Other Functions of Nucleotides:- Nucleotides Carry Chemical Energy in Cells
The phosphate group covalently linked at the 5 hydroxyl of a ribonucleotide may have one or two additional phosphates attached. The resulting molecules are referred to as nucleoside mono-, di-, and triphosphates (Fig. 8–39). Starting from the ribose, the three phosphates are generally labeled α, β, and γ. Hydrolysis of nucleoside triphosphates provides the chemical energy to drive a wide variety of cellular reactions. Adenosine 5-triphosphate, ATP, is by far the most widely used for this purpose, but UTP, GTP, and CTP are also used in some reactions. Nucleoside triphosphates also serve as the activated precursors of DNA and RNA synthesis, as described in Chapters 25 and 26.
The energy released by hydrolysis of ATP and the other nucleoside triphosphates is accounted for by the structure of the triphosphate group. The bond between the ribose and the phosphate is an ester linkage. The α, β and β, γ linkages are phosphoanhydrides (Fig. 8–40). Hydrolysis of the ester linkage yields about 14 kJ/mol under standard conditions, whereas hydrolysis of each anhydride bond yields about 30 kJ/mol. ATP hydrolysis often plays an important thermodynamic role in biosynthesis. When coupled to a reaction with a positive free-energy change, ATP hydrolysis shifts the equilibrium of the overall process to favor product formation (recall the relationship between equilibrium con stant and free-energy change described by Eqn 6–3 on p. 195).


FIGURE 8–39 Nucleoside phosphates. General structure of the nucleoside 5-mono-, di-, and triphosphates (NMPs, NDPs, and NTPs) and their standard abbreviations. In the deoxyribonucleoside phosphates (dNMPs, dNDPs, and dNTPs), the pentose is 2-deoxy-D-ribose.