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Bohr effect
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== Experimental discovery == [[File:Bohr effect.png|thumb|291x291px|The original dissociation curves from Bohr's experiments in the first description of the Bohr effect, showing a decrease in oxygen affinity as the partial pressure of carbon dioxide increases. This is also one of the first examples of [[cooperative binding]]. X-axis: oxygen partial pressure in [[mmHg]], Y-axis % [[hemoglobin|oxy-hemoglobin]]. The curves were obtained using whole [[dog]] [[blood]], with the exception of the dashed curve, for which [[horse]] [[blood]] was used.]] In the early 1900s, Christian Bohr was a professor at the [[University of Copenhagen]] in Denmark, already well known for his work in the field of respiratory physiology.<ref name=":5">{{Cite journal|last=Irzhak|first=L. I.|title=Christian Bohr (On the Occasion of the 150th Anniversary of His Birth)|journal=Human Physiology|language=en|volume=31|issue=3|pages=366–368|doi=10.1007/s10747-005-0060-x|issn=0362-1197|year=2005}}</ref> He had spent the last two decades studying the solubility of oxygen, carbon dioxide, and other gases in various liquids,<ref name=":3" /> and had conducted extensive research on haemoglobin and its affinity for oxygen.<ref name=":5" /> In 1903, he began working closely with [[Karl Albert Hasselbalch|Karl Hasselbalch]] and [[August Krogh]], two of his associates at the university, in an attempt to experimentally replicate the work of [[Gustav von Hüfner]], using whole blood instead of haemoglobin solution.<ref name=":2" /> Hüfner had suggested that the oxygen-haemoglobin binding curve was [[Hyperbolic function|hyperbolic]] in shape,<ref>G. Hüfner, "Ueber das Gesetz der Dissociation des Oxyharmoglobins und über einige daran sich knupfenden wichtigen Fragen aus der Biologie," [On the Law of the Dissociation of Oxyharmoglobin, and on some important questions arising from biology]. Arch. Anat. Physiol. (in German) (Physiol. Abtheilung) (1890), 1-27.</ref> but after extensive experimentation, the Copenhagen group determined that the curve was in fact [[Sigmoid function|sigmoidal]]. Furthermore, in the process of plotting out numerous dissociation curves, it soon became apparent that high partial pressures of carbon dioxide caused the curves to shift to the right.<ref name=":3">{{Cite journal |last=Edsall |first=J. T.|jstor=4330576|title=Blood and Hemoglobin: The Evolution of Knowledge of Functional Adaptation in a Biochemical System. Part I: The Adaptation of Chemical Structure to Function in Hemoglobin |journal=Journal of the History of Biology |volume=5 |issue=2 |year=1972 |pages=205–257 |doi=10.1007/bf00346659|pmid=11610121|s2cid=751105}}</ref> Further experimentation while varying the CO<sub>2</sub> concentration quickly provided conclusive evidence, confirming the existence of what would soon become known as the Bohr effect.<ref name=":2" /> === Controversy === There is some more debate over whether Bohr was actually the first to discover the relationship between CO<sub>2</sub> and oxygen affinity, or whether the Russian physiologist {{ill|Bronislav Verigo|ru|Вериго, Бронислав Фортунатович}} beat him to it, allegedly discovering the effect in 1898, six years before Bohr.<ref>{{Cite news|url=http://dic.academic.ru/dic.nsf/medic2/8579|title=Вериго эффект - это... Что такое Вериго эффект?|newspaper=Словари и энциклопедии на Академике|language=ru|trans-title=Verigo effect is... What is the Verigo effect?|access-date=2016-11-08}}</ref> While this has never been proven, Verigo did in fact publish a paper on the haemoglobin-CO<sub>2</sub> relationship in 1892.<ref>B. Werigo, "Zur Frage uber die Wirkung des Sauerstoffs auf die Kohlensaureausscheidung in den Lungen," [The question about the effect of oxygen on the secretion of carbonic acid in the lungs]. Pflügers Arch. ges. Physiol. (in German), 51 (1892), 321-361. </ref> His proposed model was flawed, and Bohr harshly criticized it in his own publications.<ref name=":2" /> Another challenge to Bohr's discovery comes from within his lab. Though Bohr was quick to take full credit, his associate Krogh, who invented the apparatus used to measure gas concentrations in the experiments,<ref>A. Krogh, "Apparat und Methoden zur Bestimmung der Aufnahme von Gasen im Blute bei verschiedenen Spannungen der Gase," [Apparatus and methods for the determination of the absorption of gases in the blood at different tensions of the gases]. Skand. Arch. Physiol. (in German), 16 (1904), 390-401.</ref> maintained throughout his life that he himself had actually been the first to demonstrate the effect. Though there is some evidence to support this, retroactively changing the name of a well-known phenomenon would be extremely impractical, so it remains known as the Bohr effect.<ref name=":3" />
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