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Vitamin B3, the Nicotinamide Adenine Dinucleotides and Aging

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Mechanisms of Ageing and Development 131 (2010) 287–298 Contents lists available at ScienceDirect Mechanisms of Ageing and Development journal homepage: www.elsevier.com/locate/mechagedev Review Vitamin B3, the nicotinamide adenine dinucleotides and aging Ping Xu, Anthony A. Sauve * Department of Pharmacology, Weill Medical College of Cornell University, 1300 York Avenue LC216, New York, NY, 10065, United States ARTICLE INFO ABSTRACT Article history: Organism aging is a process of time and maturation culminating in senescence and death. The molecular Available online 20 March 2010 details that define and determine aging have been intensely investigated. It has become appreciated that the process is partly an accumulation of random yet inevitable changes, but it can be strongly affected by Keywords: genes that alter lifespan. In this review, we consider how NAD+ metabolism plays important roles in the NAD random patterns of aging, and also in the more programmatic aspects. The derivatives of NAD+, such as NADPH reduced and oxidized forms of NAD(P)+, play important roles in maintaining and regulating cellular Vitamin B3 redox state, Ca2+ stores, DNA damage and repair, stress responses, cell cycle timing and lipid and energy Aging metabolism. NAD+ is also a substrate for signaling enzymes like the sirtuins and poly-ADP- Metabolism ribosylpolymerases, members of a broad family of protein deacetylases and ADP-ribosyltransferases that regulate fundamental cellular processes such as transcription, recombination, cell division, proliferation, genome maintenance, apoptosis, stress resistance and senescence. NAD+-dependent enzymes are increasingly appreciated to regulate the timing of changes that lead to aging phenotypes. We consider how metabolism, specifically connected with Vitamin B3 and the nicotinamide adenine dinucleotides and their derivatives, occupies a central place in the aging processes of mammals. ß 2010 Published by Elsevier Ireland Ltd. 1. Introduction Aging is a complex process in which tissues and organs Abbreviations: AADPR, O-acetyl-ADP-ribose; ACCa, acetyl-CoA carboxylase-alpha; accumulate changes over the lifetime of an organism. Late in life AceCS1 and 2, acetyl-coenzyme A synthetase 1 and 2; ADPR, adenosine 5’- these changes are associated with a reduced level of tissue function diphosphoribose; ANT2, 3adenine nucleotide translocator isoform 2 and 3; ARTs, and a reduced capacity for tissue self-repair (Rossi et al., 2007). ADP-ribosyl trasnferases; ARCs, ADP-ribosyl cyclases; ATP5A, human mitochon- Accompanying this condition is an increased vulnerability to drial ATP synthase alpha-subunit; Bax, BCL2-associated X protein; cADPR, cyclic ADP-ribose; CPS1, carbamoyl phosphate synthetase 1; DBC1, deleted in breast disease. In humans, age-related diseases include cancer, arthritis, cancer-1; FOXO, forkhead box transcription factor; G6PDH, glucose-6-phosphate cataracts, osteoporosis, type II diabetes, hypertension, heart dehydrogenase; GDH, glutamate dehydrogenase; HSP70, 70 kilodalton heat shock diseases, Alzheimer’s disease, stroke and other neurodegenerative proteins; IDH, isocitrate dehydrogenase; IDP, NADP+-dependent isocitrate dehy- disorders, all of which cumulatively reflect the major causes of drogenases; IP3, inositol trisphosphate; Ku70, lupus Ku autoantigen protein p70; death late in human life (except infection). Molecular and cell IDO, indoleamine 2,3-dioxygenase; LPS, lipopolysaccharides; LXR, liver X receptor; MEF2D, myocyte-specific enhancer factor 2D; MEP, NADP+-dependent malic biological studies have revealed that aging in tissues is character- enzymes; Mtap, methylthioadenosine phosphorylase; NA, nicotinic acid; NaAD, ized by hallmark features such as telomere shortening (von Figura nicotinic acid adenine dinucleotide; NAADP, nicotinic acid adenine dinucleotide et al., 2009), reduced capacities for DNA repair (Gorbunova et al., phosphate; NADKs, NAD+ kinase; NAM, nicotinamide; NaMN, nicotinic acid 2007; Rossi et al., 2007), reduced stress responses (Knight, 2001; mononucleotide; Nampt, nicotinamide phosphoribosyltransferase; NAPRTase, nicotinic acid phosphoribosyltransferase; NF-kB, nuclear factor kappa-light- Naidoo, 2009), increased sensitivity to apoptosis and decreased cell chain-enhancer of activated B cells; Nmnat, nicotinamide/nicotinc acid mono- division (Rossi et al., 2007). In addition, aging causes profound nucleotide adenylyltransferase; NMN, nicotinamide mononucleotide; NR, nicoti- changes in tissue and systemic metabolism (Finley and Haigis, namide riboside; NOX, NADPH oxidase; Nrk1,2, nicotinamide riboside kinase; 2009) and endocrine signaling, such as slowed metabolic rate and PARPs, poly-ADP-ribosyl polymerases; PGC1-a, PPAR-gamma coactivator 1; PPAR- increased insulin resistance (Arai et al., 2009). g, peroxisome proliferator-activated receptor-gamma; p53, tumor protein 53; Pol I, + RNA polymerase I; Pnp1, purine-nucleoside phosphorylase 1; Qprtase, Quinolinate Nicotinamide dinucleotide (NAD ), and its derivatives such as phosphoribosyltransferase; SUV39H1, histone-lysine N-methyl-transferase; NADH, NADP(H) are small molecules that are central cellular Smad7, SMAD family number 7; TAFI68, TBP-associated factors; TH, transhydro- metabolites and are increasingly recognized to play important genase; TNF-a, tumor necrosis factor alpha; WRN, Werner syndrome ATP- roles in the aging process (Lin and Guarente, 2003; Sinclair and dependent helicase; 6GDH, 6-gluconate phosphate dehydrogenase. Guarente, 2006; Pollak et al., 2007; Yang et al., 2007). Consistently, * Corresponding author. Tel.: +1 212 746 6224. E-mail address: [email protected] (A.A. Sauve). these metabolites function in energy metabolism, DNA repair 0047-6374/$ – see front matter ß 2010 Published by Elsevier Ireland Ltd. doi:10.1016/j.mad.2010.03.006 288 P. Xu, A.A. Sauve / Mechanisms of Ageing and Development 131 (2010) 287–298 Table 1 Cellular functions of NAD+-related metabolites and their relationship to human health. Molecules Cellular functions Effects on human health References Nicotinamide Increase NAD+ level; Lifespan extension of human cell; Hoane et al. (2006a,b); Kang et al. (2006); Lin et al. (2001) Inhibition on PARPs, sirtuins Neuroprotection from brain injury; Prevention of vascular injury, lung injury Treatment of pellagra Nicotinic acid Reduce lipids (cholesterol, triglycerides, Treatment of cardiovascular diseases; Benavente et al. (2009); Penberthy (2009); low-density lipoproteins) Scanu and Bamba (2008) Increase HDL level; Neuroprotection; Increase NAD+ level Treatment of skin diseases, including pellagra NAD+/NADH NAD+ metabolism; NAD+ depletion and cell death; Bedard and Krause (2007) Component in cellular antioxidation systems Associated with brain ischemia, diabetes, cancer, cardiovascular diseases NADP/NADPH Component in cellular antioxidation systems; Associated with brain ischemia, diabetes, Bedard and Krause, 2007 cancer, cardiovascular diseases Substrate of NADPH oxidase NAADP Ca2+ signaling Smooth muscle tension Galione (2006) cADPR Ca2+ signaling Unknown Galione (2006) AADPR Substrate of yeast Ysa1; Unknown Liou et al. (2005); Tong et al. (2009) Promotion of SIR assembly in yeast ADPR ADP-ribosylation of protein by ARTs Unknown Corda and Di Girolamo (2003); and PARPs Peralta-Leal et al. (2009) pathways and cell protection pathways as well as cell death differentiate NAD+ and NADP+ functionally, and indeed the cell pathways (Hassa et al., 2006; Pollak et al., 2007; Yang et al., 2007). maintains a high NADPH/NADP ratio, consistent with use of These metabolites can be thought of as central mediators that NADPH as a reductant and as a cell protectant against oxidative influence a variety of cellular processes known to change in aging. stress (Pollak et al., 2007). Increasing evidence suggests that the In this review, we consider the recent biological and medical absolute amounts of NAD(H) and NADP(H) are important in efforts being made to understand how NAD(P)+ metabolism and its maintaining cellular health. central effects influence aging processes. 3. Metabolic transformations of NAD+ and NADH in cells 2. Vitamin B3, distribution of NAD+ and NADP+ and their redox partners NAD+ is metabolized in several ways in cells. Its metabolites include NADH, NADP+ and NADPH, nicotinic acid adenine Vitamin B3 is the precursor of nicotinamide dinucleotide dinucleotide phosphate (NAADP), adenosine 50-diphosphoribose (NAD+), and is commonly sold as a supplement in two major forms, (ADPR), cyclic ADP-ribose (cADPR) and O-acetyl-ADPR (AADPR) namely nicotinamide (NAM) and nicotinic acid (NA). The latter (Fig. 1). ADP-ribose is also generated from NAD+, and ADP- form of vitamin B3 is commonly known as niacin. Most raw foods ribosylation of proteins is an important post-translational also provide these substances, or provide them after digestive modification, expecially for genome stability (Hassa et al., 2006). action, although diets of corn are noteworthy in being a poor A major reaction of NAD+ in metabolism is to accept a hydride source of B3 (Goldsmith et al., 1952). These substances in their equivalent to generate NADH. NADH can be re-oxidized back to unmodified forms can regulate cellular activities through receptor NAD+ by many reactions within cells, but within the mitochondria, activation and also through inhibition of signaling enzymes. They electrons from NADH provide the chemical energy by which ATP are also precursors for the biosynthesis of derivatives of NAD+. can be generated. The effects of
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