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Fish Hemoglobins 769 ISSN 0100-879X Review Brazilian Journal of Medical and Biological Research (2007) 40: 769-778 Fish hemoglobins 769 ISSN 0100-879X Review Fish hemoglobins P.C. de Souza and Departamento de Química e Ciências Ambientais, G.O. Bonilla-Rodriguez Instituto de Biociências, Letras e Ciências Exatas, Universidade Estadual Paulista, São José do Rio Preto, SP, Brasil Abstract Correspondence Vertebrate hemoglobin, contained in erythrocytes, is a globular pro- Key words G.O. Bonilla-Rodriguez tein with a quaternary structure composed of 4 globin chains (2 alpha • Hemoglobins Departamento de Química e and 2 beta) and a prosthetic group named heme bound to each one. • Fish Ciências Ambientais Having myoglobin as an ancestor, hemoglobin acquired the capacity • Root effect IBILCE-UNESP to respond to chemical stimuli that modulate its function according to • Bohr effect Rua Cristovão Colombo, 2265 15054-000 São José do Rio Preto, SP tissue requirements for oxygen. Fish are generally submitted to spatial Brasil and temporal O2 variations and have developed anatomical, physi- E-mail: [email protected] ological and biochemical strategies to adapt to the changing environ- mental gas availability. Structurally, most fish hemoglobins are tet- Research supported by FAPESP rameric; however, those from some species such as lamprey and (No. 03/00085-4) to hagfish dissociate, being monomeric when oxygenated and oligo- G.O. Bonilla-Rodriguez. meric when deoxygenated. Fish blood frequently possesses several hemoglobins; the primary origin of this finding lies in the polymor- phism that occurs in the globin loci, an aspect that may occasionally Received October 23, 2006 confer advantages to its carriers or even be a harmless evolutionary Accepted March 9, 2007 remnant. On the other hand, the functional properties exhibit different behaviors, ranging from a total absence of responses to allosteric regulation to drastic ones, such as the Root effect. Introduction to the tissues, where it functions as the final acceptor of electrons originating from oxi- Hemoglobin is surely the most studied of dative catabolic reactions (3). The function all proteins. Indeed, the molecular analysis of hemoglobin seems to be adapted to the of hemoglobin has been the testing ground different metabolic necessities of animals for many contemporary ideas and concepts and to constant environmental changes (4). in biology, particularly the understanding of Among vertebrates, fish are the animals the crystallographic structure and structure- of greatest interest to research on adaptive function relationship of proteins, ligand bind- processes due to their position in the evolu- ing, structural transitions between conform- tionary chain and to their genotypic charac- ers, allosteric interactions, and others (1,2). teristics. These features guarantee enormous The ability of the aerobic metabolism of plasticity in the selection of environments animals to satisfy the demands for oxygen is and the capacity of some species to cope only possible thanks to the role of proteins with different conditions of temperature, pres- such as hemoglobins, contained in red cells sure, salinity, and oxygen availability. They or erythrocytes, which facilitate the dissolu- are also the group of animals with the highest tion of large quantities of gas and transport it number of multiple hemoglobins (3). www.bjournal.com.br Braz J Med Biol Res 40(6) 2007 770 P.C. de Souza and G.O. Bonilla-Rodriguez Numerous studies reporting biochemical globins seem to occur in all organisms and adaptations in fish are available in the litera- tissues, exhibiting a diversity of quaternary ture. According to these reports, fish present structures and a large number of functions a high genotypic plasticity, i.e., their genetic apart from oxygen transportation and stor- material is able to undergo modifications, age, as illustrated by cytoglobins and with its expression varying according to en- neuroglobins (7,8). vironmental changes. The ability of fish to The organization of globin genes has form hybrids, such as Tambacu “created” by been well defined in mammals, birds and crossing a female Tambaqui with a male amphibians. In mammals, the alpha and beta Pacu, is confirmed by the high degree of globin genes are located on different chro- genetic variation detected among the differ- mosomes; in humans, the alpha globin gene ent species, and the existence of species that is located on chromosome 16 and the beta preserve features closely similar to those of globin gene on chromosome 11. In chickens, their ancestors (5). the alpha and beta globin genes are also on The evolution of vertebrate hemoglobins different chromosomes. In amphibians, such has been subjected to many selective restric- as the Xenopus, the genes are on the same tions. The duplication of the ancestral gene chromosome. In teleosts, including the Atlan- that gave origin to the alpha and beta sub- tic salmon, carp and zebrafish, the adult alpha units from a common ancestor occurred at globin gene is adjacently linked to the beta least 450 million years ago, and great changes globin gene and the embryonary globins are in the environment have occurred during completely different from adult globins (9). this period. The evolution of hemoglobins In vertebrates, the hemoglobin molecule was totally neutral and mainly involved se- includes four globin chains that create a lected adaptations associated with physi- stable tetramer (Figure 1) formed by two ological and environmental changes (6). alpha-like and two beta-like chains with 141 and 146 amino acid residues, respectively. The hemoglobin structure Each one contains a heme group, which allows the binding of four oxygen (O2) mol- Hemoglobin is composed by polypep- ecules in a reversible form, according to the tide chains, known as globins, each having a scheme below: prosthetic group called heme, identical in every fish species studied to date. On the Hb + 4O2 = Hb(O2)4 other hand, globins differ from species to deoxy-Hb oxy-Hb species and among isoforms. Remarkably, The main function of hemoglobin is to Figure 1. Hemoglobin molecule transport oxygen from the gas-exchange or- with four globin chains, two al- gans to peripheral tissues. It must be able to pha-like and two beta-like chains bind oxygen strongly but at the same time to each bearing a heme group, re- sponsible for the reversible bind- release it when necessary, depending on the ing of oxygen. The figure was partial pressure of the gas (10). Reversible generated by Pymol 0.99 oxygen binding is possible thanks to the (DeLano Scientific LLC, Palo Alto, CA, USA) using a pdb file heme group, specifically with the participa- (1OUU) from trout hemoglobin, tion of an iron atom in the ferrous form, Fe2+. Oncorhynchus mykiss. For some species of primitive fish such as lampreys and hagfish, reversible hemo- globin dissociation occurs in response to oxygen binding. In the case of lampreys, the Braz J Med Biol Res 40(6) 2007 www.bjournal.com.br Fish hemoglobins 771 oxygenated form is monomeric, undergoing (3). The chemical species capable of causing associations with dimers and tetramers when such alterations are denominated allosteric deoxygenating (11). Figure 2 shows a mono- effectors or agents, and these effectors pref- mer of lamprey hemoglobin. In Myxine erentially bind to one of the conformational glutinosa (hagfish) there are three mono- states, T or R, stabilizing them, and therefore meric hemoglobins in the oxygenated form, they reduce or increase O2 affinity, respec- but when they release O2, there is an associa- tively (13). Binding of the effector may oc- tion to form heterodimers and heterotetramers cur to both canonical or classic states as (12). described first for phosphates binding to the hemoglobins from dromedary (Camelus Allosteric control dromedarius) (14) and the “matrinxã” fish (Brycon cephalus) (15). The heterotropic The quaternary structure guarantees the effectors can induce different sub-states in possibility of interaction among the subunits addition to those previously mentioned, as of hemoglobins and the emergence of new was determined in hemoglobins from the properties, known as allostery. This is im- fish “tamoatá” (Hoplosternum littorale) (16). + portant when the oxygen saturation curve is Both H and CO2 also bind to hemoglo- considered. The curve has a sigmoidal struc- bin, affecting its affinity for oxygen and ture, demonstrating the existence of cooper- subtly changing the three-dimensional struc- + ativity among the globins. In other words, O2 tures. The effect of H on the O2 affinity is binding occurs with progressive facilitation called the Bohr effect. For most hemoglo- between the first and fourth subunits, repre- bins, an increase in H+ concentrations, that senting extremes of low and high affinity for is, a pH decrease, lowers the oxygen affinity the gas, respectively. The property of an O2- (3,17), characterizing the alkaline or normal binding subunit modifying the subsequent Bohr effect. Under these circumstances, he- binding of the same molecule is known as moglobin presents a lower oxygen affinity homotropic allosteric interaction (6). and releases the gas at the tissue level. In The most accepted model explaining the allosteric properties of hemoglobins is the Figure 2. A monomer of lamprey one proposed by Monod et al., in 1965. hemoglobin generated by Pymol 0.99 (DeLano Scientific LLC) According to these investigators, hemoglo- using the 1F5P.pdb file. The fig- bin presents two conformations with differ- ure shows the globin
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