Interestingly, the transmembrane helices of GPCRs are frequently tilted with varied tilt and rotation angles that depend not only on the receptor type but also on its activation state. Unfortunately, precise prediction of kinks in the TMHs of GPCRs is still limited as these helical deformations cannot be explained only by the presence of the well-known helix-breakers such as Pro, Ser, Thr or Gly residues.
Most likely these deformations are introduced by tertiary interactions which are difficult to capture without a 3D structure of the receptor Yohannan et al. So far, only members of the rhodopsin-like receptor family have been crystallized.
CXCR1 by Park et al. The above classification of the rhodopsin family is still under discussion as other methods have provided different shapes for its phylogenetic tree Surgand et al. For example, Pele et al. Even if two GPCRs are classified as members of the same subfamily, they can significantly differ in their amino acid composition see Fig. The high sequence diversity inside the rhodopsin family corresponds to the high diversity of kinks and bulges in the TM helices and distinct conformations of loops.
Histograms of sequence identity between members of four branches of rhodopsin-like family of GPCRs. Refinement of crystallization conditions yielded higher resolution data, extending the model to 2. Water molecules play critical roles in the GPCR activation process by stabilizing intramolecular interactions Wikstrom et al. Observed in key structurally sensitive areas of many GPCRs, water molecules along with amino acid side chains can form a signal transmission network extending from the ligand-binding site to the cytoplasmic surface Angel et al.
Despite their low sequence similarities, the overall folds of structurally determined GPCRs are remarkably similar. For all rhodopsin structures, RMSDs for transmembrane regions are within 1. Remarkably, preservation of only a few essential regions of a GPCR is required for activation of its cognate G protein Mirzadegan et al.
Rhodopsin in its inactive, ground state undergoes a series of photointermediate steps upon absorption of a photon and isomerization of its chromophore cis -retinylidene. These photointermediate states exhibit unique absorption maxima and can be isolated by trapping alone or with a G protein-derived peptide analog. Ultimately, the photoisomerized chromophore is hydrolyzed and released from the binding pocket yielding opsin and free all- trans -retinal Jastrzebska et al.
Our laboratory expanded upon this work with the structure which exhibits spectral qualities of Meta II, the activated state Salom et al. For rhodopsin, only the Meta II intermediate is capable of activating G t and it differs chemically from other photo-intermediates only by deprotonation of the Schiff base and uptake of a proton from bulk solvent Salom et al.
By now, the structures of most photo-intermediates have been solved by X-ray and electron crystallographic methods Breitman et al. Interestingly, the structures of those rhodopsin photo-intermediates did not exhibit large-scale movements of entire helices. Rather they showed that photoactivation was accomplished with just small-scale, local changes propagated to the cytoplasmic loops, especially the ends of helices V and VI.
An additional structure of a photoactivated rhodopsin, obtained by regenerating opsin crystals with all- trans -retinal, superposed well with opsin structures and is spectrally indistinguishable from either our photoactivated rhodopsin structure or Meta II in solution Choe et al. Moreover, structures of constitutively active mutants of rhodopsin have also been reported Standfuss et al. When one considers the agonist-bound GPCR structures, it becomes readily apparent that the dynamics of the molecule which make it both a difficult structural target and play a key role in its activation are recapitulated.
The inherent flexibility of GPCRs permits dynamic and conformational changes triggered by only a fraction of the energy derived from ligand binding. Because various agonists can bind to a GPCR leading to varying levels of activity, the small changes induced by agonist binding must account for the differences in the efficacy of such ligands in activating a given G protein Rosenbaum et al. This G protein activation is required for subsequent activation of the cascade of reactions, processes required to advance stepwise signal transduction.
Rather than to photon as in rhodopsin , most GP-CRs respond to molecules called ligands that upon binding to a particular GPCR cause a ligand-specific cellular response. The signal is attenuated by receptor phosphorylation and binding of a capping protein arrestin. In the ternary complex consisting of agonist, receptor and G protein, the affinity of the receptor for the agonist is enhanced and the specificity of the G protein for guanine nucleotides changes in favor of GTP over GDP.
These separate subunits can modulate the activity of different cellular effectors channels, kinases or other enzymes. The active state of a GPCR can be defined as that conformation that couples to and stabilizes a nucleotide-free G protein. Each family comprises various members that often show specific expression patterns. Analysis of cellular signaling processes regulated through G 12 and G 13 has been difficult because specific inhibitors of these G proteins are not available.
The cAMP produced in response to G s activation directly modulates the gating of hyperpolarization-activated, cyclic nucleotide-gated channels and activates protein kinase A PKA. Activation of PDE lowers cytosolic cGMP levels leading to a decreased probability of cGMP-regulated cation channels in the plasma membrane being open, which eventually causes hyperpolarization of photoreceptor cells Arshavsky et al.
Perhaps our most advanced understanding of this activation process is derived from rhodopsin and the visual system. This triggers nucleotide release from G protein and its subsequent activation, processes required to advance visual signal transduction. Most GPCRs respond to molecular signals in the form of ligands which upon binding elicit a ligand-specific cellular response. For most GPCRs, the ligand-binding site coincides with the retinylidene-binding pocket in rhodopsin.
Thus, ligand binding causes similar conformational changes as those triggered by chromophore photoisomerization in rhodopsin, and the remaining molecular mechanisms for signal transduction are similar for all GPCRs. Its molecular envelope is consistent with dimeric rhodopsin molecules together with one G protein heterotrimer, yielding a molar ratio of photoactivated rhodopsin to G protein Jastrzebska et al.
The heteropentameric structure for this complex was obtained from native proteins, both rhodopsin and G protein see also Jastrzebska et al. First, it was surprising that the receptor was in a monomeric state, as single particle analyses of a similar preparation indicated that the receptor exists to some degree in a dimeric form in solution Westfield et al.
But these discrepancies cannot be explained by differences in the inherent structure of either this GPCR as mentioned earlier or its G protein. Future research should elucidate this discrepancy. View along the membrane plane. Together, these constraints fix the position of the nucleotide domain with respect to the membrane. The helical domain is connected to the ras-like domain through two flexible linkers. One cannot say when GDP is released during the formation of the complex.
The question is of great importance because it represents an essential, pharmacologically relevant interaction that can regulate nearly all aspects of eukaryotic cell physiology. Moreover, an atomic-resolution understanding will explain the GPCR functional selectivity, namely the ability of different agonists to elicit distinct downstream effects from a single GPCR.
Photoactivated rhodopsin is specifically phosphorylated by G protein-coupled receptor kinases GRKs Maeda et al. Lefkowitz, as many other investigators, was interested in the mechanism of signal termination by GPCRs.
Non-visual arrestins bind the great majority of GPCRs found in different mammalian species Xiao et al. Visual arrestin shows high selectivity for its cognate receptor rhodopsin, even though several other proteins have been identified to be bound by this arrestin subtype Gurevich et al.
The structures of arrestin Fig. The receptor-binding surface is mainly localized to the side of arrestin containing these two cavities. Several exposed residues in the C- and N-domains of arrestin have been identified as being responsible for GPCR recognition Hanson, ; Vishnivetskiy et al. The two domains are linked together by a polar core of charged residues that form a network of salt bridges which stabilize their relative orientation.
It seems that non-visual arrestins can form functionally different complexes with the same receptor depending on the number of receptor-attached phosphates and their positions. These receptor-attached phosphates play a major role in arrestin recruitment, whereas their positions apparently determine the functional consequences of arrestin binding to the phosphoreceptor Tobin et al. Phosphorylation sites on rhodopsin were well defined a decade ago in vitro and in vivo Ohguro et al. Orange balls indicate regions that change upon GPCR binding but are not directly involved in the interaction with receptor.
Colored residues are important for arrestin stability a salt bridge in blue and red in polar core region or initial recognition of receptor two Lys residues in green. Arrestin-1 preferentially binds to activated and phosphorylated rhodopsin. It also specifically binds inactive phosphorylated and active non-phosphorylated forms, but with much lower affinition.
This observation was the first indication that arrestin-1 can recognize activation and phosphorylation of rhodopsin independently of each other. However, when arrestin-1 encounters phosphorylated photoactivated rhodopsin, the engagement of both primary sites allows arrestin to switch into the high-affinity rhodopsin-binding state, bringing additional arrestin elements into contact with rhodopsin.
However, there is no large relative motion of arrestin domains during complex formation, but a concerted movement of multiple flexible loops most probably helps arrestin mold itself onto the receptor Kim et al. The stoichiometry of rhodopsin complexes with its cognate proteins is a matter of debate. Historically, one-to-one binding was usually assumed Hanson et al.
However, it has also been proposed by Palczewski that a single arrestin molecule could accommodate two receptors Liang et al. Monomeric activated and phosphorylated rhodopsin in nanodiscs can bind arrestin Tsukamoto et al. At the same time, it has been reported that although arrestin requires at least a single phosphorylated photoactivated rhodopsin to bind to the membrane, a single arrestin can actually interact with a pair of receptors composed of two different photo-intermediate states.
The binding stoichiometry depends on the percentage of active receptors Sommer et al. From a physiological standpoint, the different binding modes of arrestin correspond well to the functional needs of the cell at different light intensities. In the single-photon range, arrestin binds monomeric phosphorylated photoactivated rhodopsin to quench signaling. But as the lighting level increases and photoactivates more rhodopsin, arrestin also binds dimeric photoactivated rhodopsin with only one phosphorylated protomer.
Further studies have revealed that differentiated binding preferences of the two domains of arrestin allow it to accommodate the different functional forms of phosphorylated rhodopsin Sommer et al. A general hypothesis is formed that the N-domain of arrestin mediates binding to agonist-activated receptor, whereas the less specific C-domain may serve various functions depending on the requirements of the biological system.
The clathrin-coated pit is pinched off from the plasma membrane by the motor protein dynamin, causing the desensitized receptor to enter an endosomal pool. Arrestin-2 is found in both the cytoplasm and the nucleus, whereas arrestin-3 is localized only in the cytoplasm.
Class B receptors, such as angiotensin AT1a receptor, neurotensin receptor 1 and vasopressin V2 receptor, bind arrestin-2 and -3 with equal affinity and their interaction remains intact during internalization. Receptors targeted for degradation traffic to lysosomes and are enzymatically degraded, whereas receptors for recycling traffic to acidified vesicles where they are de-phosphorylated and recycled back to the plasma membrane Tan et al.
Here they serve as adaptor or scaffold molecules that bring crucial molecular components of specific signaling pathways in close proximity to an activated GPCR. The group of Lefkowitz first found that arrestin-2 is complexed with the tyrosine kinase c-Src. In many cases, signals transmitted by arrestin binding are demonstrably independent of heterotrimeric G protein activation.
This observation has triggered the concept of biased agonists, pathway-selective ligands that activate only a subset of the GPCR signaling repertoire Kenakin, Thus, balanced ligands stabilize the conformations that are competent for signaling to all downstream pathways, whereas biased ligands stabilize only those conformations that are capable of promoting a subset of signaling effects Rajagopal et al. The latter ligands could more selectively target beneficial signaling and even block or negate detrimental or unwanted actions of full receptor activation e.
Distinct active conformations of receptor are coupled to different active conformation of arrestin which govern different functional outcomes. Treatment with GPCR agonists can be limited by the development of tachyphylaxis, a decrease in responsiveness to the same dose of a drug, together with tolerance, whereby higher drug doses are required to obtain the same effect.
CCR5-tropic viruses are the predominant species in the early stages of infection and there has been a significant interest in targeting this receptor for the treatment of HIV infection. Indeed, modified CCR5 ligands promote receptor internalization. They also function as ligand-regulated scaffolds that recruit functionally diverse proteins to GPCRs to confer novel signaling properties.
Arrestin-dependent signals are involved in different processes in vivo , such as cell migration, neurotransmission, cardiac muscle contraction, and apoptosis. These signals can be initiated or antagonized independently of G protein activation. Chemokine receptors CRs are one of the most interesting families of GPCRs due to their key role in a number of diseases that affect millions of people worldwide.
Chemokines are small chemotactic cytokines that regulate the trafficking of immune cells by binding to cell surface chemokine receptors CRs. About 50 human chemokines that interact with 22 different receptors have been identified to date.
Disregulated expression of chemokines and their receptors has been implicated in the development of many human diseases see Table 1. As a result, considerable effort has been made to solve the three-dimensional structure of chemokine receptors and to develop drugs to modulate their activities.
By the end of we have learned the connection between a disease and target protein for at least 15 CRs, developed agonists and antagonists for at least 10 different CRs and solved the 3D structures of two chemokine receptors, CXCR4 and CXCR1 see Fig.
We also present computational approaches to model CR structures and perform rational drug design. Top, view along the membrane plane, bottom, from the extracellular side. Major diseases linked to chemokine receptors adopted from Allen et al. The largest subfamily of CRs is named after their ability to bind CC chemokines — a subfamily of chemokines with four or six Cys residues forming two or three di-sulfide bonds, with two conserved Cys residues always forming a CC motif.
Members of this subfamily share substantial homology with the exception of CCR CCR1 was the first CC chemokine receptor identified. It shares a Whereas CCR1 has been implicated in multiple sclerosis, rheumatoid arthritis, psoriasis, transplant rejection, cancer and kidney disease, CCR3 has been connected only to asthma and allergic rhinitis. The first antagonists of CCR1 were reported by Hesselgesser et al.
Some reported drug candidates are potent antagonists with reported K i values of around 1 nM, e. BX Liang et al. A few of the most potent and selective CCR1 inhibitors have progressed to clinical development, but despite their high therapeutic potential none have yet passed clinical trials Gladue et al. The story of CCR3 antagonists is similar, although the first chemical compound showing high activity against it SB was reported later White et al.
As of the end of more than thirty different compounds interacting with CCR3 derived from at least ten different scaffolds have been identified, some with high CCR3 affinity EC 50 values of 1—5 nM Naya et al. More dual inhibitors have been reported Dhanak et al.
Though some of those inhibitors were tested in patients with various diseases, none have yet succeeded. They also interact with some of the same chemokines and both have been linked to the same immunologic and cardiovascular diseases Zhao, The first CCR2 antagonists were described in the literature in , six years after the successful cloning of this receptor Forbes et al. By the end of several pharmaceutical companies have disclosed more than fifty with different molecular scaffolds including piperidine, spiropiperidine, aminopyrrolidine, compounds with bisubstituted cyclohexane groups, and others.
Some of the optimized ligands evidenced CCR2 inhibition in the nanomolar range, with the lowest IC 50 subnanomolar value reported by Teijin company for one of their optimized homopiperazine derivatives Moree et al. This compound bound to the receptor at nanomolar concentrations and inhibited HIV cellular entry in vitro.
Unfortunately, it also exhibited a poor oral bioavailability, but still was used as a CCR5 inhibitor model by other pharmaceutical companies. In the last ten years at least forty different CCR5 antagonists have been identified and, due to the importance of AIDS, many were optimized to display inhibitory efficacy at nanomolar or subnanomolar concentrations. CXCR4 is one of the two chemokine receptors together with CCR5 used by HIV to enter human cells, a finding that has greatly accelerated structural research aimed at this protein Oberlin et al.
Moreover, the relatively large amount of data for this receptor provided insights into the binding modes of a number of antagonists Vabeno et al. The most striking differences were the positions and rotations of helices 1, 2 and 6 that resulted in a much looser packing of all helices, and also the lengths of helices 5 and 7.
Surprisingly, the binding cavity of CXCR4 also was larger, more open and located much closer to the extracellular surface than that of other known GPCRs. This investigation not only provides invaluable information about the structure of CXCR4 and possibly other CR-family members, but it also serves as a platform for rational drug design, contributing to understanding of the HIV-1antagonists entry process along with the elucidated structures of native HIV-1 gp trimers Liu et al.
The first dual antagonist for this pair of proteins was shown to inhibit acute and chronic models of arthritis in the rabbit Podolin et al. Two such dual antagonists have passed all clinical trials and are now marketed — reparixin which attenuates inflammatory responses and promotes recovery of function after spinal cord injury Gorio et al.
CXCR1 is also the second chemokine receptor with a known three-dimensional structure. In its structure was obtained by using rotationally aligned solid-state NMR Park et al. That novel method involved the use of NMR for the first time in GPCR structural studies and provided important information about this receptor in its natural phospholipid bilayer environment.
A comparison of its structure with that of CXCR4 Four charged residues in the helical region of CXCR1 form a polar cluster in the ligand-binding site of this receptor. They most likely are important for ligand binding, similar to the three polar residues found in CXCR4.
Chemokine receptors are a family of GPCRs that have always attracted much attention from researchers and health professionals. Immediately after their recognition as potential targets in various diseases, these proteins became the focus of numerous programs run by pharmaceutical companies. Of these recent innovations, the possibility of obtaining three-dimensional structures by using crystallography or NMR was one of the most important. There are still many questions about the structure of other CRs, their probable oligomerization and dynamics in human cells, but our knowledge of this family of proteins is now increasing exponentially.
The GPCR research will further shape the field of pharmacology and medicine ne a functional channel mediating activation in the decades ahead. Thus it will be challenging but necessary to determine high resolution structures of native GPCRs alone and in complexes with their G protein, receptor kinase and arrestin Jastrzebska et al. Moreover, GPCRs and G proteins are critically altered by several post-translational modifications which should be taken into account in structural studies.
In addition to these modifications, water molecules have been shown to play a key role in all proteins, including transmembrane ones. It will be essential to determine the exact location of water molecules in these receptors and their re-arrangements during GPCR activation.
Understanding the energy landscape of GPCRs corresponding to their folding pathway and activation is highly anticipated. Identification of key residues responsible for folding and membrane insertion is needed to explain the etiology of many human diseases associated with receptor mutations. Moreover, GPCRs are not monomeric — they have a propensity to interact not only with each other but also with other transmembrane proteins. Understanding the structural complementarities of GPCR homo- and hetero-oligomerization offers several novel pharmacological opportunities to explore the high specificity of these interactions.
A large number of GPCRs communicate with only a limited number of G proteins and even a smaller number of effectors such as enzymes and channels. It will be important to determine the intersections of different relevant GPCR signaling pathways in native tissues of interest. Modern 3D structural electron microscopy cryo-electron microscopy and tomography and hybrid microscopic techniques will facilitate obtaining high resolution structures of signaling complexes between GPCRs, their cognate G proteins and effector molecules in native tissues.
Such studies can determine how the GPCR signaling complexes are compartmentalized within cells to evoke their local effects. We are confident that the discovery of mutations responsible for genetic diseases will be dramatically accelerated due to the DNA and RNA sequencing of whole personal genomes as well as tissue transcriptomes. This new methodology will enable evaluation of the impact of mutations and polymorphisms affecting GPCR expression levels on human conditions.
Hence, GPCRs will certainly play a central role in drug research in the foreseeable future. Hord Professor of Pharmacology. National Center for Biotechnology Information , U. Acta Biochim Pol. Author manuscript; available in PMC Feb Author information Copyright and License information Disclaimer. Copyright notice. The publisher's final edited version of this article is available free at Acta Biochim Pol. See other articles in PMC that cite the published article.
Abstract The years and witnessed milestones in current understanding of G protein-coupled receptor GPCR structural biology. Open in a separate window. Figure 1. Figure 2. Figure 3. Figure 4. Figure 5. Crystal structure of arrestin PDB id: 1CF1 with characteristic elements indicated Orange balls indicate regions that change upon GPCR binding but are not directly involved in the interaction with receptor.
Figure 6. Figure 7. Figure 8. Table 1 Major diseases linked to chemokine receptors adopted from Allen et al. Chemokine: receptor structure, interactions, and antagonism. Annu Rev Immunol. Conserved waters mediate structural and functional activation of family A rhodopsin-like G protein-coupled receptors.
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The case of the Maramures County, Romania. Concas, Sisinnio : Highway capital expenditures and induced vehicle travel. Conrad, Daren A. Proceedings : pp. Cooke, Edgar F. User Manual 2. User Manual 1. Corsini, Lorenzo and Spataro, Luca : Savings for retirement under liquidity constraints: a note. Published in: Procedia Economics and Finance 1 , Vol. Coulombe, Harold and Wodon, Quentin : Benefit incidence of public health spending for public and faith-inspired health facilities in Ghana.
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Forthcoming in: Review of New Political Economy. Dai, Darong : Learning Nash Equilibria. Published in: Economic Research Guardian , Vol. Dai, Darong : On the existence and stability of Pareto optimal endogenous matching with fairness. Dai, Darong and Shen, Kunrong : A new stationary game equilibrium induced by stochastic group evolution and rational Individual choice. Dang, Duc Anh : Cooperation makes beliefs: climate variation and sources of social trust in Vietnam.
Dang, Duc Anh : On the sources of risk preferences in rural Vietnam. Dang, Jianwei and Motohashi, Kazuyuki : Patent value and liquidity: evidence from patent-collateralized loans in China. Das, Debasish Kumar : Determinants of current account imbalances in the global economy: A dynamic panel analysis.
Das, Debasish Kumar and Dutta, Champa Bati : Global financial crisis and foreign development assistance shocks in least developing countries. Published in: Modern Economy , Vol. Das, Subhendu : Moneyless Economy. Davutyan, Nurhan and Bilsel, Murat and Tarcan, Menderes : Risk-Adjusted Mortality, varieties of congestion and patient satisfaction in Turkish provincial general hospitals. De Borger, Bruno and Fosgerau, Mogens : Information provision by regulated public transport companies.
De Koning, Kees : Pension savings and economic growth. The U. De Koning, Kees : When capitalism no longer works - a profit warning. De Koning, Kees : The savings paradox or managing financial, economic or financial risks. De Koning, Kees : The world's dream: economic growth : the balance sheet approach. Network analysis of international co-inventions. De Prato, Giuditta and Nepelski, Daniel : A framework for assessing innovation collaboration partners and its application to India.
Published in: Lifelong Learning in Europe , Vol. De Vijlder, Nicolas : A macroeconomic analysis of the land market in the count of Flanders and the duchy of Brabant. Published in: Journal of Conflict Resolution. Decker, Stephanie : The silence of the archive: post-colonialism and the practice of historical reconstruction from archival evidence. Deepak, Shah : Financial and economic crisis: implications for agricultural sector in India. Deepak, Shah : Implementation of national food security mission for pulse crops in Maharashtra.
Deepak, Shah : Special economic zones in India: investment, trade, employment generation and impact assessment. Published in: Managerial Finance , Vol. Delis, Manthos D and Tsionas, Efthymios : A new method to estimate the risk of financial intermediaries.
Delisle, R. Jared and Lee, Bong Soo and Mauck, Nathan : The dynamic relation between short sellers, option traders, and aggregate returns. Demachi, Kazue : The effect of crude oil price change and volatility on Nigerian economy. Demichelis, Stefano : Evolution towards efficient coordination in repeated games, preliminary version. Demir, Firat and Dahi, Omar S. Vanessa : On the epidemic of financial crises. Dennis, Wesselbaum : Stochastic Volatility in the U.
Labor Market. Denny, Eleanor and Keane, Andrew : A smart integrated network for an offshore island. Dentcheva, Darinka and Ruszczynski, Andrzej : Common mathematical foundations of expected utility and dual utility theories. Desiderio, Saul and Chen, Siyan : Long-run consequences of debt in a stock-flow consistent network economy. Destefanis, Sergio : Alcune considerazioni sul mercato del lavoro italiano alla luce della ricostruzione delle serie storiche territoriali per il mercato del lavoro, Devereux, Michael B.
Published in: Canadian Journal of Economics , Vol. Dey, Jaya and Tsai, Yi-Chan : Explaining the durable goods co-movement puzzle with non-separable preferences: a bayesian approach. Dezhina, Irina and Simachev, Yuri : Partnering universities and companies in Russia: effects of new government initiative.
Di Berardino, Claudio : La distribuzione spaziale della migrazione interna in Polonia: un'analisi delle differenze tra aree urbane e rurali. Di Gaetano, Luigi : A Model of corporate donations to open source under hardware—software complementarity. Diakantoni, Antonia and Escaith, Hubert : Reassessing effective protection rates in a trade in tasks perspective: Evolution of trade policy in "Factory Asia".
Diallo, Ibrahima Amadou : The effects of real exchange rate volatility on productivity growth. Diamondopoulos, John : To what extent are financial crises comparable and thus predictable? Dietrich, Franz : Judgment aggregation and the discursive dilemma. Dietrich, Franz and List, Christian : Mentalism versus behaviourism in economics: a philosophy-of-science perspective. Dietrich, Franz and List, Christian : Reasons for prior belief in bayesian epistemology.
Dietrich, Franz and Spiekermann, Kai : Independent opinions? On the causal foundations of belief formation and jury theorems. Dincecco, Mark and Katz, Gabriel : State capacity and long-run performance. Forthcoming in: Journal of Economic Growth. Dissanayake, D. Dizioli, Allan and Pinheiro, Roberto B. Dmitriev, Mikhail and Hoddenbagh, Jonathan : The optimal design of a fiscal union.
Dobre, Ana Maria : Intangible assets as a source of competitiveness in the post-crisis economy. The role of trademarks. Published in: Journal of Economic Structures , Vol. Doko Tchatoka, Firmin and Dufour, Jean-Marie : Identification-robust inference for endogeneity parameters in linear structural models. Published in: International Business Research , Vol. Published in: Frontiers of Economics in China , Vol. Doran, Justin : Are different forms of innovation complements or substitutes?
Doran, Justin : An analysis of the interdependence of demographic factors, labour effort and economic growth in Ireland. Doran, Justin and Fingleton, Bernard : Economic shocks and growth: spatio-temporal perspectives on Europe's economies in a time of crisis. Forthcoming in: Papers in Regional Science. Published in: Industry and Innovation , Vol. Doretti, Marco : Modelli di scoring per il rischio paese. Published in: Economy Transdisciplinarity Cognition , Vol. XV, No.
Dorsch, Michael and Maarek, Paul : Inefficient predation, information, and contagious institutional change. Douillet, Mathilde : Trade and agricultural policies in Malawi: Not all policy reform is equally good for the poor. Douillet, Mathilde : Trade policies and agricultural exports of Sub-Saharan African countries: Some stylized facts and perspectives. Dr Bhawna, Rathore : Impact of demographic features on economic development of India from - Drakopoulos, Stavros A.
Drichoutis, Andreas and Lusk, Jayson : Judging statistical models of individual decision making under risk using in- and out-of-sample criteria. Drichoutis, Andreas and Lusk, Jayson : What can multiple price lists really tell us about risk preferences? Drichoutis, Andreas and Nayga, Rodolfo : Do risk and time preferences have biological roots? Du, Chuang : Solving payoff sets of perfect public equilibria: an example.
Published in: Southwestern Economic Review , Vol. Duddy, Conal : Condorcet's principle and the strong no-show paradoxes. Duffy, Sean and Smith, John : Cognitive load in the multi-player prisoner's dilemma game. Duffy, Sean and Smith, John : Cognitive load in the multi-player prisoner's dilemma game: Are there brains in games?
Dumitriu, Ramona and Stefanescu, Razvan and Nistor, Costel : Reactions of the capital markets to the shocks before and during the global crisis. Dutcher, E. An Experiment. Dutta, Mousumi and Husain, Zakir : Use of hospital services and socio-economic status in urban India: Does health insurance ensure equitable outcomes? Evidence from Syndicated Lending. May Eagle, David M.
MS Ecchia, Stefania : Mercati informali del credito agrario nella Palestina di fine Impero Ottomano: un'analisi dell'evoluzione dei contratti bay-wafa, salam e muzaraah nel distretto di Haifa Eckel, Catherine and Johnson, Cathleen and Montmarquette, Claude : Human capital investment by the poor: Informing policy with laboratory experiments. Forthcoming in: Journal of Economic Behavior and Organization.
Edwards, Jeffrey A. Published in: Natural Resources Journal , Vol. Eichengreen, Barry and Gupta, Poonam : The global financial crisis and indian banks: survival of the fittest? Ekong, Christopher N. Vol 2, No. No 4 October : pp. Elshaer, Ibrahim : What is the Meaning of Quality? Emara, Noha : Inflation volatility, financial institutions and sovereign debt rating. Ercolano, Salvatore and Gaeta, Giuseppe Lucio and Romano, Oriana : Environmental fiscal reform and willingness to pay for the environment: an empirical analysis on European micro data.
Erten, Irem and Okay, Nesrin : Re-examining Turkey's trade deficit with structural breaks: Evidence from Erten, Irem and Tuncel, Murat B. Published in: Current Issues in Tourism , Vol. Escobari, Diego and Lee, Sang-Yeob : Demand shifting across flights and airports in a spatial competition model. Forthcoming in: Letters in Spatial and Resource Sciences. Espinosa, Miguel and Rondon, Carlos and Romero, Mauricio : The use of mathematics in economics and its effect on a scholar's academic career.
Estrada, Fernando : Asymmetric information and financial markets. Estrada, Fernando : Brief note about the dilemmas of public election. Estrada, Fernando : Economy and power to tax. Estrada, Fernando : Estado y poder fiscal. IV, No. Estrada, Fernando : Heuristic in the economic: a note on Robert Nozick. Estrada, Fernando : Transaction costs, externalities and innovation. Estrada, Fernando : The logic of violence in the civil war: the economics perspective.
Ettah, Bassey E. Evangelista, Rui and Santos, Daniel : The treatment of housing co-operatives in a house price index. Ewa, Lechman : Cross national technology convergence. An empirical study for the period Ewa, Lechman : Social development — a multidimensional approach to social development analysis. Country level evidence for year Published in: in Onafowokan O. Oluyombo eds. Fan, Haichao and Lai, Edwin L.
Fan, Jianqing and Liao, Yuan : Endogeneity in ultrahigh dimension. Farzanegan, Mohammad Reza : Does the Iranian oil supply matter for the oil prices? Fauceglia, Dario and Shingal, Anirudh and Wermelinger, Martin : "Natural hedging" of exchange rate risk: The role of imported input prices. Favaro, Donata and Ninka, Eniel and Turvani, Margherita : Productivity in innovation: the role of inventor connections and mobility. Fe, Eduardo : Efficient estimation in regression discontinuity designs via asymmetric kernels.
Fe, Eduardo and Hollingsworth, Bruce : Estimating the eect of retirement on mental health via panel discontinuity designs. Federici, Daniela and Parisi, Valentino : Corporate taxation and exports. Fedotenkov, Igor and van Groezen, Bas and Meijdam, Lex : International trade with pensions and demographic shocks.
Feige, Edgar L. Forthcoming in: Bundesbank Conference Volume. Fenske, James : Does land abundance explain African institutions? Fenske, James : Ecology, trade and states in pre-colonial Africa. Fenske, James : "Rubber will not keep in this country": Failed development in Benin, Fenske, James : Trees, tenure and conflict: Rubber in colonial Benin. Ferreira Lima, Luis Cristovao : The determinants of the academic outcome: a Bayesian approach using a sample of economics students from the University of Brasilia, Brazil.
Filippova, Irina and Sumcov, Victor : Social responsibility as an obligatory element of the institutional system. Forthcoming in: Filippova, Irina H. XIV, No. Fioriti, Linda and Marchini, Andrea and Diotallevi, Francesco and Pampanini, Rossella : Obesity epidemic: the role of retailing sector in promoting fruit and vegetable consumption.
Fischer, Justina A. Fischer, Justina AV : Globalization and political trust. ISSN No. Forte, Antonio : Italy after the crisis: a case of recoveryless credit growth. Forte, Antonio and Cepparulo, Alessandra : Microeconomic determinants of losses in financial institutions during the crisis. Fortun Vargas, Jonathan M. Fosgerau, Mogens and Fukuda, Daisuke : Valuing travel time variability: Characteristics of the travel time distribution on an urban road. Fosgerau, Mogens and Small, Kenneth E.
Published in: Journal of Urban Economics , Vol. Cambiare paradigma per salvare il servizio: il caso ligure. Published in: Territori No. Franco, Daniel and LoFazio, Aurelio : Da governo a governance: il possibile ruolo della partecipazione.
Francq, Christian and Meintanis, Simos : Fourier--type estimation of the power garch model with stable--paretian innovations. Francq, Christian and Wintenberger, Olivier and Zakoian, Jean-Michel : Garch models without positivity constraints: exponential or log garch?
Fratini, Saverio M. Friberg, Richard and Huse, Cristian : How to use demand systems to evaluate risky projects, with an application to automobile production. Friedrich, T. Fry, John : Exogenous and endogenous crashes as phase transitions in complex financial systems. Fujii, Hidemichi and Iwata, Kazuyuki and Kaneko, Shinji and Managi, Shunsuke : Corporate environmental and economic performances of Japanese manufacturing firms: Empirical study for sustainable development.
Forthcoming in: Business Strategy and the Environment. Fujii, Hidemichi and Managi, Shunsuke : Decomposition of toxic chemical substance management in three U. Forthcoming in: Journal of Industrial Ecology. Fujii, Hidemichi and Managi, Shunsuke : Productive inefficiency analysis and toxic chemical substances in US and Japanese manufacturing sectors.
Fujisaki, Seiya : Taylor rules and equilibrium determinacy in a two-country model with non-traded goods. Fullerton, Thomas M. Published in: Empirical Economics Letters , Vol. Doyle and Walke, Adam G. Summer June : pp. Fulli-Lemaire, Nicolas and Palidda, Ernesto : Swapping headline for core inflation: an asset liability management approach. Furukawa, Yuichi : Perpetual leapfrogging in international competition. Gabrielsen, A. Gabrisch, Hubert and Orlowski, Lucjan T. Gagen, Michael : Using strong isomorphisms to construct game strategy spaces.
Gagnin, Cristiano and Havas, Attila and Saritas, Ozcan : Future-oriented technology analysis: Its potential to address disruptive transformations. Ekonomiczne nauki August : pp. Galimberti, Jaqueson K. Published in: Research in Experimental Economics , Vol. Gallurt, Jesus and Pombo, Pablo and Ramirez, Jesus and Molina, Horacio : La asimetria de la informacion en la crisis financiera, el racionamiento del credito y la garantia como mecanismo simbiotico del sistema.
Garcia-Martinez, Jose A. Forthcoming in: Psychologia Spoleczna. Gavazza, Alessandro and Lizzeri, Alessandro and Rokestkiy, Nikita : A quantitative analysis of the used-car market. Forthcoming in: Journal of Industrial Economics. Forthcoming in: Economics of Transportation. Gechert, Sebastian : The multiplier principle, credit-money and time.
Gemignani, Regina and Tsimpo, Clarence and Wodon, Quentin : Making quality care available for the poor: faith-inspired health facilities in Burkina Faso. Gemignani, Regina and Wodon, Quentin : How do households choose between health providers?
Published in: Erudition , Vol. Georgiadis, Georgios : The panel conditionally homogenous vectorautoregressive model. Ghafele, Roya : Financing University Research. Ghafele, Roya and D. Ghafele, Roya and Gibert, Benjamin : Promoting intellectual property monetization in developing countries : a review of issues and strategies to support knowledge-driven growth. WPS July : pp. Ghassan, Hassan B. Ghiurca, Ana-Andreea and Lamasanu, Andreea and Mihai, Florin-Constantin : Environmental education in rural areas - a real support for sustainable development,.
Evidence from Indian firm-level data. Published in: Journal of International Entrepreneurship , Vol. Ghosh, Saibal : Foreign banks in India: liabilities or assets? Published in: Economic Papers , Vol. Gill, David and Prowse, Victoria : Cognitive ability and learning to play equilibrium: A level-k analysis.
Gill, David and Prowse, Victoria : Gender differences and dynamics in competition: the role of luck. Gill, David and Stone, Rebecca : Desert and inequity aversion in teams. Gimeno, Ricardo and Gonzalez, Clara I. Giocoli, Nicola : British economists on competition policy Giocoli, Nicola : Old lady charm: explaining the persistent appeal of Chicago antitrust.
Girardi, Daniele : Do financial investors affect the price of wheat? Girardi, Daniele : Il settore delle costruzioni in Italia tra crisi e trasformazione. Girardi, Daniele : A brief essay on the financialization of agricultural commodity markets. Giuranno, Michele G. Givens, Gregory and Salemi, Michael : Inferring monetary policy objectives with a partially observed state. Gluschenko, Konstantin : Myths about Beta-Convergence.
Gnidchenko, Andrey A. Goagara, Daniel and Vasilescu, Laura and Nitu, Cornelia : National and international exigencies on increasing the quality of accounting information generated by evaluation by comparison. Evidence from Malaysia. Gojinetchi, Sergiu : Consumer protection in the shadow of the global financial crisis - a study on the way forward of consumer protection in European Union, Republic of Moldova and P.
XX, No. XX : X-X. Published in: Przeglad Organizacji , Vol. Gong, Liutang and Zou, Heng-fu : Risk-taking, fiscal policies, asset pricing, and stochastic growth with the spirit of capitalism. Gong, Ting and Wu, Alfred M. Evidence from China. Published in: European Economic Review , Vol. The impact of aid on Latin American inequality. Goodwin, Barry K. Gorbanev, Mikhail : Sunspots, unemployment, and recessions, or Can the solar activity cycle shape the business cycle?
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Published in: India Development Report June : pp. Grady, Patrick : The parent and grandparent immigration program in Canada: costs and proposed changes. Graefe, Andreas and Armstrong, J. Forthcoming in: Journal of Behavioral Decision Making. Grech, Aaron George : Evaluating the possible impact of pension reforms on future living standards in Europe.
Greco, Esteban M. Greco, Giulio : Governance codes and types of issuer. An empirical research on a global sample. Greco, Giulio : Ownership structures, corporate governance and earnings management in the European Oil Industry. Green, Kesten C. Scott : Evidence on the effects of mandatory disclaimers in advertising. Green, Mitchell R. Published in: Oeconomicus , Vol.
Groll, Thomas and Ellis, Christopher J. Groth, Christian and Wendner, Ronald : Embodied learning by investing and speed of convergence. Grubel, Herbert and Grady, Patrick : Fiscal transfers to immigrants in Canada: responding to critics and a revised estimate. Grydaki, Maria and Bezemer, Dirk J. Fall : pp. Guerrazzi, Marco : Asymptotic relations in Cournot's game. Guerrazzi, Marco : On involuntary unemployment: notes on efficiency-wage competition.
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Activation of G-protein-coupled receptors correlates with the formation of a continuous internal water pathway. Endogenous fluorophores enable two-photon imaging of the primate eye. Noninvasive two-photon microscopy imaging of mouse retina and retinal pigment epithelium through the pupil of the eye.
Nat Med. Identification and characterization of novel inhibitors of Mammalian aspartyl aminopeptidase. Time-resolved fluorescence spectroscopy measures clustering and mobility of a G protein-coupled receptor opsin in live cell membranes. J Am Chem Soc. STRA6 is critical for cellular vitamin A uptake and homeostasis. DICER1 is essential for survival of postmitotic rod photoreceptor cells in mice.
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Two carotenoid oxygenases contribute to mammalian provitamin A metabolism. Photoreceptor cells are major contributors to diabetes-induced oxidative stress and local inflammation in the retina. Cellular retinaldehyde binding protein-different binding modes and micro-solvation patterns for high-affinity 9-cis- and cis-retinal substrates.
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A hybrid structural approach to analyze ligand binding by the serotonin type 4 receptor 5-HT4. Mol Cell Proteomics. QLT, a 9-cis-retinal analog, is well-tolerated by retinas of mice with impaired visual cycles. Asymmetry of the rhodopsin dimer in complex with transducin. Retinal degeneration in animal models with a defective visual cycle. Drug Discov Today Dis Models. Maeda A, Palczewski K. Expression of mammalian G protein-coupled receptors in Caenorhabditis elegans.
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Melanopsin is highly resistant to light and chemical bleaching in vivo. Substrate-induced changes in the dynamics of rhodopsin kinase G protein-coupled receptor kinase 1. The physiological impact of microRNA gene regulation in the retina. Cell Mol Life Sci. Insights into substrate specificity and metal activation of mammalian tetrahedral aspartyl aminopeptidase. Imaging of protein crystals with two-photon microscopy.
Retinal cone and rod photoreceptor cells exhibit differential susceptibility to light-induced damage. J Neurochem. Primary amines protect against retinal degeneration in mouse models of retinopathies. Post-translational modifications of the serotonin type 4 receptor heterologously expressed in mouse rod cells. Mechanism of all-trans-retinal toxicity with implications for stargardt disease and age-related macular degeneration. The significance of G protein-coupled receptor crystallography for drug discovery.
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Enzymatic properties and regulation of the native isozymes of retinal membrane guanylyl cyclase RetGC from mouse photoreceptors. Heterogeneous N-terminal acylation of retinal proteins results from the retina's unusual lipid metabolism. Bereta G, Palczewski K. Key enzymes of the retinoid visual cycle in vertebrate retina. Biochim Biophys Acta. Role of bulk water in hydrolysis of the rhodopsin chromophore. Retinyl ester storage particles retinosomes from the retinal pigmented epithelium resemble lipid droplets in other tissues.
Serial sectioning for examination of photoreceptor cell architecture by focused ion beam technology. J Neurosci Methods. Role of membrane integrity on G protein-coupled receptors: Rhodopsin stability and function. Prog Lipid Res. Toll-like receptor 3 is required for development of retinopathy caused by impaired all-trans-retinal clearance in mice. Activation of G protein-coupled receptor kinase 1 involves interactions between its N-terminal region and its kinase domain.
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Noninvasive multiphoton fluorescence microscopy resolves retinol and retinal condensation products in mouse eyes. Blind dogs that can see: pharmacological treatment of Leber congenital amaurosis caused by a defective visual cycle. Arch Ophthalmol. Conformational changes in the g protein-coupled receptor rhodopsin revealed by histidine hydrogen-deuterium exchange. Vitamin A deficiency results in meiotic failure and accumulation of undifferentiated spermatogonia in prepubertal mouse testis.
Biol Reprod. Characterization of the secondary binding sites of Maclura pomifera agglutinin by glycan array and crystallographic analyses. Electrostatic compensation restores trafficking of the autosomal recessive retinitis pigmentosa EK opsin mutant to the plasma membrane. Pulagam LP, Palczewski K. An acyl-covalent enzyme intermediate of lecithin:retinol acyltransferase. Golczak M, Palczewski K.
Structural characterization of the rod cGMP phosphodiesterase 6. J Mol Biol. Beta,beta-carotene decreases peroxisome proliferator receptor gamma activity and reduces lipid storage capacity of adipocytes in a beta,beta-carotene oxygenase 1-dependent manner. Trends Biochem Sci. Retinoids for treatment of retinal diseases. Complexes between photoactivated rhodopsin and transducin: progress and questions.
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Phagocytosis of retinal rod and cone photoreceptors. Physiology Bethesda. Kevany BM, Palczewski K. Importance of membrane structural integrity for RPE65 retinoid isomerization activity. ISX is a retinoic acid-sensitive gatekeeper that controls intestinal beta,beta-carotene absorption and vitamin A production. Adv Exp Med Biol. Molecular biology and analytical chemistry methods used to probe the retinoid cycle.
Visualization of retinoid storage and trafficking by two-photon microscopy. Imanishi Y, Palczewski K. Multiple pathways ensure retinoid delivery to milk: studies in genetically modified mice. Am J Physiol Endocrinol Metab. Increased adiposity in the retinol saturase-knockout mouse. A functional kinase homology domain is essential for the activity of photoreceptor guanylate cyclase 1.
Crystal structure of native RPE65, the retinoid isomerase of the visual cycle. Activation of retinoic acid receptors by dihydroretinoids. Use of thallium to identify monovalent cation binding sites in GroEL. Structural waters define a functional channel mediating activation of the GPCR, rhodopsin. Limited roles of Rdh8, Rdh12, and Abca4 in all-trans-retinal clearance in mouse retina. Phospholipids are needed for the proper formation, stability, and function of the photoactivated rhodopsin-transducin complex.
Structure of cone photoreceptors. Evaluation of potential therapies for a mouse model of human age-related macular degeneration caused by delayed all-trans-retinal clearance. Modulation of molecular interactions and function by rhodopsin palmitylation.
Conserved waters mediate structural and functional activation of family A rhodopsin-like G protein-coupled receptors. Clarin-1, encoded by the Usher Syndrome III causative gene, forms a membranous microdomain: possible role of clarin-1 in organizing the actin cytoskeleton. Usher syndrome IIIA gene clarin-1 is essential for hair cell function and associated neural activation.
Loss of cone photoreceptors caused by chromophore depletion is partially prevented by the artificial chromophore pro-drug, 9-cis-retinyl acetate. Involvement of all-trans-retinal in acute light-induced retinopathy of mice. Chemokine receptors and other G protein-coupled receptors. Lodowski DT, Palczewski K. Comparative analysis of GPCR crystal structures. Photochem Photobiol. Focus on molecules: guanylate cyclase-activating proteins GCAPs. Exp Eye Res.
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Topology of class A G protein-coupled receptors: insights gained from crystal structures of rhodopsins, adrenergic and adenosine receptors. Mustafi D, Palczewski K. Isolation and functional characterization of a stable complex between photoactivated rhodopsin and the G protein, transducin. Vision Res. Impact of retinal disease-associated RPE65 mutations on retinoid isomerization. Effects of long-term administration of 9-cis-retinyl acetate on visual function in mice.
Retinopathy in mice induced by disrupted all-trans-retinal clearance. Retinyl ester homeostasis in the adipose differentiation-related protein-deficient retina. Trafficking of membrane-associated proteins to cone photoreceptor outer segments requires the chromophore cis-retinal. The molecular basis of retinoid absorption: a genetic dissection. Structures of rhodopsin kinase in different ligand states reveal key elements involved in G protein-coupled receptor kinase activation.
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Activation of G protein-coupled receptors: beyond two-state models and tertiary conformational changes. Retinyl ester formation by lecithin: retinol acyltransferase is a key regulator of retinoid homeostasis in mouse embryogenesis. Redundant and unique roles of retinol dehydrogenases in the mouse retina. Efficient coupling of transducin to monomeric rhodopsin in a phospholipid bilayer.
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Delivery of retinoid-based therapies to target tissues. International Union of Basic and Clinical Pharmacology. Recommendations for the recognition and nomenclature of G protein-coupled receptor heteromultimers. Crystal packing analysis of Rhodopsin crystals. Visual rhodopsin sees the light: structure and mechanism of G protein signaling.
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Topology and membrane association of lecithin: retinol acyltransferase. Sequestration of retinyl esters is essential for retinoid signaling in the zebrafish embryo. The role of the cis-retinal ring methyl substituents in visual pigment formation. Crystal structure of a photoactivated deprotonated intermediate of rhodopsin. Retinol dehydrogenase RDH12 protects photoreceptors from light-induced degeneration in mice.
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