The Root effect is a physiological phenomenon that occurs in

fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...

hemoglobin Hemoglobin (haemoglobin, Hb or Hgb) is a protein containing iron that facilitates the transportation of oxygen in red blood cells. Almost all vertebrates contain hemoglobin, with the sole exception of the fish family Channichthyidae. Hemoglobin ...

, named after its discoverer R. W. Root. It is the phenomenon where an increased

proton A proton is a stable subatomic particle, symbol , Hydron (chemistry), H+, or 1H+ with a positive electric charge of +1 ''e'' (elementary charge). Its mass is slightly less than the mass of a neutron and approximately times the mass of an e ...

carbon dioxide Carbon dioxide is a chemical compound with the chemical formula . It is made up of molecules that each have one carbon atom covalent bond, covalently double bonded to two oxygen atoms. It is found in a gas state at room temperature and at norma ...

concentration (lower pH) lowers hemoglobin's affinity and carrying capacity for

oxygen Oxygen is a chemical element; it has chemical symbol, symbol O and atomic number 8. It is a member of the chalcogen group (periodic table), group in the periodic table, a highly reactivity (chemistry), reactive nonmetal (chemistry), non ...

. The Root effect is to be distinguished from the

Bohr effect The Bohr effect is a phenomenon first described in 1904 by the Danish physiologist Christian Bohr. Hemoglobin's oxygen binding affinity (see oxygen–haemoglobin dissociation curve) is inversely related both to acidity and to the concentration of ...

where only the affinity to oxygen is reduced. Hemoglobins showing the Root effect show a loss of cooperativity at low pH. This results in the Hb-O₂ dissociation curve being shifted downward and not just to the right. At low pH, hemoglobins showing the Root effect don't become fully oxygenated even at oxygen tensions up to 20kPa. This effect allows hemoglobin in fish with

swim bladder The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ (anatomy), organ in bony fish that functions to modulate buoyancy, and thus allowing the fish to stay at desired water depth without having to maintain lift ...

s to unload oxygen into the swim bladder against a high oxygen gradient.Verde, C., A. Vergara, D. Giordano, L. Mazzarella, and G. di Prisco. 2007. The Root effect - a structural and evolutionary perspective. Antarctic Science 19:271-278. The effect is also noted in the choroid rete, the network of

blood vessel Blood vessels are the tubular structures of a circulatory system that transport blood throughout many Animal, animals’ bodies. Blood vessels transport blood cells, nutrients, and oxygen to most of the Tissue (biology), tissues of a Body (bi ...

s which carries oxygen to the

retina The retina (; or retinas) is the innermost, photosensitivity, light-sensitive layer of tissue (biology), tissue of the eye of most vertebrates and some Mollusca, molluscs. The optics of the eye create a focus (optics), focused two-dimensional ...

. In the absence of the Root effect, retia will result in the diffusion of some oxygen directly from the arterial blood to the venous blood, making such systems less effective for the concentration of oxygen. It has also been hypothesized that the loss of affinity is used to provide more oxygen to red muscle during acidotic stress.

References

{{diversity of fish Respiratory physiology