ATP/ITP metabolism
ATP plays the important role in a metabolism. This compound is a
universal energy source for all biochemical processes occurring in live
systems. Knowledge of Inosine metabolism has led to advances in
immunotherapy in recent decades.
ATP is often used as a phosphate source, e.g., in the reaction with
Neopterin diphosphate (NDP) that results in formation of ADP
and Neopterin-3'-triphosphate (NTP), as well as in the reaction
with Inosine diphosphate (IDP) in which Inosine triphosphate (ITP)
is formed. These reactions are catalyzed by similar enzymes, Nucleoside
diphosphate kinase (NDPK complex) [1],
[2], [3], [4],
[5], Non-metastatic cells 4, protein
expressed in (NDPK D (mitochondrial)) [6],
[7], [8],
Non-metastatic cells 3, protein expressed in (NDPK C) [7],
[9], [10],
Nucleoside non-metastatic cells 6, protein expressed in
(nucleoside-diphosphate kinase) (NDPK 6) [8],
[11], Non-metastatic cells 7, protein
expressed in (nucleoside-diphosphate kinase) (NDPK 7) [8]
and non-metastatic cells 2, protein (NM23B) expressed in, pseudogene 1 (NDPK
8) [12].
Hydrolysis of ATP to ADP proceeds in two ways and
catalyzed by specific groups of enzymes. The first group consists of
Ectonucleoside triphosphate diphosphohydrolase 2 (ENTPD2-alpha) [13],
[14], Ectonucleoside triphosphate
diphosphohydrolase 1 (ENP1) [15], [16],
[17], [18],
Ectonucleoside triphosphate diphosphohydrolase 3 (ENP3) [19],
[20], [21],
Acylphosphatase 1, erythrocyte (common) type (ACYP1) [22],
[23], Acylphosphatase 2, muscle type (ACYP2)
[22], [23], Acid
phosphatase 5, tartrate resistant (PPA5) [24],
[25], Epoxide hydrolase 2, cytoplasmic (EPHX2)
[26], and Alkaline phosphatase, placental
(Regan isozyme) (ALPP) [26]. The
second group consists of Acid phosphatase 2, lysosomal (PPAL) [27],
[28], [29], Acid
phosphatase 5, tartrate resistant (PPA5) [30],
[31], [32], [33],
and Acid phosphatase, prostate (PPAP) [34],
[35], [36], [37].
Second group also catalyzes further hydrolysis of ADP to AMP
and AMP to release Adenosine.
There are two processes that lead to ITP hydrolysis. The first is
a reaction catalyzed by Ectonucleoside triphosphate diphosphohydrolase 1
(ENP1) [15], [18],
[19], Ectonucleoside triphosphate
diphosphohydrolase 3 (ENP3) [19], [20],
[21], and Ectonucleoside triphosphate
diphosphohydrolase 6 (putative function) (ENTPD6) [38],
[39]. It results in formation of IDP.
These enzymes also participate in the following hydrolysis of IDP
to Inosine monophosphate (IMP). And in the second case ITP
is hydrolyzed directly to IMP by the action Inosine
triphosphatase (nucleoside triphosphate pyrophosphatase) (ITPA) [40],
[41], [42], [43].
Yet another process leading to formation of ADP is the reaction
of ATP with AMP catalyzed by Adenylate kinase 5 (AK5)
[44], Adenylate kinase 1 (AK1) [44],
[45], Adenylate kinase 2 (AK2) [44],
[45], Adenylate kinase 3-like 1 (AK3) [44],
[46], TAF9 RNA polymerase II, TATA box
binding protein (TBP)-associated factor, 32kDa (KAD6) [47],
[48], and Adenylate kinase 7 (KAD7) [49],
[50].
ADP can participate in reaction of formation of 2 '-deoxy-ADP (dADP).
This reaction is catalyzed by Ribonucleotide reductase. This
enzyme is involved in one more reaction of formation of 2 '-deoxy-IDP (dIDP)
from IDP [51], [52],
[53], [54], [55].
dADP and dIDP take part in the dATP/dITP metabolism.
AMP can be hydrolyzed to IMP via two pathways. The first
is a direct hydrolysis catalyzed by AMP deaminase [56],
[57], Adenosine monophosphate deaminase 2
(isoform L) AMP deaminase 2 [58], [59],
[60], Adenosine monophosphate deaminase 1
(isoform M) AMP deaminase 1 [61], [62],
[63], [64], and
Adenosine monophosphate deaminase (isoform E) (AMD3) [65],
[66], [67], [68].
The second is represented by a chain of consecutive reactions: formation
of Adenylo-succinate catalyzed by Adenylosuccinate lyase (ADSL)
[69], [70], [71],
[72], [73]
followed by formation of IMP in the presence of Adenylosuccinate
synthase (ADSS) [74], [75],
[76], [77] and
Adenylosuccinate synthase like 1 (ADSSL1) [74],
[75], [76], [77],
[78]. IMP also takes part in IMP
biosynthesis and GTP-XTP metabolism.
AMP can directly form Adenine, this reaction occurs in the
presence of Adenine phosphoribosyltransferase (APRT) [79],
[80], [81].
Similar reaction proceeds for IMP from Hypoxanthine under
the action of Hypoxanthine phosphoribosyltransferase 1 (HPRT) [82],
[83], [84], [85],
[86]. Adenine and Hypoxanthine
participate in other processes, e.g., dATP/dITP metabolism and in
GTP-XTP metabolism.
Nucleoside phosphorylase (PNPH) catalyzes the formation of Adenine
from Adenosine [87], [88],
[89] and Inosine from Hypoxanthine [87],
[88], [89], [90],
[91], [92]. Inosine
can also be produced as a result of hydrolysis of Adenosine by
Adenosine deaminase (ADA) [93], [94],
[95], Adenosine deaminase, RNA-specific (ADAR1)
[96], Adenosine deaminase, RNA-specific, B1
(RED1 homolog rat) (ADAR2) [96]
Adenosine deaminase, RNA-specific, B2 (RED2 homolog rat) (ADAR3) [96].
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