ライフサイエンスコーパス: unwinding

1 endent ATPase activity and the extent of RNA unwinding.

2 ell it tolerates potential roadblocks to DNA unwinding.

3 s the degree to which re-winding counteracts unwinding.

4 gions that depend strongly on eIF4A-mediated unwinding.

5 have mostly been ignored with regard to DNA unwinding.

6 significantly affecting ATP-dependent duplex unwinding.

7 ance of the SEW model for hexameric helicase unwinding.

8 the coupling between ATP hydrolysis and RNA unwinding.

9 into the Brr2 helicase active site ready for unwinding.

10 o be directly involved in regulating U1/5'ss unwinding.

11 ble-stranded RNA, resulting in gradual helix unwinding.

12 08-like modules that are responsible for RNA unwinding.

13 spliceosomal activation in addition to U4/U6 unwinding.

14 creating torsional strain and leading to DNA unwinding.

15 nsuring that synthesis is tightly coupled to unwinding.

16 shaped hexamers that encircle DNA for duplex unwinding.

17 ment of the GINS helicase activator, and DNA unwinding.

18 displaced strand and duplex coordinates DNA unwinding.

19 with blunt-ends or 3'-ssDNA overhangs by DNA unwinding.

20 ' ssDNA overhang is required to initiate DNA unwinding.

21 d is a less understood characteristic of DNA unwinding.

22 to gain access to ssDNA and facilitating DNA unwinding.

23 tations, blocks on either strand inhibit CMG unwinding.

24 iation that functions beyond its role in RNA unwinding.

25 ifically with the excluded DNA strand during unwinding.

26 nterstrand distance and can only elongate by unwinding.

27 trand cross-link in the dsDNA that prevented unwinding.

28 ng structural and mechanistic aspects of DNA unwinding.

29 elucidate the structural basis of duplex DNA unwinding.

30 ivities, including DNA binding, nicking, and unwinding.

31 g several kinetic parameters associated with unwinding.

32 the Thermobifida fusca type I-E Cascade: (1) unwinding 11 bp of dsDNA at the seed-sequence region to

33 ith the PIC and plays important roles beyond unwinding 5'-UTR structure is consistent with a recent p

34 helicase exhibited greater processivity when unwinding a DNA fork compared to a ss/ds DNA junction su

35 x RNA helicase that is responsible for U4/U6 unwinding, a critical step in spliceosomal activation.

36 '-ended single-stranded DNA during continued unwinding; a crevice between RecB and RecC increasingly

37 by enhancing the coupled cap-binding and RNA-unwinding activities of eIF4F.

38 ase with strong 5' --> 3' and weak 3' --> 5' unwinding activities.

39 Srs2 also possesses a DNA unwinding activity and here we show that assembly of mor

40 insight into the regulation of nucleic acid unwinding activity and introduces a monomeric superhelic

41 Cas9 proteins have limited dsDNA binding and unwinding activity and promiscuous guide RNA specificity

42 s with oligomerization and thereby modulates unwinding activity and RNA affinity of the helicase.

43 king of a helicase monomer with undetectable unwinding activity converts it into a superhelicase that

44 ATP-dependent DNA unwinding activity has been demonstrated for recombinant

45 at MOV10L1 exhibits 5'-to-3' directional RNA-unwinding activity in vitro and that a point mutation th

46 mutant in vivo showed reduced stimulation of unwinding activity in vitro.

47 purified protein exhibits a "hyper-helicase" unwinding activity in vitro.

48 ormed a complex with Dhr1 and stimulated its unwinding activity in vitro.

49 r; however, the contextual regulation of its unwinding activity is not fully described.

50 e used to study the assembly pathway and DNA unwinding activity of the bacteriophage T4 helicase-prim

51 an activated gene, but also impairs the DNA unwinding activity of XPD and the nucleotide excision re

52 RNA substrates and had significantly reduced unwinding activity on DNA.

53 Tim stimulates DDX11 unwinding activity on forked DNA substrates up to 10-fol

54 Based on its in vitro unwinding activity on G-quadruplex (G4) DNA, the Bloom s

55 otein, the C-terminal helicase domain had no unwinding activity on RNA substrates and had significant

56 eal that UvrD exhibits two distinct types of unwinding activity regulated by its stoichiometry.

57 We conclude that AdnAB's processive DSB unwinding activity suffices for AdnAB function in HR.

58 n order to engage with DNA, regulate its DNA-unwinding activity, and maintain genome stability.

59 alanine causes an even greater reduction in unwinding activity, which is somewhat surprising as this

60 d protein possesses branch migration and DNA unwinding activity.

61 d triplex DNA structures and inhibited ChlR1 unwinding activity.

62 h the excluded strand act as a regulator for unwinding activity.

63 f the arch domain has minimal effect on Mtr4 unwinding activity.

64 e and the KH domain is required for its full unwinding activity.

65 ase/CDC7-Dbf4 kinase and exhibit reduced DNA unwinding activity.

66 teraction with Srs2 blocks the helicase (DNA unwinding) activity of Srs2.

67 Instead of leaving after G4 unwinding, allowing it to refold, or going beyond to unw

68 Combined DNA unwinding and annealing activities are typical of certai

69 strand-exchange involves active coupling of unwinding and annealing reactions by the TWINKLE.

70 as well as events corresponding to stepwise unwinding and annealing.

71 bed, obtaining data and analyzing results of unwinding and ATPase assays takes approximately 4 h.

72 no acid change in the KH domain, had reduced unwinding and binding activates on RNA and DNA substrate

73 mechanism that controls eIF4AI-mediated mRNA unwinding and can guide further mechanistic studies on o

74 its Cas3, a nuclease-helicase that catalyzes unwinding and cleavage of foreign double-stranded DNA (d

75 tional analysis allowed us to split the mRNA unwinding and codon selection activities of DHX29.

76 zyme complexes are highly processive, duplex unwinding and degrading machines that require tight regu

77 which MOV10L1 RNA helicase activity promotes unwinding and funneling of the single-stranded piRNA pre

78 to identify interactions that promote U4/U6 unwinding and have studied their impact in yeast.

79 play complex motion, which includes winding, unwinding and helix inversion, as dictated by their init

80 chanism of nucleosome unfolding in which DNA unwinding and histone protein disassembly are coupled.

81 r data reveal conserved mechanisms for U4/U6 unwinding and indicate telestem dynamics are critical fo

82 t duplex unwinding, suggestive of a cycle of unwinding and inhibition by Dbp2.

83 re-based mutants providing insights into the unwinding and loading mechanism of RNAs.

84 ghly conserved phenylalanine for Pur-alpha's unwinding and neuroprotective function.

85 leic acid processing enzymes involved in DNA unwinding and polymerization, our results suggested the

86 st that mycobacterial HR can proceed via DSB unwinding and protein capture of the displaced 3'-OH sin

87 t couple the energy of ATP hydrolysis to the unwinding and remodeling of structured DNA or RNA, which

88 ng optical tweezers, we measure the rates of unwinding and rewinding for these two states and show th

89 rgets branched DNA substrates and drives the unwinding and rewinding of DNA strands.

90 rein RecQ melts dsDNA internally to initiate unwinding and subsequently assembles at the fork into a

91 The helicase RTEL1 promotes t-loop unwinding and suppresses telomere fragility to maintain

92 naA assembles at the origin and promotes DNA unwinding and the assembly of a replication initiation c

93 trands encircled by the helicase ring during unwinding and the ring orientation at the replication fo

94 in plays critical roles in NTP-dependent RNA unwinding and translocation during viral replication.

95 -hairpin), showed a decrease in DNA binding, unwinding, and annealing, as expected for a functional R

96 angular and linear motions such as torsion, unwinding, and sliding in addition to the previously rep

97 ated mechanic effects like local elongation, unwinding, and softening of the DNA often remain in ques

98 o cleavage complex, comet, DNA intercalating/unwinding, and Topo IIalpha-mediated ATP hydrolysis assa

99 the frequency of initiation and the rate of unwinding are highly dependent on RecQ concentration.

100 entify binding sites for ssDNA during SsoMCM unwinding as well as validating the importance of the SE

101 As evidenced by an alkaline comet unwinding assay, 3 induces extensive DNA damage, suggest

102 Novel single molecule FRET binding and unwinding assays show an interaction of the excluded str

103 lecule Forster resonance energy transfer and unwinding assays to identify interactions that promote U

104 e mechanism of its helicase function, RecBCD unwinding at low adenosine triphosphate (ATP) (2-4 muM)

105 helicase-primase complex is required for DNA unwinding at the replication fork and synthesis of prime

106 s of RHA are responsible for such repetitive unwinding behavior in addition to providing an increased

107 allele of MCM10 that stimulates initial DNA unwinding but is defective in replication elongation and

108 d in the crystal structure, is vital for DNA unwinding but not for DNA binding.

109 dly during heat stress, which eliminates RNA unwinding (but not rewinding) competence.

110 o a stepwise decrease of Brr2-mediated U4/U6 unwinding, but that unwinding is largely restored by a B

111 DSB unwinding by AdnAB in vitro is stringently dependent on

112 al protein, Mcm10, drives initial origin DNA unwinding by an unknown mechanism.

113 rved aromatic loop (AL) is important for DNA unwinding by bacterial RecQ [23, 24] and truncated RECQ1

114 We show that Topo IIIalpha stimulates DNA unwinding by BLM in a manner that is potentiated by RMI1

115 o position the RNA guide for DNA binding and unwinding by Cas8c.

116 d a fluorescence assay to monitor RNA duplex unwinding by DEAD-box helicases in real time.

117 ability of G4 substrates affects binding and unwinding by DHX36.

118 stability of the G4 substrates and rates of unwinding by DHX36.

119 eIF4G strongly stimulates the rate of duplex unwinding by eIF4A on the IRES.

120 , DnaA filaments assemble and promote duplex unwinding by engaging and stretching a single DNA strand

121 for the same substrate, hence prominent DNA unwinding by hDNA2 alone can only be observed using the

122 generalizing a previous model of processive unwinding by helicases, we provide a unified framework f

123 rnative to the current general model for DNA unwinding by hexameric helicases.

124 a directional contrahelicase, blocking mtDNA unwinding by the mitochondrial helicase TWINKLE.

125 DNA unwinding by the replicative helicase may continue durin

126 eat sequence DNA also stimulated binding and unwinding by these enzymes.

127 ondrial replication machinery, including DNA unwinding by Twinkle helicase.

128 Most strikingly, G4 unwinding by WRN was inhibited approximately 50% for all

129 e coupling between ATP hydrolysis and duplex unwinding can be determined.

130 amics was coupled to RecBCD entering into an unwinding-competent state that required a sufficiently l

131 nal dynamics of the RecBCD-DNA complex in an unwinding-competent state, arising, in part, by an enzym

132 Transcription requires unwinding complementary DNA strands, generating torsiona

133 The efficiency of unwinding correlates with the stability of the 'closed-s

134 structure, with the helicases preferentially unwinding D-loops.

135 For RNA unwinding, DDX3X shows a greater preference than Ded1p f

136 s recombinogenic single-stranded DNA ends by unwinding DNA and cutting it a few nucleotides to the 3'

137 is the replicative helicase responsible for unwinding DNA during archaeal and eukaryal genome replic

138 cialized helicases play an important role in unwinding DNA structures to maintain genome stability.

139 tive and thus has all features essential for unwinding DNA.

140 randed nucleic acid and couple the motion to unwinding double-strands of a duplex nucleic acid.

141 ion, could provide RecQ with a mechanism for unwinding duplex and other DNA structures.

142 ys ATP hydrolysis activity and is capable of unwinding duplex DNA.

143 erhelicase with strong RNA affinity and high unwinding efficiency on a broad range of targets.

144 hown to unwind DNA via a SEW mode to enhance unwinding efficiency.

145 on with one strand of the plasmid origin DNA unwinding element (DUE) contribute to strand-specific re

146 The DNA unwinding element (DUE)-binding protein (DUE-B) binds to

147 RECQ1 is an ATP-dependent DNA-unwinding enzyme (helicase) [8, 9] with roles in replica

148 tures by preferentially interacting with and unwinding exposed RNA secondary structure.

149 These results support the SEW model of unwinding for EcDnaB that expands on the existing SE mod

150 obases involves multiple monomers at the DNA unwinding fork.

151 We establish that unwinding forks can initiate internally by melting dsDNA

152 the formation and progression of individual unwinding forks, we observed that both the frequency of

153 have also been shown to be important for DNA unwinding, giving rise to the steric exclusion and wrapp

154 in this work classify them into a low-force unwinding group of fimbriae together with the CFA/I and

155 Moreover, the role of TFB2M in promoter unwinding has not been discriminated from that of TFAM.

156 45 and the outer edge of the Mcm2-7 ring for unwinding have remained unexplored.

157 found that deficiency of BLM, or another G4-unwinding helicase, the Werner syndrome-associated helic

158 dimerization, substrate orientation or flap unwinding impair the structure-specific nuclease activit

159 k biochemical assays show that Yra1 inhibits unwinding in a concentration-dependent manner by prevent

160 h fork rotation and precatenation facilitate unwinding in hard-to-replicate contexts, they intrinsica

161 ation is detrimental to helicase loading and unwinding in the presence or absence of Gp32.

162 As and inhibits Brr2-mediated U4/U6 di-snRNA unwinding in vitro.

163 re-winding of the opened DNA strongly limits unwinding, independent of the 3'-tail.

164 r the 43S PIC requires an ATP-dependent, but unwinding-independent, activity of eIF4A.

165 cation of eIF4G that stimulates eIF4A duplex unwinding independently of eIF4E.

166 Further, the repetitive unwinding induces an efficient annealing of a complement

167 In addition, we show that the probability of unwinding into each state is dependent on the applied fo

168 itate or suppress access to DNA by promoting unwinding into one state or the other.

169 complex must be regulated to ensure that DNA unwinding is coupled with DNA synthesis.

170 bination in vivo, whereas unregulated duplex unwinding is detrimental for recombination precision.

171 e of Brr2-mediated U4/U6 unwinding, but that unwinding is largely restored by a Brr2 cofactor, the C-

172 Unwinding is stimulated by the U6 telestem, which transi

173 Unwinding is then completed by the partially displaced U

174 engages with the telomere to promote t-loop unwinding is unclear.

175 w they engage with a replication fork during unwinding is unknown.

176 mechanism that couples ATP hydrolysis to DNA unwinding is unknown.

177 inds DNA downstream from the location of DNA unwinding, it cannot function using a conventional helic

178 at hDNA2 is a processive helicase capable of unwinding kilobases of dsDNA in length.

179 Hexameric helicases are processive DNA unwinding machines but how they engage with a replicatio

180 that, in addition to stimulating initial DNA unwinding, Mcm10 stabilizes Cdc45 and GINS association w

181 fluorescence assays, we elucidated a unique unwinding mechanism of RNA helicase A (RHA) that entails

182 ealing how this exonuclease uses a novel DNA-unwinding mechanism to separate the polynucleotide stran

183 ments, lead us to suggest a replication fork unwinding mechanism whereby the N-terminal and AAA+ tier

184 an opposite structural twist, suggesting an unwinding mechanism.

185 ength SNARE complex, suggesting a processive unwinding mechanism.

186 nized dynamic facets of replicative helicase unwinding mechanisms.

187 e II (Pol II) open complex (OC) requires DNA unwinding mediated by the transcription factor TFIIH hel

188 sDNA) and stabilize the complex in a forward unwinding mode.

189 Current DNA unwinding models propose that motor translocation is tig

190 nces that differ for G4 disruption and dsDNA unwinding, most likely arising from differences in the r

191 GCTGGTGG 3') during DNA unwinding, RecBCD's unwinding, nuclease, and RecA-loading activities change

192 us a third domain (D3) that is important for unwinding nucleic acid duplexes.

193 hows that duplex DNA enters the helicase and unwinding occurs in the central channel.

194 This conformation requires an unwinding of a portion of the C-terminal helix of the Hi

195 A in a 'closed' preinitiation complex (PIC); unwinding of about 15 base pairs of the promoter DNA to

196 ment of alphaM4 and hypothesize that winding/unwinding of alphaM4 represents a trigger for high-affin

197 w that the expansion does not involve either unwinding of CTPR3 helices or unraveling of interactions

198 olved in protein binding, translocation, and unwinding of DNA and RNA substrates.

199 ated by the DnaA protein, which promotes the unwinding of DNA at oriC We demonstrate that the binding

200 Helicases catalyze the unwinding of double-stranded nucleic acids where structu

201 In this study, recombinant nsP2 demonstrated unwinding of double-stranded RNA in a 5'-3' directionall

202 A prerequisite for DNA replication is the unwinding of duplex DNA catalyzed by a replicative hexam

203 In addition, unwinding of duplex siRNA bound to AGO1 requires passeng

204 However, Pif1 unwinding of duplexes occurs at a much faster rate than

205 kely resulting from enhanced eIF4A-dependent unwinding of G-quadruplexes in the 5' untranslated regio

206 RN involved in the binding and ATPase-driven unwinding of G4 DNA.

207 HA may help in explaining how the repetitive unwinding of helicases contributes to their biological f

208 ned to quantitatively measure long-range DNA unwinding of individual DNA helicases from the archaeons

209 Each cell division requires the unwinding of millions of DNA base pairs to allow chromos

210 Moreover, the NER factor XPA activates unwinding of normal DNA by Core7, but inhibits the Core7

211 The unwinding of nucleic acid secondary structures within ce

212 UvsW-catalyzed unwinding of recombination intermediates such as D-loops

213 n contrast to E. coli DinG, MtDinG catalyzes unwinding of replication fork and Holliday junction stru

214 We show that the preferential unwinding of RNA:DNA hybrids is due to neither specific

215 a RecQ dimer, and that continued processive unwinding of several kilobases involves multiple monomer

216 Recent reports suggest that unwinding of some helicases display repetitive activity,

217 ue to a central bubble that represents local unwinding of specific sequences.

218 verhangs, it could also catalyze significant unwinding of substrates containing 3' overhangs.

219 tant, PcalRG is able to induce ATP-dependent unwinding of synthetic Holliday junctions and ATP-stimul

220 to the transcription start site facilitates unwinding of the DNA duplex downstream of the transcript

221 plified by RNA-dependent ATPase activity and unwinding of the DNA-RNA duplex.

222 cking of one of the template DNA strands and unwinding of the duplex prior to subsequent leading stra

223 he 3' vRNA in the active site and results in unwinding of the duplexed region of the promoter.

224 or sequence complementarity, and (2) further unwinding of the entire protospacer to form a full R-loo

225 The unwinding of the helical structure at the grid walls dri

226 ric structural transition, involving partial unwinding of the helix-X between heme a and a3, thereby

227 he level of inward-open SERT and may lead to unwinding of the TM5 helix to allow phosphorylation.

228 onal and conformational changes triggered by unwinding of the U4 and U6 (U4/U6) snRNAs.

229 Brr2 catalyzes an ATP-dependent unwinding of the U4/U6 RNA duplex, which is a critical s

230 This has complicated the unwinding of their unique non-redundant roles.

231 Moreover, WRN unwinding of these recombination structures is further e

232 wound using DNA oligonucleotides by coupling unwinding of U4/U6 stem II with strand invasion of stem

233 The extent of unwinding of Y-shaped DNA structures was approximately 3

234 ateaus that arise from unzipping, as well as unwinding, of helical bundles.

235 w that E1 generates strikingly heterogeneous unwinding patterns stemming from varying degrees of repe

236 ther, acting as a wedge with an external DNA unwinding point during translocation.

237 th strands enter the helicase and the duplex unwinding point is internal, followed by exclusion of th

238 In contrast, the unwinding processivity always increases when the force i

239 ave coevolved with helicases to increase the unwinding processivity even if the velocity remains unaf

240 along single-stranded DNA, and the increased unwinding processivity may contribute directly to the an

241 We predict that the sensitivity of unwinding processivity to external force is a universal

242 Interestingly, the increase in unwinding processivity with force depends on whether the

243 The lag phase in the unwinding progress curve was reduced for the forked DNA

244 se activity of the complex is activated once unwinding progresses.

245 Human RecQ helicases are DNA unwinding proteins that are critical responders to DNA d

246 g and has a modest stimulatory effect on the unwinding rate of RecQ.

247 ngly, these mutations also increased the DNA unwinding rate, suggesting that electrostatic contacts w

248 se stimulates the helicase by increasing the unwinding rate-constant (kcat), consequently the combine

249 stead of DNA also dramatically increased the unwinding rate.

250 from the excluded strand to regulate the DNA unwinding rate.

251 that the protease cleavage, RNA binding, and unwinding rates of NS3 are minimally affected in vitro.

252 monstrated the utility of the probe in a DNA unwinding reaction using a helicase from Bacillus and fo

253 ue to an increase in the processivity of the unwinding reaction.

254 ctively, these observations show that during unwinding, RecBCD binds to DNA in a dynamic mode that is

255 bination hotspot (5' GCTGGTGG 3') during DNA unwinding, RecBCD's unwinding, nuclease, and RecA-loadin

256 randed DNA and how ATP hydrolysis drives DNA unwinding remain open questions.

257 the course of hours whereas steps involving unwinding-rewinding of the helix proceeded over the cour

258 ithout dissociation, resulting in repetitive unwinding/rewinding cycles.

259 the velocity at which the force required for unwinding rises exponentially with increased speed) of 1

260 y active form that is capable of immediately unwinding RNA duplexes without undergoing rate-limiting

261 Instead, Pif1 is capable of unwinding RNA:DNA heteroduplexes with moderately greater

262 A with tight affinity are incapable of fully unwinding short duplex RNAs.

263 Interestingly, although MtRecD-catalyzed unwinding showed a markedly higher preference for duplex

264 The initiation of substrate unwinding showed some sequence dependency, while DNA bin

265 ould have implications for understanding the unwinding specificity of pNS3h, which is active only on

266 substeps consisting of binding, activation, unwinding, stalling and reactivation stages.

267 rminal domain is shown to play a role in DNA unwinding, strand annealing, and Holliday junction (HJ)

268 atalyze strand-exchange reaction between the unwinding substrate and a homologous single-stranded DNA

269 main plays a previously unrecognized role in unwinding substrates.

270 TFAM inhibits Twinkle unwinding, suggesting other replisome proteins may be re

271 zing this interaction accelerates RNA duplex unwinding, suggesting that it is present in solution and

272 ons as an inhibitor of Dbp2-dependent duplex unwinding, suggestive of a cycle of unwinding and inhibi

273 ggests a mechanism for DNA translocation and unwinding, suggests how the enzyme binds specifically to

274 ove beyond the site of DNA synthesis, likely unwinding template DNA.

275 ontributions with the excluded strand during unwinding, termed steric exclusion and wrapping (SEW).

276 helicase activity, which is responsible for unwinding the DNA while it is being transported to a rec

277 s in providing molecular information without unwinding the duplex.

278 hromosome ends, effectively block FANCJ from unwinding the forked duplex telomeric substrate.

279 sruption to chromatin structure generated by unwinding the parental DNA strands.

280 accepted 'steric exclusion' model for dsDNA unwinding, the active 3' ssDNA strand is pulled through

281 hUPF1 must bind a ssNA segments to initiate unwinding they also raise the possibility that hUPF1 has

282 on, initiating directional target DNA strand unwinding to allow segmented base-pairing with crRNA.

283 of the traditional steric exclusion model of unwinding to also include significant contributions with

284 vere DNA bending, leading to spontaneous DNA unwinding to form a seed-bubble.

285 (Cas) protein Cas9 begin with RNA-guided DNA unwinding to form an RNA-DNA hybrid and a displaced DNA

286 the existing SE model of hexameric helicase unwinding to include contributions from the excluded str

287 ly hindered by H-DNA and has difficulties in unwinding triplex DNA.

288 p that we assign to nonspecific deformation (unwinding/"twisting") of DNA by Rad4.

289 Here, we identify, during RecBCD's DNA unwinding, two Chi-stimulated conformational changes inv

290 case that preserves chromosomal stability by unwinding unimolecular G4 DNA likely to form in transien

291 ecule experiments have demonstrated that the unwinding velocities of some helicases increase dramatic

292 The differing responses of the unwinding velocity and processivity to force have lacked

293 e ATP hydrolysis to nucleic acid binding and unwinding via molecular mechanisms that remain poorly de

294 Brr2-mediated U4/U6 unwinding was strongly inhibited by mutations in U4/U6 d

295 iochemical assays for RNA binding and duplex unwinding, we show that JFH-1 NS3 binds RNA much more ra

296 s unclear how bound proteins influence U4/U6 unwinding, which regions of the U4/U6 duplex the helicas

297 e replisome, must efficiently coordinate DNA unwinding with priming and synthesis to complete duplica

298 The mechanism for coupling DNA unwinding with synthesis is starting to be elucidated, h

299 under replication stress by coordinating DNA unwinding with synthesis of both strands.

300 ere mutated and shown to generally lower DNA unwinding without negatively affecting the ATP hydrolysi