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The Root Effect – a Structural and Evolutionary Perspective

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The Root Effect – a Structural and Evolutionary Perspective Antarctic Science 19 (2), 271–278 (2007) © Antarctic Science Ltd Printed in the UK DOI: 10.1017/S095410200700034X The Root effect - a structural and evolutionary perspective CINZIA VERDE1*, ALESSANDRO VERGARA2,3, DANIELA GIORDANO1, LELIO MAZZARELLA2,3 and GUIDO DI PRISCO1 1Institute of Protein Biochemistry, CNR, Via Pietro Castellino 111, I-80131 Naples, Italy 2Dipartimento di Chimica, Università degli Studi di Napoli “Federico II”, Complesso Universitario di Monte S. Angelo, Via Cinthia, I-80126, Napoli, Italy 3Istituto di Biostrutture e Bioimmagini, CNR, Via Mezzocannone 6, I-80134 Napoli, Italy *[email protected] Abstract: Haemoglobin carries oxygen from the environment to tissues; in vertebrates, it is contained in specialized cells, called erythrocytes. Over the last century, the study of the chemical properties of this haemoprotein has provided a wealth of information. One of its most important and ancient physiological features is the Root effect, found in many teleost fish (and some amphibians). The Root effect corresponds to an extreme pH sensitivity and can be described as an exaggerated Bohr effect: it dictates to what extent the oxygen tension can be raised in acid-producing tissues. It is likely that the eye choroid rete represents the most ancient anatomical structure associated with the presence of Root effect haemoglobins. This review describes our overall understanding of the molecular properties, biological occurrence, physiological role and evolutionary origin of Root effect haemoglobins. The current knowledge of the structural properties of Root effect haemoglobins is discussed in the light of recent results obtained on the haemoglobins of the cold- adapted notothenioids Trematomus newnesi and T. bernacchii. Received 29 June 2006, accepted 4 September 2006 Key words: Antarctica, evolution, fish, haemoglobin, pH-regulation, structure Introduction (Monod et al. 1965, Perutz et al. 1987). The main concept The biochemistry of oxygen transport in polar fish offers a of the two-state allosteric model of Monod, Wyman and wealth of important indications on physiological Changeux (MWC) was that the Hb molecule can exist only adaptations of the organism to the climatic conditions of the in two quaternary states, corresponding to a low-affinity habitat and the remarkable functional flexibility of structure called T (tense) and a high affinity structure called haemoglobin (Hb) in fish physiology. Hence, studies on R (relaxed) (Monod et al. 1965). According to the MWC Hbs of polar fish are helping to improve the general model, the cooperative O2 binding arises from a shift in the understanding of the extremely complex structure-function population from the T to R structure as binding increases. relationship in this ancient and versatile protein. This model further postulates that the heterotropic effects, The Hb molecule has evolved structural and functional such as the Bohr effect, are ascribed to shifts of the diversity to adapt and modify its features under selective allosteric equilibrium. pressures of all types, but both the predominantly helical Fish Hbs, like other vertebrate Hbs, are tetrameric structure and a large number of amino acid residues are well proteins consisting of two α- and two β-subunits, each of conserved. The primary role of Hb, that of carrying oxygen which contains one O2-binding haem group. These subunits to vertebrate tissues, is probably the origin of its adaptation α β α β are paired in two dimers, 1 1 and 2 2. Within different to widely different environmental conditions. Its specialized species the transport of O2 can be modulated by changes in function imposes severe structural constraints on the the Hb structure and allosteric-ligand concentration (ATP molecule. It is not therefore surprising that only a small for most teleost fish), and by changes in the expression of fraction of the residues of the polypeptide chains have been multiple Hbs likely to display different functional features. allowed to be replaced during evolution. According to the During evolution, complex and sophisticated molecular species-adaptation theory of Perutz (1983), the replacement mechanisms, e.g. modulation by pH, carbon dioxide, of a few key amino acid residues leads to functional organophosphates and temperature, have been developed to modulation. The first protein crystal structures of regulate O2 transport by Hb. myoglobin and Hb provided the basis for understanding the The decreased oxygen affinity of Hb at lower pH values relationship between changes in amino acid sequence and in the physiological range is known as the alkaline Bohr protein structure (Kendrew et al. 1958, Perutz et al. 1965). effect, reviewed by Riggs (1988). In many Hbs from teleost The quaternary structure, assembling the four globin fishes, when the pH is lowered, the O2 affinity decreases to subunits, also provided the classic source of theories and such an extent that Hbs cannot be fully saturated even at structural studies of allosteric conformational transitions very high O2 pressure. In addition, cooperativity is totally 271 Downloaded from https://www.cambridge.org/core. Open University Library, on 05 May 2019 at 04:49:16, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S095410200700034X 272 CINZIA VERDE et al. lost and the O2 capacity of blood undergoes reduction of quaternary transition between R and T conformations. 50% or more of its value at alkaline pH. This feature is Allosteric effectors (organophosphates and protons) may known as the Root effect, reviewed by Brittain (2005). Root modulate these steric effects within both conformations. It effect Hbs are so strongly pH dependent that they are able to is feasible that the Root effect does not completely depend unload a large amount of bound O2 at low pH and against a on the primary structure, but also on how and to what extent pressure gradient. The structural basis of the Root effect has the Hb molecule binds effectors in the T and R states been addressed in many studies. Nevertheless, it remains a (Yonetani et al. 2002). puzzle in several aspects. Originally, it was proposed that As pointed out by Brittain (2005), a detailed definition of inter-subunit salt bridges stabilize the T quaternary determinants for the Root effect requires structural structure, lower the O2 affinity of T relative to R, and are information on R and T states at different pH values. responsible for the Bohr and Root effects (Perutz & Brunori Following this line, we review recent advances in 1982). The ongoing debate on the structural interpretation understanding the structure, function and evolution of the of the Root effect suggests that the classical model is an Root effect in fish Hbs. This review also describes our oversimplified explanation of the pH modulation in fish general knowledge of the structural basis of the Root effect Hbs. in Hbs of Antarctic notothenioids, e.g. T. newnesi Hb C (see The C-terminal residue of His in β146 appears to be below) and T. bernacchii Hb. involved in the Root effect in some Hbs but not in all. Bovichtidae, Pseudaphritidae, Eleginopidae, Nototheniidae, Moreover, a fundamental difference between Root effect Harpagiferidae, Artedidraconidae, Bathydraconidae and Hbs and Hbs with normal Bohr effect is that in the former Channichthyidae are the families of the dominant suborder α β the 1 2 interface remains stable in the T state upon Notothenioidei, thought to have arisen in Antarctica through oxygenation, whereas in the latter the switch to the R state adaptive radiation of the ancestral stock. Channichthyidae occurs. are devoid of Hb, and in the other seven families the Recent studies on the Root effect Hb of the spot concentration of the single major Hb component is highly Leiostomus xanthurus have suggested that the reduced. In contrast, the nototheniid T. newnesi has two destabilization of the R state (and not the stabilization of the functionally distinct major Hbs. The Hb system of T state) at low pH, inducing an R→T transition, may T. newnesi (D’Avino et al. 1994), a semipelagic, active fish, explain the molecular basis for the Root effect is made of Hb C (20–25% of the total), and Hb 1 (70–75%; (Mylvaganam et al. 1996). In the R state of spot Hb, the it has the α chain in common with Hb C) and Hb 2 (5%). central cavity is more narrow than in human Hb (Hb A), due Only Hb C displays pH regulation. Cooperativity of oxygen to the presence of some specific residues in the binding is present in Hb 1 and Hb 2 in the whole β chains. The same features have also been found in the physiological pH range, but is completely lost in Hb C at Antarctic notothenioid Trematomus bernacchii Hb lower pH (a typical feature of Root effect Hbs). This Hb displaying the Root effect (Ito et al. 1995). However, these system can ensure oxygen binding at the gills (via Hb 1) and structural properties by themselves are not sufficient to controlled delivery to tissues (via Hb C) also when active explain the absence of the Root effect in Hb 1 of the other behaviour may produce acidosis. Even if Hb C has not been Antarctic notothenioid, Trematomus newnesi (D’Avino subjected to selection, its expression can nonetheless be et al. 1994) that has all the residues believed to be important activated as a consequence of needs arising from the fish to evoke the Root effect. Interestingly, the sequence-identity life style. High levels of Hb C, conceivably redundant in between these two major cold-adapted Hbs is very high. other notothenioids (which have only traces of Hb C, but Recently, O2-binding experiments under a wide range of rely on the Root-and Bohr-effects of both Hb 1 and Hb 2), medium conditions have brought about a new model of compensate for the lack of regulation of Hb 1 and Hb 2 by allosteric regulation of Hb, the Global Allostery Model, protons and other effectors.
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