rich atmosphere can release phosphorus and iron from rock, by weathering, and these elements then become available for sustenance of new species whose metabolisms require these elements as oxides.[2]

References
  1. Holland, H. D. (2006). "The oxygenation of the atmosphere and oceans". Philosophical Transactions of the Royal Society B: Biological Sciences. 361 (1470): 903–915. doi:10.1098/rstb.2006.1838. PMC 1578726. PMID 16754606.
  2. Zimmer, Carl (3 October 2013). "Earth's Oxygen: A Mystery Easy to Take for Granted". New York Times. Retrieved 3 October 2013.
  3. Dutkiewicz, A.; Volk, H.; George, S. C.; Ridley, J.; Buick, R. (2006). "Biomarkers from Huronian oil-bearing fluid inclusions: an uncontaminated record of life before the Great Oxidation Event". Geology. 34 (6): 437. Bibcode:2006Geo....34..437D. doi:10.1130/G22360.1.
  4. Anbar, A.; Duan, Y.; Lyons, T.; Arnold, G.; Kendall, B.; Creaser, R.; Kaufman, A.; Gordon, G.; Scott, C.; Garvin, J.; Buick, R. (2007). "A whiff of oxygen before the great oxidation event?". Science. 317 (5846): 1903–1906. Bibcode:2007Sci...317.1903A. doi:10.1126/science.1140325. PMID 17901330. S2CID 25260892.
  5. Dole, M. (1965). "The Natural History of Oxygen". The Journal of General Physiology. 49 (1): Suppl:Supp5–27. doi:10.1085/jgp.49.1.5. PMC 2195461. PMID 5859927.
  6. Frei, R.; Gaucher, C.; Poulton, S. W.; Canfield, D. E. (2009). "Fluctuations in Precambrian atmospheric oxygenation recorded by chromium isotopes". Nature. 461 (7261): 250–253. Bibcode:2009Natur.461..250F. doi:10.1038/nature08266. PMID 19741707. S2CID 4373201. Lay summary.
  7. Butterfield, N. J. (2007). "Macroevolution and macroecology through deep time". Palaeontology. 50 (1): 41–55. doi:10.1111/j.1475-4983.2006.00613.x. S2CID 59436643.
  8. Freeman, Scott (2005). Biological Science, 2nd. Upper Saddle River, NJ: Pearson – Prentice Hall. pp. 214, 586. ISBN 978-0-13-140941-5.
  9. Butterfield, N. J. (2009). "Oxygen, animals and oceanic ventilation: An alternative view". Geobiology. 7 (1): 1–7. doi:10.1111/j.1472-4669.2009.00188.x. PMID 19200141. S2CID 31074331.
  10. Berner, R. A. (Sep 1999). "Atmospheric oxygen over Phanerozoic time". Proceedings of the National Academy of Sciences of the United States of America. 96 (20): 10955–10957. Bibcode:1999PNAS...9610955B. doi:10.1073/pnas.96.20.10955. ISSN 0027-8424. PMC 34224. PMID 10500106.
  11. Emsley, John (2001). "Oxygen". Nature's Building Blocks: An A-Z Guide to the Elements. Oxford, England, UK: Oxford University Press. pp. 297–304. ISBN 978-0-19-850340-8.
  12. Polet, Delyle (2011). "THE BIGGEST BUGS: AN INVESTIGATION INTO THE FACTORS CONTROLLING THE MAXIMUM SIZE OF INSECTS". 2 (1): 43–46. Cite journal requires |journal= (help)
  13. Haldane, J.B.S., On being the right size, paragraph 7
  14. Navarro-González, Rafaell; McKay, Christopher P.; Nna Mvondo, Delphine (Jul 2001). "A possible nitrogen crisis for Archaean life due to reduced nitrogen fixation by lightning" (PDF). Nature. 412 (5 July 2001): 61–64. Bibcode:2001Natur.412...61N. doi:10.1038/35083537. hdl:10261/8224. PMID 11452304. S2CID 4405370.
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