IndraLab

Statements


2 11 4 | 3 156 322

sparser
"Further studies showed an interaction between mTOR and Rheb in the mock-infected cells (Pearson’s co-localization coefficient: 0.95), and FMDV infection enhanced the interaction between mTOR and Rheb (Pearson’s co-localization coefficient: 0.98) ( xref )."

sparser
"It has been reported that binding of mTOR to RAG GTPases and Rheb results in enhanced mTOR activity ( xref )."

sparser
"In mTORC1, Rheb binds to the α-solenoid domain of mTOR, which then induces a conformational change making the α-solenoid move inwards to form a new interaction with the FAT domain."

reach
"In this state, Rheb binds and activates mTOR at the lysosome [41,42,43]."

sparser
"As expected, Rheb and mTOR interacted in the presence of growth factors in the medium ( xref B, lanes 2 and 4)."

sparser
"Meanwhile, the interaction between Rheb and mTOR was abrogated, which agrees with previously published data [ xref ]."

sparser
"This recruitment to the lysosome surface is critical for mTOR-Rheb interaction, allowing Rheb to phosphorylate and activate mTOR ( xref )."

sparser
"MTOR-dependent mechanisms include activation of AMPK/ TSC2 axis by energy depletion [68], activation of TSC2 by regulated in development and DNA damage responses 1 (REDD1) [69] and promyelocytic leukemiamediated (PML) inhibition of mTOR-Rheb-GTP interaction, and consequent nuclear translocation of mTOR [70]."

sparser
"Neither rapamycin nor torin-1 was able to disrupt the physical interaction of mTOR and Rheb, as seen by PLA ( xref )."

reach
"As Rheb, which directly binds and activates mTOR, mediates amino acid induced mTORC1 activity, we hypothesized that RasGRP1 may promote mTORC1 activity by interacting with Rheb in the brain."

sparser
"Similarly, while neither baf nor the LMP-inducer siramesine disrupted the mTOR-Rheb interaction, which is critical for mTORC1 growth signaling, this interaction was completely disrupted by DQ661 ( xref )."

sparser
"Stroke pathology-induced miR-155 upregulation mechanistically disrupts the Rheb and mTOR pathway interaction promoting autophagy and inflammasome activation [ xref ]."

sparser
"The unique ability of DQ661 to disrupt the mTOR-Rheb interaction is therefore not shared by allosteric or catalytic inhibitors of mTORC1, nor by other prototypical lysosomally directed cationic amphiphilic drugs (CADs)."

sparser
"Rheb directly interacts with mTOR Complex1, thus stimulating its ability to signal downstream to S6K1 [ 33 ]."

sparser
"P1_WT bound to RHEB ( K D = 0.14 μM) and inhibited RHEB-mTOR N-heat interaction (IC 50 = 0.33 μM) in vitro."

sparser
"Binding of the mTOR activator, the small GTPase Rheb, causes a motion of the N-heat solenoid (comparable to the ATM Spiral in its location in the primary sequence but not in its structure)."

reach
"Mechanistically, schweinfurthins and their analogs arrested trans-Golgi-network trafficking, an intracellular vesicular trafficking system, resulting in the induction of endoplasmic reticulum stress and the suppression of both lipid raft mediated PI3K activation and mTOR and RheB complex formation, which collectively led to an effective inhibition of mTOR and AKT signaling."

sparser
"In vivo evidence of DQ661-induced disruption of the mTOR-Rheb interaction was demonstrated by PLA ( xref )."

sparser
"However no connection between mTORC1 signaling and ASM inhibition was made in this previous report, and our data indicates that siramesine in nutrient-replete conditions cannot break the mTOR-Rheb interaction in the same manner as DQ661."

sparser
"It was also described that lysosomal MTOR localization brings it in close vicinity to its main regulator, RHEB (Ras homolog, mTORC1 binding), and that as a result, the MTOR-RHEB interaction can activate MTOR kinase activity leading to the phosphorylation of downstream effectors [ xref ]."

sparser
"However, Rag interaction with mTORC1 does not directly activate the mTOR kinase; rather Rag proteins modulate cellular translocation of mTORC1 to the lysosome that facilitates an mTOR and RHEB interaction, which in the presence of elevated AA concentrations, results in RHEB-induced activation of mTORC1."

reach
"Rheb directly binds to mTOR and activates it in a GTP dependent manner."

sparser
"The other is linked to prevention of the interaction between v-ATPase and the Ragulator complex, blockage of MTOR lysosomal recruitment, impairment of MTOR-RHEB interaction and finally the inhibition of MTOR signaling pathway."

reach
"It is activated by Akt and PKB, which phosphorylates tuberin (TSC2) at multiple sites relieving the inhibitory effects of the hamartin-tuberin (TSC1-TSC2) complex on Rheb and mTOR complex 1 (mTORC1) [XREF_BIBR - XREF_BIBR]."

sparser
"Furthermore, in response to anabolic stimuli (resistance exercise and protein feeding), we report a reduction in Rheb-TSC2 colocalization along with a reciprocal increase in mTOR-Rheb colocalization."

sparser
"The results showed that mTOR ΔN bound with RHEB by a K D value of 7.27 μM ( xref C), corresponding to the value reported by the BLI analysis ( K D ∼ 6.5 μM). xref Then, we evaluated the effect of P1_WT to inhibit RHEB-mTOR ΔN interaction by preincubating RHEB with a series of different concentration of P1_WT before adding mTOR ΔN . As a result, P1_WT inhibited RHEB binding to mTOR ΔN with an IC 50 value similar to that of mTORC1 activity inhibition (∼0.3 μM; xref B–C and xref D)."

sparser
"A new modality of cancer therapy has been emerging by blocking PPIs involved in mTORC1 activation. xref In this study, we aimed to inhibit mTORC1 by interfering with the RHEB-mTOR interaction on the lysosome."

sparser
"The large PPI area of RHEB-mTOR limits the efficacy of small molecules for inhibition."

sparser
"Previously, we studied the molecular interactions of RHEB with full-length mTOR and mTOR ΔN , yielding K D values of ∼2.4 and 6.4 μM, respectively. xref In the RHEB-mTOR ΔN interface, α5- and α7-helices of mTOR ΔN stabilize the mTORC1-RHEB complexation followed by the kinase activations. xref The residues (aa 63–79) of RHEB in the switch II region lie between α5- and α7-helices, where α5 interacts with Q72, T73, S75, I76, and D77 residues of RHEB ( xref D), while α7 interacts with the same residues with the extension to bind with N79 and N-terminal resides (K5, S6, R7, and K8; xref E). xref "

sparser
"Once on the lysosome, Rheb binds to the subunit mTOR resulting in a conformational change that exposes active site resides allowing TORC1 to bind and phosphorylate a wide array of substrates ( xref )."

sparser
"Furthermore, we assayed the inhibitory effect of P1_WT for the PPI of RHEB-mTOR ΔN using the AlphaLISA system (PerkinElmer, USA). xref We used the 6xHis tagged RHEB (∼20 kDa) and mTOR ΔN (∼13 kDa) prepared by the BL21(DE3) E. coli overexpression system. xref RHEB was first de-tagged by thrombin and then labeled with biotin using a NHS-(PEG) 24 -biotin reagent."

reach
"RHEB binds to mTOR distally from the kinase active site, yet causes a global conformational change that allosterically realigns active-site residues, accelerating catalysis."

sparser
"Instead, we employed small peptides to disturb the PPI based on the structural characteristics of the α5- and α7-helices of N-heat domain of mTOR, which directly interact with RHEB."

sparser
"This resulted in the improved inhibition of mTORC1 activity by P1_WT compared to that of the other selected variants because P1_WT bound with RHEB at the sub-micromolar level, which corresponds to its inhibitory activity on RHEB-mTOR ΔN interaction."

sparser
"Recent studies illustrate a direct interaction between Rheb and mTOR, which stimulates its kinase activity [14] ."

sparser
"It has also been shown that in contrast to raptor, Rheb binds to the mTOR catalytic domain and that its binding is reversibly inhibited by amino acid withdrawal, regardless of GTP or GDP loading ( Lon[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Inhibition of Rheb binding to mTOR is critical for the inhibitory effect of amino acid withdrawal on mTOR signaling [ xref ]."

sparser
"Alternatively, the ER may be the site of RHEB-mTOR contact but static targeting of RHEB to the ER may be problematic perhaps due to a need for exchange with the prominent cytosolic pool of RHEB to somehow maintain RHEB in an activated state.[ xref ]"

sparser
"To evaluate the effect of P1_WT to inhibit RHEB-mTOR ΔN interaction, we used the AlphaLISA-based assay as previously described. xref , xref First, we evaluated RHEB-mTOR ΔN interaction by mixing different concentrations of 6xHis-mTOR ΔN with 1 μM biotinylated RHEB in a 384-well OptiPlateTM (PerkinElmer, USA) followed by adding 100 μg/mL streptavidin-coated donor beads and 200 μg/mL anti-6xHis-coated acceptor beads."

sparser
"The effect of P1_WT on RHEB-mTOR ΔN interaction was evaluated using the same method with some modifications."

sparser
"Interestingly, the increase in glutathione in P47S cells leads to increased activation of mTOR due to increased Rheb-mTOR association; this leads to improved fitness and muscle recovery in P47S mice, suggesting that this allele may have been selected for at one time ( xref )."

reach
"Rheb binds and regulates the mTOR kinase."

sparser
"While activation of mTORC1 requires GTP-bound Rheb, the interaction of Rheb with mTOR is independent of the nucleotide binding state of Rheb [51,52] ."

sparser
"Rheb can bind to mTOR regardless of its guanylyl nucleotide binding state."

reach
"In addition, Rheb physically interacts with mTOR both at baseline and during GD (XREF_SUPPLEMENTARY), thus indicating that Rheb directly regulates mTORC1 in CMs."

sparser
"While CHAPS can retain the interaction of Rheb and mTOR, NP40 disrupts the interaction."

reach
"Our results indicate that Rheb interacts with mTOR, that Rheb is inhibited during GD and myocardial ischemia, and that its inactivation is required for mTORC1 inhibition."

sparser
"Similarly, culture of myocytes with alcohol for 24 hours antagonizes the ability of leucine to promote phosphorylation of mTOR, S6K1, S6 and 4E-BP1 as well as blunts the interaction between mTOR and Rheb ( xref )."

sparser
"In addition, Rheb physically interacts with mTOR both at baseline and during GD ( xref ), thus indicating that Rheb directly regulates mTORC1 in CMs."

sparser
"In contrast, the interaction between mTOR and Rheb increased with the transfection of the GTP-bound RAGA-Q66L and the GDP-bound RAGC-S75N ( xref )."

reach
"Rheb binds directly to mTOR, inducing conformational changes, suggesting an allosteric mechanism for activating TORC1 [32]."

sparser
"Our results indicate that Rheb interacts with mTOR, that Rheb is inhibited during GD and myocardial ischemia, and that its inactivation is required for mTORC1 inhibition."

sparser
"However, the interaction between Rheb and components of TORC1 is not nucleotide dependent: the GDP-bound or nucleotide-free Rheb interacts with mTOR more strongly than does the GTP-bound Rheb [70,72] [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"The binding of Rheb to mTOR causes a conformational change to expose the mTOR kinase domain to substrate, such as PS6K and 4EBP1, thereby increasing the substrate accessibility of mTORC1 ."

sparser
"Rheb binds directly to the mTOR catalytic domain, and Rheb-GTP enables TORC1 to attain an active configuration."

sparser
"Spinal Rheb-mTOR signal plays an important role in regulation of spinal sensitization in neuropathic pain, and targeting mTOR may give a new strategy for pain management."
| PMC

sparser
"Sancak and coworkers had previously suggested that Rheb localizes to late endosomes, and that amino acid-stimulated mTORC1 translocation facilitates mTOR-Rheb interactions. xref We find that in TSC2 −/− cells overexpressing Rab5CA, endogenous Rheb is hyperactivated but still cannot activate mTORC1."

reach
"TSC2 negatively regulates mTOR by acting as a GTPase-activating protein (GAP) for the small GTPase Rheb, which binds and activates mTOR [18,19] ."

sparser
"RHEB binds mTOR, activating mTORC1 kinase activity."

sparser
"They are unimpaired in their ability to bind GTP or mammalian (m)TOR in vivo or in vitro, and the mTOR polypeptides retrieved with these inactive Rheb mutants exhibit kinase activity in vitro comparable with mTOR bound to wild type Rheb."

sparser
"A follow-up article examines the RhebmTOR interaction and its dependency on extracellular amino acids and/or leucine [98] ."

sparser
"Interaction of Rheb with mTOR has been reported [ xref , xref ]."

sparser
"Effects of amino acid starvation on the interaction of Rheb and mTOR were reported [ xref ]."

sparser
"Moreover, knocking down RAGA or RAGC in WDR24-KO cells, restored the interaction between mTOR and Rheb while decreasing the interaction between TSC2 and Rheb ( xref , xref )."

sparser
"This may agree with the previous observations that the interaction between Rheb and mTOR is rather weak."

sparser
"Rheb interacts directly with mTOR and activates mTOR kinase ( xref ), whereas Rabin8 interacts with Rheb and inhibits the phosphorylation of Ser235/Ser236 in small ribosomal subunit protein S6 ( xref )."

reach
"GTP-bound RHEB directly binds and activates mTOR in the lysosome in response to nutrient levels, resulting in metabolic reprogramming and cell growth ."

sparser
"Our data suggest that the hybrid early/late endosomes produced by overexpression of Rab5CA or knockdown of hVps39 cannot support mTORC1 signaling due to reduced mTOR-Rheb interactions ( xref )."

sparser
"In conclusion, this study indicates that spinal Rheb-mTOR signal paly an important role in regulation of spinal sensitization in neuropathic pain, and suggests that targeting mTOR pathway may provide a new strategy for pain management."
| PMC

sparser
"Alternatively, changes in endosomal composition might lead to conformational changes in endosomal proteins that facilitate mTOR-Rheb interactions."

reach
"Under the induction of amino acids and insulin, the formation of the RHEB and mTOR complex in the skeletal muscle of newborn piglets may be related to the activation of mTORC1 that regulates skeletal muscle protein synthesis [XREF_BIBR]."

sparser
"EtOH also decreased the interaction of Rheb with mTOR, and this was prevented by Leu."

sparser
"Rheb binds directly to mTOR, an interaction that appears to be essential for mTORC1 activation."

sparser
"Co-immunoprecipitation assays of mTOR and Rheb in MC3T3-E1 cells demonstrated that the level of co-immunoprecipitated mTOR was decreased in the presence of D-PDMP or L-PDMP ( Fig. 2 C), suggesting tha[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Recent studies illustrate a direct interaction between Rheb and mTOR, which stimulates its kinase activity [14] ."

sparser
"The contribution of Rheb's regulation of PL-D1 and FKBP38 to mTORC1 activation, relative to Rheb's direct binding to mTOR, remains to be fully defined."

reach
"Rheb binding to mammalian target of rapamycin (mTOR) is regulated by amino acid sufficiency."

sparser
"MTOR-WGA (cell periphery marker) and mTOR-Rheb co-localization was greater in LEAA compared to PLA irrespective of time-point (p<0.05)."

sparser
"Rheb binds directly to the mTOR catalytic domain and activates it in a manner dependent on the state of Rheb nucleotide charging ( xref )."

sparser
"Further supporting this model, it has been reported that AAs promote Rheb-mTOR binding [ xref , xref ]."

sparser
"PRO and KET+PRO increased (Interaction: P <0.0001) mTOR-Rheb co-localization at 120-minutes vs. basal; however, KET+PRO resulted in a sustained increase in mTOR-Rheb co-localization at 300-minutes that was greater than KET and PRO."

sparser
"However, we have been unable to experimentally verify whether NR1 acts by blocking the interaction of Rheb and mTOR due to the well-known technical challenges in detecting this interaction in a definitive manner."

reach
"Suggested leucine mediated mechanisms include promotion of Rheb binding to mTOR, stabilization of the activated mTOR and raptor complex, or mTOR and raptor activation through an alternative class 3 PI3K (hVps34) parallel to that controlled through insulin."

sparser
"It is important to note that when activated, Rheb binds directly to mTOR ( xref ), an interaction that is essential for activation of the mTOR complex 1."

sparser
"The translocation of mTOR results in the impaired interaction between mTOR and Rheb, an mTOR activator, leading to mTORC1 inactivation."

sparser
"Moreover, several studies have demonstrated the role of the RhebmTOR axis in synaptic morphogenesis and axon elongation [ xref , xref ]."
| PMC

reach
"14 In TSC, angiogenesis is thought to be mediated via increased signal transduction through Rheb and mammalian target of rapamycin complex 1 (mTORC1)."

reach
"Inhibition of Rheb binding to mTOR is critical for the inhibitory effect of amino acid withdrawal on mTOR signaling [XREF_BIBR]."

sparser
"Since the localization of mTOR on the LE/Ly membranes is required for the interaction of mTORC1 with its activator Rheb [10,11] , we investigated whether PDMP weakens the interaction between mTOR and [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"GTP-charged Rheb binds directly to the mTOR catalytic domain and activates the mTORC1 cascade, leading to promotion of cell growth [14] ."

sparser
"The activation of the RhebmTOR axis acts as an important signaling pathway for the maintenance of long-term potentiation (LTP) and learning and memory functions [ xref , xref ]."
| PMC

sparser
"Overall, the evidence reported to date suggests that RhebmTOR signaling is associated with neuronal loss that occurs in different neurodegenerative diseases and could therefore be a potential therapeutic target for a wide range of such disorders."
| PMC

sparser
"A mechanistic understanding for the allosteric activation of mTORC1 is now available: Rheb binds to mTOR with its switch I and II regions, which have been previously observed crystallographically to undergo nucleotide-dependent conformational changes ( xref )."

sparser
"The activity of Rheb-mTOR signaling is modulated by both GTPase activity of Rheb and its expression level."

sparser
"Additionally, these results reveal an evolutionary role of Rheb-mTOR in BAT-bone metabolism."

sparser
"However, it is unclear whether imbalance of Rheb-mTOR signaling in BAT will result in bone abnormalities in other metabolic disorders with BAT whitening, such as obesity and type 2 diabetes mellitus. xref , xref "

sparser
"Although S100A8/A9 is known to be secreted mainly by myeloid cells such as neutrophils and macrophages, xref , xref multiple lines of evidence suggest that S100A8/A9 derived from BADs, but not myeloid cells, may play a major role in the suppression of osteogenesis, at least in the initiating stage. (1) Rheb BAD and Rheb AD KO mice displayed markedly enhanced S100A8/A9 expression and secretion in primary BADs in comparison with their control littermates. (2) Rheb BAD mice exhibited markedly higher S100A8/A9 level in mature BAD fraction than in the SVF fraction of BAT, which mainly includes myeloid cells, lymphocytes, preadipocytes, and endothelial cells. xref , xref (3) Both primary BADs and induced C2C12 BADs with Rheb-mTOR inactivation showed markedly increased S100A8/A9 secretion in vitro . (4) Macrophages with Rheb knockout in vitro did not display significant alteration in S100A8/A9 expression. (5) There were far less infiltrating macrophages than BADs in the BAT (less than 5% of CD45 + leukocytes and even lesser macrophages), which does not change during aging. xref Additionally, no significant differences were observed in the number of either total, M1, or M2 macrophages between the control and Rheb knockout mice. (6) We further revealed whitened BAT of aged mice secrete much more S100A8 than bone marrow per se."

reach
"Rheb, like Rac1, directly binds to mTOR independent of the GTP and GDP bound state."

sparser
"Taken together, these data suggest that BAD may serve as one of the primary sources of S100A8/A9 secretion upon BAT dysfunction induced by Rheb-mTOR inactivation."

sparser
"On the contrary, low energy or DNA damage suppresses Rheb activity via activation of TSC. xref , xref Regulation of Rheb-mTOR signaling has been implicated in energy metabolism disorders."

sparser
"Mice with high-fat diet-induced obesity is associated with hyperactivation of Rheb-mTOR signaling. xref Paradoxically, mice with ectopic Rheb expression in pancreatic β cells showed improved glucose tolerance and resistance to hyperglycemia induced by obesity. xref While Rheb activates mTOR signaling to suppress lipolysis, it inhibits white-beige transformation of fat and thermogenesis in an mTORC1-independent but PDE45-CAMP-dependent mechanism, xref further complicating the regulatory role of Rheb-mTOR in metabolism."

sparser
"In this regard, understanding tissue- or cell-type specific role(s) of Rheb-mTOR in these pathological conditions is of fundamental importance to perform targeted inhibition or activation of the pathway without side effects."

sparser
"Combining the results from Rheb ablation in BAD only ( Ucp1-Cre ), or in both BAD and WAD ( Fabp4-Cre ), we demonstrated that Rheb-mTOR inactivation in BAT led to BAT malfunction, outburst of inflammatory batokines including S100A8/A9, and consequent reduction of osteogenesis."

reach
"Moreover, GAPDH also interacts with the member of Ras superfamily of GTPases Rheb, which prevents binding of Rheb to mTOR and thereby inhibits mTOR signaling."

reach
"Whereas the regulators of GTP loading onto Rheb remain to be established, the interaction between Rheb and mTOR is sufficient both in vitro and in vivo to promote the phosphorylation of mTORC1 substrates [XREF_BIBR, XREF_BIBR]."

reach
"Rheb can bind to mTOR regardless of its guanylyl nucleotide binding state."

reach
"The results showed that mTOR bound with RHEB by a K value of 7.27 μM (Figure 4C), corresponding to the value reported by the BLI analysis (K ∼ 6.5 μM)."

reach
"As a result, P1_WT inhibited RHEB binding to mTOR with an IC value similar to that of mTORC1 activity inhibition (∼0.3 μM; Figures 3B–C and 4D)."

reach
"In addition to the interactions between Rheb, Rag GTPases, and mTOR, several protein complexes have been found to act upstream of Rheb and Rag GTPases to control their nucleotide loading status : either by promoting the hydrolysis of GTP and increasing the ratio of GDP/GTP by working as the GAP, or by stimulating the exchange between GDP and GTP by working as the GEF."

sparser
"More importantly, our studies also confirm the key role of Rheb in mTORC1 activation and suggest that the discovery of small molecules that target the Rheb-mTOR protein-protein interaction would be a fruitful avenue to pursue."

reach
"PA decreases AMPK / TSC2 phosphorylation and increases mTOR binding with Rheb and Rag GTPases."

reach
"The addition of PA increased the binding of Rheb with mTOR, while EtOH exposure decreased this interaction (XREF_FIG)."

reach
"There, mTOR interacts with Rheb, resulting in the activation of the mTOR kinase."

reach
"Under low-glucose conditions, GAPDH inhibits mTOR by interacting with the mTOR activator Rheb to prevent the binding of Rheb to mTOR in HEK293 cells or mouse embryonic fibroblasts."

reach
"Because TSC2 functions as a GAP for Rheb, inhibition of its activity promotes the binding of Rheb with mTOR, thereby enhancing mTORC1 function."

sparser
"Long et al. have found that Rheb-regulates mTOR signaling through the direct binding of Rheb-GTP with mTOR [ xref ]."

reach
"However, in terms of leucine stimulation, the latest report by Suryawan et al. found that neonatal pigs fed with a low protein diet supplemented with leucine or its metabolite beta-hydroxy-beta-methylbutyrate (HMbeta) failed to promote the Rheb and mTOR complex formation, but this effect was not observed in pigs fed a high protein diet combined with leucine supplementation."

sparser
"Rheb, like Rac1, directly binds to mTOR independent of the GTP/GDP bound state ( xref )."

sparser
"Ras homolog enriched in the brain (Rheb) is a GTPase, and both mTOR and Rheb share a common pathway because the GTP-binding domain of Rheb unswervingly interacts with the mTOR complex, thereby kindling its kinase activity."

reach
"The action of insulin and growth factors converges on the small GTPase Rheb, which binds directly to the MTOR polypeptide in MTORC1 and, in its GTP bound state, initiates kinase activation."

sparser
"Rheb GTP , not Rheb GDP , binds and activates MTOR."

reach
"Long X, Ortiz-Vega S, Lin Y, Avruch J. Rheb binding to mammalian target of rapamycin (mTOR) is regulated by amino acid sufficiency."

sparser
"2012.06.013 [14] Long X, Lin Y, Ortiz-Vega S, Yonezawa K, Avruch J. Rheb binds and regulates the mTOR kinase."

sparser
"Co-IP analysis of the effect of dhBBR on the binding of exogenous Rheb and mTOR."

sparser
"As shown in Fig.  xref C, Rheb bound to mTOR, and dhBBR inhibited this binding."

sparser
"GTP-loaded RHEB interacts with the mTOR catalytic domain and activates mTORC1 [19] ."

sparser
"Co-IP assay was also used to examine the interaction between Rheb and mTOR."

reach
"The addition of PA to myocytes increased mTORC1 dependent S6K1 phosphorylation, and this was associated with increased binding of mTOR with Rheb and Rag GTPase."

sparser
"Moreover, GAPDH also interacts with the member of Ras superfamily of GTPases Rheb, which prevents binding of Rheb to mTOR and thereby inhibits mTOR signaling."

trips
"They are unimpaired in their ability to bind GTP or mammalian (m)TOR in vivo or in vitro, and the mTOR polypeptides retrieved with these inactive Rheb mutants exhibit kinase activity in vitro comparable with mTOR bound to wild type Rheb."

sparser
"Although direct interaction between Rheb and mTOR has not been ruled out, it appears more likely that Rheb-GTP recruits additional protein(s) to the mTOR complex to modulate its activity."

sparser
"The lead compound they developed is a dimeric quinacrine (DQ661), which is derived from Lys05, has improved lysosomal targeting capability and impairs the activity of palmitoyl-protein thioesterase (PPT1), which is required for mTOR interaction with Rheb at the lysosome [ xref ]; inhibition of PPT1 also prevents mTORC1 from associating with the lysosomal membrane, causing mTOR inhibition [ xref ]."

sparser
"Our data suggest that protective effect of tetrandrine against lung fibrosis might be through promoting Rheb-mTOR and NRF2-SQSTM1 mediated autophagy."

trips
"Rheb binding to mammalian target of rapamycin (mTOR) is regulated by amino acid sufficiency."

reach
"The TOR pathway is regulated by activation of the Ras related protein Rheb, which can bind mTOR."

reach
"For instance, FMDV infection promotes the translocation of mTOR to lysosomes, enhances the interaction between mTOR and Rheb, and activates the PI3K/AKT/TSC2/Rheb/mTOR/p70S6K1 signaling pathway to facilitate viral replication [8]."

reach
"Rheb, a small GTPase, directly associated with mTOR and stimulated the signaling pathway."

sparser
"By contrast, mTOR bound to the RHEB mutant Rheb(Gln64Leu), which is nearly 90% GTP charged, is found predominantly in a GTP bound state and shows higher activity levels."

trips
"Moreover, the binding of mTOR to Rheb mutants that are unable to bind guanyl nucleotide in vivo is also inhibited by amino withdrawal."

sparser
"Akt has major relevance in activating mTORC1 by phosphorylation of TCS2 protein and inhibiting the AMP-PK enzyme that, in turn, activates Rheb and mTOR complex ( xref )."

sparser
"This method was used to characterize the interaction of Rheb and mTOR [ xref ]."

reach
"Active Rheb binds mTOR promoting its activation."

reach
"GTP binding Rheb could bind to the catalytic domain, and then activate the mTOR."

reach
"2009; 296 : E592-602; PMID :18765678; http://dx.doi.org/10.1152/ajpendo.90645.2008[13] Long X, Ortiz-Vega S, Lin Y, Avruch J. Rheb bindingto mammalian target of rapamycin (mTOR) is regu lated by amino acid sufficiency."

sparser
"These protective effects of TET on pulmonary fibrosis are associated with activation of autophagy through promoting NRF2-SQSTM1 axis and Rheb-mTOR signaling."

sparser
"The best characterized mTOR regulator is the tuberous sclerosis complex (TSC), as its activation downregulates the Rheb-mTOR axis under periods of nutrient stress."

sparser
"These scenarios demonstrate how signals transduced through the TSC are mediated by Rheb to affect mTOR, although the precise interactions between Rheb and mTOR have not yet been entirely elucidated."

reach
"Herein we show that the binding of Rheb to endogenous and recombinant mTOR is reversibly inhibited by withdrawal of all extracellular amino acids or just leucine."

reach
"Withdrawal of all extracellular AAs or of leucine alone results in inhibition of the binding of Rheb to mTOR."

reach
"For example, the TSC1 and TSC2 complex inhibits mTOR functions by inactivating Rheb; Rheb binds and activates mTOR only when the former is in its GTP bound form."

reach
"This is in agreement with a previous observation showing that AAs promote the association between Rheb and mTOR."

sparser
"The presence of GAPDH substrate, Gly-3-P, disrupted the interaction between Rheb-GAPDH and increased Rheb-mTOR binding."

sparser
"A potential mechanism of amino acid sensing was suggested by the observation that amino acids, leucine in particular, promote the association of Rheb with MTOR (Long et al. xref )."

sparser
"TET Activated SQSTM1-Mediated Selective Autophagy via Rheb-mTOR Signaling."

reach
"In its GTP bound state, rheb binds to mTOR near its kinase domain and directly stimulates its activity [XREF_BIBR]."

reach
"There is some evidence indicating that Rheb, a small guanosine triphosphate (GTP)-binding protein, interacts with mTOR and activates the enzymatic activity of mTOR by binding the Rheb-GTP to mTOR directly."

sparser
"Taken together, TET activated autophagy through Rheb-mTOR signaling."

sparser
"The binding of IL-37d to Rheb switch-II subregion destabilizes the Rheb-mTOR and mTOR-S6K interactions, further halting mTORC1 signaling."

sparser
"Binding of Rheb to mTOR requires a significant movement of the N-HEAT domain towards the M-HEAT domain ( xref A)."

sparser
"The conformational change induced by Rheb binding precludes the existence of a mixed mTORC1 dimer in which only one mTOR monomer is bound to Rheb ( xref C)."

sparser
"We found that TET can enhance the interaction of SQSTM1 with MAP1LC3-II and ubiquitinated proteins due to NRF2-mediated SQSTM1 transcription and Rheb-mTOR signaling activation, thus dramatically inducing SQSTM1-selective autophagy and directly leading to Col-I degradation in lysosome ( xref )."

sparser
"The binding of Rheb to mTOR causes a conformational change to expose the mTOR kinase domain to substrate, such as PS6K and 4EBP1, thereby increasing the substrate accessibility of mTORC1 xref ."

sparser
"For instance, similar to Rheb binding to mTOR, binding of DNA/Ku70/80 to DNA-PKcs (which together form the DNA-PK complex) has been proposed to stimulate DNA-PKcs activity via an allosteric mechanism that involves the movement of the N-HEAT towards the FAT domain, leading to conformational changes in the KD [ xref ]."

sparser
"Lysosomal IL-37d can suppress Rheb activity independently of TSC2, disrupt Rheb-mTOR interaction, and inhibit interactions of mTOR and its substrate, S6K, thereby terminating mTORC1 activation."

reach
"Together, MA promotes the K48‐linked ubiquitination of Rheb by regulating deubiquitinase USP4 in TDSCs.2.6 MA inhibits the mTORC1 signalling pathway by intercepting the interaction of Rheb and mTOR ."

reach
"IP experiments were performed to investigate whether MA, aside from promoting the degradation of Rheb protein, affects the interaction between Rheb and mTOR to inhibit the mTORC1 signalling pathway."

sparser
"Leucine-enriched amino acids maintain peripheral mTOR-Rheb localization independent of myofibrillar protein synthesis and mTORC1 signaling post-exercise."

reach
"In this context, GR-mTOR crosstalk might be a key for creating an interdisciplinary research area that bridges nutrition and medicine.The biochemical rationale for the usage of BCAA as a therapeutic t[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Rheb binds directly to mTOR which is a required step in mTORC1 activation."

reach
"Further support for the interaction between Rheb and mTOR comes from FRET-FLIM imaging studies."

sparser
"The mTOR pathway senses amino acids through the Rag family of GTPases and the TSC1/2 complex to promote the interaction of mTOR with Rheb( xref ; xref )."

sparser
"Rheb binds directly to mTOR, inducing conformational changes, suggesting an allosteric mechanism for activating TORC1 [ xref ]."

sparser
"Spinal Rheb-mTOR signal plays an important role in regulation of spinal sensitization in neuropathic pain, and targeting mTOR may give a new strategy for pain management."

sparser
"The third mechanism blocks the interaction mTOR-RHEB through the proteins Bcl2/adenovirus E1B 19-kDa protein-interacting protein 3 (BNIP3) [ xref ], and promyelocytic leukemia protein (PML) [ xref ]."

sparser
"We found tunicamycin treatment has abolished the RhebmTOR interaction but the Rheb–PERK interactions remains unaffected ( xref )."

sparser
"Rheb interacts with mTOR’s catalytic domain, and when in its GTP-bound state, Rheb can directly activate mTOR kinase activity [ xref , xref ]."

sparser
"Thus, depending upon the cellular status, such as ER stress, Rheb-mTOR interaction would be dismantled, but Rheb-PERK interactions might be stabilized."

sparser
"A canonical cascade for protein synthesis involves interactions of Rheb with mTOR to activate the mTORC1 protein synthesis pathway ( xref )."

sparser
"The gene products of TSC1 (called hamartin) and TSC2 (called tuberin) assemble the TSC1/2 complex, which physiologically suppresses mTOR by inhibiting Rheb, an essential activator of the mTORC. xref Patients with TSC present with epilepsy early in life and have the characteristic presence of “tubers,” tumorlike focal lesions all over the body including in brains. xref Children with TSC can present with sleep abnormalities such as increased incidences of night waking, parasomnias, severe difficult waking up early in the morning, and daytime sleepiness. xref These abnormalities were not significantly different between epileptic and nonepileptic subgroups, representing inherent sleep and circadian disruptions in TSC patients. xref In a polysomnography study on children with TSC, sleep architecture was found to be severely disrupted, with sleep fragmentation, a shorter total sleep time, reduced sleep efficiency, and decreased rapid eye movement sleep. xref Interestingly, in this study, nocturnal seizures were recorded in some patients and associated with a more severe sleep fragmentation. xref In an adult cohort of TSC patients, similar sleep abnormalities were found, namely insomnia and excessive daytime sleepiness, which, unlike in the pediatric group, was positively correlated with their seizure history and antiseizure drug use. xref To illustrate the scale of sleep abnormalities in TSC, a large natural history study of more than 2000 patients with TSC reported sleep abnormalities as the second most reported behavioral problem in about 40% of the patients. xref Thus, it is clear that patients with TSC present with chronic sleep abnormalities, regardless of their seizure history, which indicates an intrinsic circadian dysfunction."

sparser
"Whether phosphorylation or other post-translational modifications, such as acetylation, differentiate Rheb’s interactions with mTOR vs. PERK remains to be seen."

sparser
"Rheb binds and regulates the mTOR kinase."

sparser
"Therefore, we further evaluated IL-37d role in controlling Rheb-mTOR interaction."

sparser
"Contrary to expectation, distinct from TSC2, IL-37d inhibited both the interactions of Rheb-mTOR and mTOR-S6K, suggesting that the binding of IL-37d to Switch II of Rheb leads to mTOR disassociations from Rheb and its substrate, thereby facilitating mTOR signal termination (Fig.  xref )."

sparser
"Herein, we proved that IL-37d suppressed lysosomal Rheb activity, Rheb-mTOR interaction, and mTOR substrate recruitment, thereby terminating mTORC1 activation."

reach
"MTOR can bind Rheb regardless of its GDP/GTP content, even though only GTP-Rheb can activate mTORC1 activity [6,32]."

reach
"Recruitment of mTOR to the surface of the lysosome promotes the activation of mTOR through phosphorylation by lysosome bound RHEB."

sparser
"Studies in Rheb knock-out mice showed that RHEB activity is the rate-limiting step for mTOR activation in the brain and that neuronal functioning in particular is sensitive to increased RHEB-mTOR signaling [ xref ]."

sparser
"The evidence for an association of Rheb with mTOR was obtained by using GST-Rheb proteins as baits to pull down endogenous mTOR."

reach
"MTOR interaction with Rheb is restricted in basal conditions due to the enzymatic activity of the tuberous sclerosis complex proteins TSC1 and TSC2, which maintain Rheb in a GDP bound state 16."

reach
"These scenarios demonstrate how signals transduced through the TSC are mediated by Rheb to affect mTOR, although the precise interactions between Rheb and mTOR have not yet been entirely elucidated."

sparser
"As previously reported, the binding ability of Rheb to mTOR is independent of its GTP- or GDP-status but its switch II segment plays a key role in promoting Rheb-mTOR direct binding xref , xref ."

reach
"Rheb binds and regulates the mTOR kinase."

sparser
"We found that, distinct from TSC2, IL-37d disrupts interactions of Rheb-mTOR and mTOR-S6K, thereby further enhancing the suppression of mTORC1 via targeting the Rheb switch II subregion."

reach
"MTOR can be activated by : (i) growth factors, through the PI-3kinase and Akt pathway; 91 (ii) mechanical loading, through the removal of the inhibitor TSC2 from the mTOR and Rheb complex; 92 and (iii) feeding, through the GATOR/Rag/Ragulator pathway."

sparser
"Moreover, the switch II subregion of Rheb can also play a unique regulation of mTOR by recruiting other unknown coactivators, which is independent of Rheb activity and Rheb-mTOR interaction."

sparser
"The modulation of the TSC complex on Rheb-mTOR is generally considered to be necessary for mTORC1 activation when responding to amino acid stimulation."

sparser
"However, in terms of leucine stimulation, the latest report by Suryawan et al. found that neonatal pigs fed with a low-protein diet supplemented with leucine or its metabolite β-hydroxy-β-methylbutyrate (HMβ) failed to promote the RhebmTOR complex formation, but this effect was not observed in pigs fed a high-protein diet combined with leucine supplementation ( xref )."

sparser
"All three regulatory events inhibit the interaction of mTOR with Rheb, which in turn reduces the phosphorylation/activation of S6K. Inhibition of the mTOR/S6K pathway through these mechanisms results in decreased translation of HIF-1 α and expression of VEGF [ xref – xref ]."

sparser
"In view of the fact that the human experimental studies by xref and xref have observed consistent results, there seems to be reason to believe that both aerobic exercise and resistance exercise can downregulate the expression of TSC2, promote the dissociation of TSC2-Rheb, and enhance the lysosomal co-localization of mTOR-Rheb, which subsequently inhibits the conversion of Rheb to inactive state loaded with GDP and promotes the activation of mTORC1."

sparser
"As mentioned above, GTP-Rheb interaction with mTOR is an important way for activation of mTOR on the lysosomal surface."

sparser
"Moreover, RagC phosphorylation is associated with destabilization of mTORC1 (Yang et al , xref ), while Rheb directly binds to mTOR in its GTP bound, which is required for mTORC1 kinase activity (Saci et al ,  xref )."

reach
"TSC negatively regulates the GTPase Ras homolog enriched in brain (RHEB), which associates with mammalian target of rapamycin complexes (mTOR) and promotes mTORC1 signaling, translation, and cell grow[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Considering the characteristic morphologies of mitochondria and Golgi, it is apparent that mTOR is not present on these organelles.Since the localization of mTOR on the LE/Ly membranes is required for[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"The translocation of mTOR results in the impaired interaction between mTOR and Rheb, an mTOR activator, leading to mTORC1 inactivation."

sparser
"Mutants of Rheb that cannot bind GTP or GDP can interact with mTOR complexes."

sparser
"The binding of the GDP:Rheb complex to mTORC1 inhibitors mTOR."

reach
"The TSC heteromeric complex formed by TSC1, TSC2 and TBC1D7 acts as a GTPase activating protein (GAP) for the small Ras related GTPase Rheb, which, in active GTP bound form, binds and activates mTOR."

sparser
"Meanwhile, upstream of mTORC1 and mTORC2, Ras homolog enriched in brain (Rheb), a GTPase, also directly binds mTOR but results in the activation of mTORC1 [ xref ]."

reach
"When PDE4D binds Rheb it inhibits the ability of Rheb to activate mTORC1 and hence it is plausible that PDE4DIP (also called myomegalin) impacts on this relationship, as an increased interaction between Rheb and mTOR should promote growth."

sparser
"Localization of RHEB and mTORC1 to the surface of lysosomes facilitates RHEB-mTOR binding."

sparser
"Rheb GTPase directly binds to mTOR and activates mTORC1 in response to growth factor signals [ xref ]."

sparser
"Rheb-GTP binds to mTOR complex 1 (mTORC1) to activate its kinase activity toward the S6Ks, 4E-BP1, and other substrates, leading to enhancement of multiple anabolic biosynthetic pathways that enable production of the building blocks (e.g. nucleotides) and macromolecules (e.g. ribosomes) required for cell size increase and mitosis ( xref )."

sparser
"The nucleotide binding state and the interaction of Rheb with mTOR is coordinately controlled by the tuberous sclerosis proteins TSC1 (aka hamartin) and TSC2 (aka tuberin)."

sparser
"Rheb binds and regulates the mTOR kinase."

reach
"Recent data obtained by cryo-electron microscopy suggested that binding of Rheb to the mTOR kinase allosterically realigned active-site residues and relieved the auto-inhibition [ 40 ]."

sparser
"The TOR pathway is regulated by activation of the Ras-related protein Rheb, which can bind mTOR."

sparser
"We showed that this increase in glutathione directly regulates the interaction of mTOR with Rheb; consequently, human and mouse cells containing the S47 variant show increased mTOR activity ( xref )."

sparser
"Our results demonstrate that Rheb regulation of mTOR signaling occurs, at least in part, through the direct binding of Rheb-GTP to mTOR, an interaction that promotes activation of the mTOR kinase acti[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Also, PML was identified as a repressor of mTOR through inhibition of Rheb-mTOR interaction in hypoxia ( xref ) [ xref ]."

sparser
"Thus, PML might oppose mTOR activity by controlling Akt, Rheb-mTOR interaction and/or PTEN in stem cells ( xref )."

sparser
"Notably, the inactive Rheb switch 1 mutant Thr38Met binds endogenous mTOR in a manner equivalent to wild-type GST-Rheb, whereas mTOR binding to Rheb Ile39Lys and to GST-Rheb Gln64Leu is reduced as com[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"No binding of Rheb to mTOR 1–670 and 1–1967 was observed; however, the mTOR fragments 1967–2191, encompassing the FKBP12-rapamycin binding domain, and 2148–2549, encompassing the kinase catalytic doma[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"GTPases primarily responsible for signal transduction include the Ras and Rho subfamilies, while Rheb and RagA/Gtr subfamily GTPases interact with the mTORC nutrient sensing machinery."

sparser
"TP53 missense mutations also play a gain-of-function role in stimulating mTORC1 activation by favoring the interaction between mTOR and Rheb [ xref ], an upstream positive regulator of mTOR activity [ xref ]."

sparser
"In fact, charging with GTP appears to substantially diminish the binding of Rheb to mTOR 2148–2549 as well as to the c-Raf1 1–257 fragment, as compared to nucleotide-free Rheb."

sparser
"Moreover, the diminished binding of mTOR 2148–2549 to GTP-charged Rheb as compared to nucleotide-free or GDP-charged Rheb is also consistent with the lesser ability of recombinant Rheb(Asn64Leu) as co[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Studies have shown that FMDV-infected PK15 cells promote the transfer of mTOR to lysosomes to enhance the interaction between mTOR and Rheb and activate the PI3K/AKT/TSC2/Rheb/mTOR/p70S6K1 pathway to promote viral replication [52], which is consistent with the results of RNA‒seq in this study.Interleukin, interferon, the tumour necrosis factor superfamily and other cytokines act in concert with specific cytokine inhibitors and soluble cytokine receptors to regulate the human immune response [53]."

sparser
"Key enzymes in the glucose metabolism pathway, hexokinase and GAPDH, have been shown to interact with mTOR or Rheb and decreases mTOR signaling when glucose is low ( xref , xref )."

sparser
"Rheb binding to its putative effector mTOR does not require Rheb guanyl nucleotide charging."

sparser
"Recombinant Rheb binds to the mTOR catalytic domain specifically, in the sense that other small GTPases most closely related to Rheb (i.e., Ras and Rap1) exhibit much weaker binding to mTOR."

sparser
"As expected, amino acid replenishment enhanced the interaction between endogenous Rheb and mTOR ( xref )."

sparser
"Surprisingly, the ability of Rheb to bind mTOR does not require Rheb nucleotide charging; in fact, nucleotide-free Rheb binds to mTOR more tightly than does Rheb-GTP, both in vivo and in vitro."

sparser
"A number of studies have looked at the involvement of Rheb in the cellular response to amino acids (Garami et al , xref ; Inoki et al , xref ; Tee et al , xref ; Zhang et al , xref ; Long et al , xref ; Smith et al , xref ; Roccio et al , xref ; Bai et al , xref ; Sancak et al , xref ), with some disagreement on whether amino acids affect Rheb GTP‐loading or RhebmTOR binding."

reach
"Akt has major relevance in activating mTORC1 by phosphorylation of TCS2 protein and inhibiting the AMP-PK enzyme that, in turn, activates Rheb and mTOR complex (75)."

sparser
"Amino acid stimulation also increased the level of Ub-Rheb and enhanced Rheb-mTOR interaction in the high-density cell fraction (HDM), which contains lysosome ( xref )."

sparser
"Reciprocally, mTOR polypeptides bound to recombinant Rheb(Gln64Leu), a mutant reported to be approximately 90% GTP charged in vivo [ 40 ], exhibit a substantially higher kinase specific activity in vi[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"The association of mTOR with GTP-charged Rheb increases mTOR kinase activity."

sparser
"Based on these results, we propose that the association of mTOR with Rheb-GTP is a necessary step in the physiologic activation of the mTOR kinase."

reach
"Binding of Rheb to mTOR requires a significant movement of the N-HEAT domain towards the M-HEAT domain (Figure 4A)."

sparser
"We find that amino-acid withdrawal inhibits the binding of Rheb to mTOR (X.L., S.O.-V., Y.L., and J.A., unpublished data)."

sparser
"If the association of endogenous Rheb with mTOR is required for the maintenance as well as for the initiation of TOR kinase activation, then the ability of amino-acid withdrawal to inhibit the Rheb-mT[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"If, however, the Rheb-mTOR interaction is only necessary for the initiation of mTOR kinase activation, then some additional site of Rheb action is required, within the TOR complex or elsewhere, to exp[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Previous studies demonstrated that the interaction between Rheb and mTOR was independent of Rheb’s GTP/GDP-loading status in vivo ( xref ; xref ), and the GDP-bound Rheb D60I mutant displayed a higher binding affinity to mTOR compared to wild-type or GTP-bound Rheb mutants in vivo ."

sparser
"Although Rheb GTP charging weakens the association of Rheb with the mTOR catalytic domain, the configuration of GTP-charged Rheb enables the TOR complex to adopt a form that is both catalytically acti[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Thus, there may be one or more Rag, Ragulator, and/or endosomal mechanisms by which sepsis disturbs the interaction of mTOR and Rheb culminating in leucine resistance."

reach
"It has been demonstrated that the small GTPase Rheb (Ras homolog enriched in brain) directly interacts with and activates mTOR [XREF_BIBR]."

sparser
"Rheb directly interacts with mammalian target of rapamycin to increase its activity."

reach
"To test the possibility that Rheb acts directly on mTOR, we first examined whether Rheb is capable of binding mTOR."

reach
"The results illustrate the power of GFP-technology combined with FRET-FLIM imaging in the study of the interaction of signalling components in living cells, here providing evidence for a direct physical interaction between mTOR and Rheb and between mTOR and raptor in living cells for the first time."

reach
"As shown in Figure 6C, many red dots were observed in control OSCC cells, suggesting the vibrant interaction between mTOR and RHEB and activation of the mTOR pathway."

reach
"From the PLA assay, the reduced interaction of mTOR and RHEB was observed more intuitively."

reach
"Rheb binds directly to mTOR which is a required step in mTORC1 activation."

sparser
"Upstream of mTORC1, feeding increased the association of mTOR with Rheb, which together with Akt phosphorylation are indices of insulin signaling, in the LD muscle in the INT-PP group compared with the INT-PA group ( P < 0.01; xref )."

reach
"In fact, charging with GTP appears to substantially diminish the binding of Rheb to mTOR 2148-2549 as well as to the c-Raf1 1-257 fragment, as compared to nucleotide-free Rheb."

reach
"It is also possible that Rheb may bind to and activate mTOR interacting proteins such as rictor, raptor or mLST8 rather than interacting with and activating mTOR directly [XREF_BIBR]."

reach
"Surprisingly, the ability of Rheb to bind mTOR does not require Rheb nucleotide charging; in fact, nucleotide-free Rheb binds to mTOR more tightly than does Rheb-GTP, both in vivo and in vitro."

reach
"Reciprocally, mTOR polypeptides bound to recombinant Rheb (Gln64Leu), a mutant reported to be approximately 90% GTP charged in vivo [40], exhibit a substantially higher kinase specific activity in vit[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"It is interesting to note that both Rheb and raptor interact strongly with mTOR in live cells as demonstrated in previous studies 21."

reach
"We find that amino-acid withdrawal inhibits the binding of Rheb to mTOR (X.L., S.O.-V., Y.L., and J.A., unpublished data)."

reach
"Hence the dynamic aspects of the interaction between mTOR, Rheb and raptor in living cells requires new approaches for the detection of interaction, such as provided by the fluorescence resonance energy transfer - fluorescence lifetime imaging (FRET-FLIM) technique for observation of appropriately labelled materials in an active environment [XREF_BIBR - XREF_BIBR]."

reach
"Using the sensitive advanced imaging technique of time correlated single photon counting coupled with fluorescence lifetime imaging and molecular GFP fusion technology, it was possible for the first time to probe directly the nature of the interaction between Rheb, mTOR and raptor as well as raptor and mTOR whilst providing new insights into their sub-cellular localization."

sparser
"The association of Rheb and mTOR is promoted by amino acids availability xref ."

sparser
"The phosphorylation and inhibition of TSC2 cause TSC to release its inhibitory hold on mTOR, allowing mTOR to associate with Rheb on the lysosomal membrane, leading to mTORC1 activation ( xref ; xref ; xref )."

sparser
"The effect of amino acids in enhancing the ubiquitination of Rheb and its binding to mTOR was diminished in the Rheb-4Rs mutant ( xref ), suggesting that amino acids promote the Rheb-mTOR interaction by enhancing the polyubiquitination of at least the four lysine residues."

reach
"Structural studies show that RHEB loaded with GTP binds to mTOR distally from the kinase site and causes a global conformational change of the complex [30]."

reach
"Intracellularly, schweinfurthins and their analogs arrested trans-Golgi-network trafficking, likely by binding to oxysterol-binding proteins, leading to an effective inhibition of mTOR/AKT signaling through inducing endoplasmic reticulum stress and suppressing both lipid raft-mediated PI3K activation and mTOR/RheB complex formation."

reach
"Moreover, RagC phosphorylation is associated with destabilization of mTORC1 (Yang et al, 2018), while Rheb directly binds to mTOR in its GTP bound, which is required for mTORC1 kinase activity (Saci et al, 2011)."

sparser
"However, the activation of this pathway is blunted in skeletal muscle of the preterm piglet, with hyperinsulinemic-euaminoa cidemic-euglycemic clamps showing lower phosphorylation of Akt and TSC2 and reduced Rheb-mTOR association in response to insulin in muscle of preterm compared to term neonates ( xref )."

reach
"The average excited state lifetime of the EGFP-mTOR in the presence of DsRed-Rheb was determined to be 2200 +/- 50 ps in three independent experiments, the reduced level due to a direct interaction between Rheb and mTOR, due to energy transfer from EGFP to DsRed."

reach
"However, Rag interaction with mTORC1 does not directly activate the mTOR kinase; rather Rag proteins modulate cellular translocation of mTORC1 to the lysosome that facilitates an mTOR and RHEB interaction, which in the presence of elevated AA concentrations, results in RHEB induced activation of mTORC1."

reach
"Quenching in the lifetime of the donor (EGFP-mTOR) from 2400 +/- 100 ps to 2000 +/- 100 ps in the nuclear regions indicates that EGFP-mTOR and DsRed-Rheb interact (due to excited state energy transfer) differently (i.e. they are closer) in the cell nucleus than in the cytoplasm, likely brought about by a different conformation of the mTOR and Rheb complex."

sparser
"This binding does not require GTP and nucleotidedeficient RHEB mutants can bind mTOR in vivo and in vitro."

reach
"This study is the first to show a direct interaction between Rheb and mTOR in specific regions of living cells."

sparser
"Opposite, the anti-apoptotic ATF6 pathway, which was shown to enhance tumor cell survival through activation of the Rheb-mTOR signaling pathway xref or by induction of GRP78 expression xref , has resulted inhibited."

reach
"The final step in this activation process is dependent on Rheb, a small GTPase that binds to mTOR and allosterically activates its kinase activity."

sparser
"The effect of Leu on LD muscle was independent of any role of insulin or insulin signaling upstream of mTORC1 because plasma insulin concentrations, Akt phosphorylation, and the association of mTOR with Rheb were not different among groups administered the Ala and Leu pulses."

sparser
"The less mTOR interacts with RHEB, the less mTOR is phosphorylated."

sparser
"IP experiments were performed to investigate whether MA, aside from promoting the degradation of Rheb protein, affects the interaction between Rheb and mTOR to inhibit the mTORC1 signalling pathway."

sparser
"In the presence of MG132, which inhibited proteasome activity to block the protein degradation pathway, MA markedly suppressed the interaction between mTOR and exogenous Rheb in HEK293T cells (Figure  xref )."

sparser
"This effect was validated in TDSCs, where MA significantly inhibited the interaction between endogenous Rheb and mTOR induced by IL‐1β (Figure  xref )."

reach
"The active Rag complex (composed of guanosine triphosphate [GTP]-bound RagA/B and guanosine diphosphate [GDP]-bound RagC/D) binds to Raptor of mTORC1 and recruits mTOR to the lysosome, where it binds to active GTP bound Rheb 23 to inhibit autophagy."

sparser
"These results indicate that MA inhibits the mTORC1 signalling pathway by intercepting the interaction of Rheb and mTOR."

sparser
"Activation most likely requires a direct interaction between Rheb and mTOR, but the exact mechanism remains unclear."

reach
"However, we have been unable to experimentally verify whether NR1 acts by blocking the interaction of Rheb and mTOR due to the well-known technical challenges in detecting this interaction in a definitive manner."

reach
"Consequently, we demonstrated that RHEB binds to the whole mTOR both in the presence or absence of GTPgammaS, with five-fold weaker affinity in the presence of GTPgammaS."
| PMC

reach
"The stoichiometry obtained, however, did not involve the kinetics of RHEB binding to mTOR by means of protein–protein interaction (PPI) assays [15,16]."
| PMC

sparser
"The investigation demonstrated that MA inhibited the mTORC1 signalling pathway by promoting the degradation of Rheb protein and intercepting interaction between Rheb with mTOR."

sparser
"Consequently, we demonstrated that RHEB binds to the whole mTOR both in the presence or absence of GTPγS, with five-fold weaker affinity in the presence of GTPγS. In addition, RHEB bound to the truncated mTOR fragments of N-heat domain (∆N, aa 60–167) or M-heat domain (∆M, aa 967–1023) with the same affinity in the absence of GTP."
| PMC

reach
"As a result, we observed that RHEB bound to mTOR in the presence or absence of GTPγS (Merck Millipore, Darmstadt, Germany), although only RHEB–GTP was shown to activate mTORC1 [4]."
| PMC

sparser
"On the other hand, growth factors and nutrients promote the binding of the RHEB to mTOR, thereby promoting the kinase activity of mTORC1 [ xref ]."
| PMC

sparser
"The stoichiometry obtained, however, did not involve the kinetics of RHEB binding to mTOR by means of protein–protein interaction (PPI) assays [ xref , xref ]."
| PMC

reach
"When the utilization rate of blood glucose decreases, ATP synthesis decreases, while AMP increases, increased AMP binds and activates AMPK, following serine at positions 1345 of tumor suppressor (TSC2) is phosphorylated by activated AMPK and has GTPase activity, leading to GTP hydrolyze to GDP, which weakens the binding of GTP-dependent Rheb to mTOR molecules, exerting the effect of inhibiting mTOR, thereby enhancing the autophagic activity of cells."

reach
"BLItz Measurements of RHEB Interactions with mTOR Truncates."
| PMC

sparser
"As a result, we observed that RHEB bound to mTOR in the presence or absence of GTPγS (Merck Millipore, Darmstadt, Germany), although only RHEB–GTP was shown to activate mTORC1 [ xref ]."
| PMC

reach
"The results show that the binding of RHEB to mTOR involves multiple binding sites with a variety of biding affinities, suggesting that RHEB regulates the kinase activity of mTOR through multiple mechanisms."
| PMC

sparser
"The activity of mTORC1 in mammalian cells is critically dependent on the small GTPase Rheb, which binds directly to the mTOR catalytic domain and, in its GTP-bound state, promotes mTORC1 catalytic activity ( xref )."

sparser
"Amino acid withdrawal does diminish the binding of recombinant Rheb to mTOR."

sparser
"Biochemical studies have demonstrated that active Rheb directly associates with mTOR and potently stimulates its kinase activity in vitro xref , xref , xref ."

reach
"Rheb binds and regulates the mTOR kinase."

sparser
"As a result, RHEB bound to all the mTOR fragments with different affinities irrespective to the endogenous GTP levels."
| PMC

sparser
"The cryo-EM analysis revealed that RHEB interacted with three different mTOR fragments of aa 60–167 in N-heat domain (∆N), aa 967–1023 in M-heat domain (∆M), and aa 1240–1360 in FAT domain (∆FAT) ( xref a) [ xref ]."
| PMC

reach
"RHEB bound to whole mTOR with 5 times weaker affinity in the presence of GTP than in its absence."
| PMC

sparser
"RhebmTOR Signaling in the Brain."

reach
"We observed that Rheb is associated with LEs in hippocampal neurons and recent structural studies suggest a central position of activated Rheb in the complex associating Rag and mTORC1 to the lysosome surface [24, 31, 32] and show that Rheb binds to mTOR at a site remote from its kinase active site resulting in a conformational change that favors mTOR activity [26]."

sparser
"RHEB bound to whole mTOR with 5 times weaker affinity in the presence of GTP than in its absence."
| PMC

sparser
"The results show that the binding of RHEB to mTOR involves multiple binding sites with a variety of biding affinities, suggesting that RHEB regulates the kinase activity of mTOR through multiple mechanisms."
| PMC

sparser
"Direct binding of Rheb-GTP to the mTOR kinase subunit of mTORC1 allosterically activates the kinase by inducing a large-scale conformational change xref ."

sparser
"Rheb:GTP, but not Rheb:GDP, binds mTOR and stimulates its protein kinase activity [ xref ]."

reach
"Importantly, under unstressed physiological condition, endogenous SAG bound to endogenous RHEB, but neither SAG nor RHEB binds to UBE2F or mTOR in these liver cancer cell lines (Fig. 4A,B)."

sparser
"The mTOR-Rheb interaction is largely maintained in both intermediate and active states, however, the residues Met1255 and Lys1256 of the FAT domain of mTOR are disordered in the intermediate state but engaged with Rheb in the active state ( xref )."

reach
"Moreover, interaction between Rheb and mTOR was also sensitive to FTI-277 treatment, whereas Rheb was not found to associate with GbetaL under tested conditions (data not shown)."

sparser
"For instance, FMDV infection promotes the translocation of mTOR to lysosomes, enhances the interaction between mTOR and Rheb, and activates the PI3K/AKT/TSC2/Rheb/mTOR/p70S6K1 signaling pathway to facilitate viral replication [ xref ]."

reach
"This strongly implicates the Golgi as the platform for the assembly of the functional Rheb and mTOR complex."

reach
"Thus, one possible explanation is that sleep deprivation reduces cAMP, which would elicit more PDE4D interaction with Rheb and thereby decrease the interaction between Rheb and mTOR."

sparser
"Because overexpression of Rheb, but not hyperactivation of endogenous Rheb, can rescue the inhibition of mTORC1 by Rab5CA, we infer that the localization of mTOR to these hybrid endosomes decreases the interaction of mTOR with endogenous Rheb ( xref )."

sparser
"Rheb, a small GTPase, directly associated with mTOR and stimulated the signaling pathway."

sparser
"This explains why overexpressed Rheb activates mTOR in the absence of amino acids; presumably the Rheb-mTOR interaction is driven by mass action in this case, and no longer requires the amino acid–stimulated movement of mTOR to late endosomes."

sparser
"These data suggest that overexpressed Rheb can interact with mTOR in the hybrid endosomes formed in Rab5CA-expressing cells, whereas endogenous Rheb cannot."

sparser
"Unfortunately, attempts to measure the binding of endogenous mTOR and Rheb has been unsuccessful even in control cells, and the localization of endogenous Rheb by fractionation or immunofluorescence has not yet been achieved, as has been described elsewhere ( xref ; xref )."

sparser
"Adding credence to this model are recent biochemical data showing that GTP-bound Rheb can bind to and activate mTOR in an amino-acid-sensitive manner [48–50] ."

reach
"Inhibition of TSC2 and PRAS40 allows the small GTPase Ras homolog enriched in brain (Rheb) to bind to mTOR and increase mTOR kinase activity (XREF_FIG)."

reach
"Amino acid stimulation may also convert Rheb guanosine diphosphate to Rheb-guanosine triphosphate, thereby enhancing the association between Rheb and mTOR, and resulting in mTOR activation."

sparser
"Recent data obtained by cryo-electron microscopy suggested that binding of Rheb to the mTOR kinase allosterically realigned active-site residues and relieved the auto-inhibition [ 40 ]."

sparser
"Although it has been proposed that the GTP-bound, active from of RHEB physically binds to mTORC1 and enhances its kinase activity [ xref ], the detailed molecular mechanism of mTORC1 activation by RHEB has been elusive until recently; the cryo-electron microscopy (cryo-EM) structure of the mTORC1-RHEB complex has revealed that RHEB directly binds to mTOR distantly from the kinase catalytic site and causes a conformational change that allosterically realigns the catalytic site, thereby enhancing mTORC1 activity [ xref , xref ]."

sparser
"A recent study has shown that mTOR is required to interact with late endosomes that are enriched in TMEM127 proteins, as binding to the mTOR activators Rheb and PA is necessary for its activation. [ xref ]."

sparser
"Then the activated Rheb interacts with mTOR to regulate downstream molecules ( Goncharova et al., 2011; Mills et al., 2008 ) ( Table 2 )."

reach
"GAPDH regulates the binding of Rheb to mTOR in a manner that is dependent upon glycolytic intermediates and is independent of the nucleotide charged status of Rheb."

sparser
"Using this approach, we first immunohistochemically probed levels of phospho-S6 (pS6; a ribosomal protein), a well-established marker for RHEB-mTOR activity ( xref ), in the NAc of the Rheb knockout and control mice ( xref )."

sparser
"Herein we show that the binding of Rheb to endogenous and recombinant mTOR is reversibly inhibited by withdrawal of all extracellular amino acids or just leucine."

sparser
"Moreover, the binding of mTOR to Rheb mutants that are unable to bind guanyl nucleotide in vivo is also inhibited by amino withdrawal."

reach
"Long X, Ortiz-Vega S, Lin Y, Avruch J. Rheb binding to mammalian target of rapamycin (mTOR) is regulated by amino acid sufficiency."

reach
"From these studies, a revised Rheb binding to mTOR is proposed to include Rheb nuclear localization."

reach
"In summary, Rheb, mTOR and raptor mainly reside on Golgi/ER like structures, where mTOR directly interacts with both Rheb and raptor, shown here for the first time in living cells using FRET-FLIM methodology."

sparser
"The subsequent Akt-mediated phosphorylation and inactivation of tuberous sclerosis complex 2 enables Ras homolog enriched in brain (Rheb) to associate with mTOR on the lysosomal membrane, leading to mTORC1 activation [ xref ]."

reach
"Rheb and mammalian target of rapamycin complex 1 (mTORC1) pathway is chronically activated in TSC deficient cells, and this activation can be diminished using the appropriate inhibitors."

reach
"It has been reported that delayed post-Golgi trafficking induces ER stress, suppresses both PI3K activation and mTOR/RheB complex formation, and subsequently leads to the inhibition of PI3K/AKT/mTOR signaling."

reach
"Notably, the apo and Rheb bound mTOR conformations are incompatible in a mixed dimer."

reach
"Activation most likely requires a direct interaction between Rheb and mTOR, but the exact mechanism remains unclear."

sparser
"PPT1 has been reported to regulate lysosomal function and is essential for the mTOR-RHEB (Ras homolog enriched in brain) interaction on the lysosomal membrane, which allows RHEB to phosphorylate and activate mTOR ( xref )."

sparser
"We first immunohistochemically probed levels of phospho-S6 (pS6; a ribosomal protein), a well-established marker for RHEB-mTOR activity ( xref ), in the NAc of the Rheb knockout and control mice ( xref )."

sparser
"In this state, Rheb binds and activates mTOR at the lysosome [ xref , xref , xref ]."

sparser
"Further, the development of new tools that enable combined in vivo single-neuron calcium imaging with CRISPR perturbations will be crucial to resolving how RHEB-mTOR signaling processes both appetitive and aversive aspects of motivation at the level of neural dynamics ( xref )."

sparser
"This allows for interaction between Rheb and mTOR to occur due to a change in location of the TSC2 protein such that it cannot interact with its target, Rheb, rather than inhibition of its intrinsic GAP activity."

sparser
"Thus, one possible explanation is that sleep deprivation reduces cAMP, which would elicit more PDE4D interaction with Rheb and thereby decrease the interaction between Rheb and mTOR ( xref )."

reach
"We therefore wondered if peripheral redistribution of lysosomes in acid limits the ability of RHEB to activate lysosome bound mTOR, explaining why TSC2 deletion could enhance, but not fully rescue, mT[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"As shown in xref , many red dots were observed in control OSCC cells, suggesting the vibrant interaction between mTOR and RHEB and activation of the mTOR pathway."

sparser
"Rheb-mTOR is the key signaling pathway component that regulates cognitive impairment caused by autophagy dysfunction in the hippocampus during brain aging."

reach
"Detailed analysis indicated that FMDV infection promoted the transfer of mTOR to lysosome to enhance interaction between mTOR and Rheb, and activated PI3K/AKT/TSC2/Rheb/mTOR/p70S6K1 pathway to promote viral replication."

reach
"RHEB binds mTOR, activating mTORC1 kinase activity."

reach
"FMDV infection promoted the transfer of mTOR to the lysosome and enhanced interaction between mTOR and Rheb."

reach
"The GTP bound form of Rheb directly binds the catalytic domain of mTOR and, therefore, activates mTORC1 [XREF_BIBR, XREF_BIBR]."

sparser
"We found that activated microglia accelerate heat stress-induced neuronal autophagy through transmission of exosomal miR-155 to neurons, with the subsequent Rheb-mTOR signaling pathway suppressed and neuronal autophagy enhanced."

reach
"Further studies showed an interaction between mTOR and Rheb in the mock-infected cells (Pearson’s co-localization coefficient: 0.95), and FMDV infection enhanced the interaction between mTOR and Rheb (Pearson’s co-localization coefficient: 0.98) (Fig 9B)."

sparser
"Activated Rheb binds and activates the mechanistic target of rapamycin (mTOR), specifically, when it is interacting with Raptor and PRAS40 as part of a protein complex referred to as mTOR complex 1 (mTORC1)."

sparser
"From the PLA assay, the reduced interaction of mTOR and RHEB was observed more intuitively."

reach
"We further investigate the interaction between mTOR and Rheb in the context of viral infection using a coimmunoprecipitation assay."

reach
"The results showed that the interaction between mTOR and Rheb was enhanced in the FMDV-infected cells compared to that in the mock-infected cells (Fig 9C)."

sparser
"Cryo-electron microscopy has revealed that Rheb-GTP binds allosterically to mTOR, resulting in a global conformational change that allosterically realigns the active-site residues to enable substrate phosphorylation xref ."

reach
"Additionally, FMDV-induced interaction between mTOR and Rheb was reduced in the SLC38A8 cells compared to the WT cells (Fig 9D)."

reach
"Altogether, these results indicated that FMDV-induced upregulation of amino acids promoted the transfer of mTOR to the lysosome and increased interaction between mTOR and Rheb."

reach
"Meanwhile, in the FMDV-infected cells, the phosphorylation of p70S6K1 and rpS6 was also decreased in the SLC38A8 cells compared to that in the WT cells (Fig 10C), suggesting that FMDV-induced the upregulation of aspartate was responsible for the increase of the phosphorylation of p70S6K1 and rpS6.As described above, amino acid stimulation enhanced interaction between Rheb and mTOR, resulting in the activation of the mTOR/p70S6K1 signaling axis."

reach
"Meanwhile, upstream of mTORC1 and mTORC2, Ras homolog enriched in brain (Rheb), a GTPase, also directly binds mTOR but results in the activation of mTORC1 [24]."

sparser
"It is believed that Rheb physically binds with mTOR, leading to its activation [ xref ]."

reach
"On the other hand, the upregulation of amino acids enhanced the interaction between mTOR and Rheb to activate the mTOR/p70S6K1 signaling axis to promote viral replication."

sparser
"Accordingly, we considered the possibility that STAT6 interferes with Rheb-mTOR interactions, thereby suppressing mTOR activity."

sparser
"To further examine the effects of STAT6 on Rheb-mTOR binding, we analyzed the interactions of STAT6 with Rheb or mTOR."

sparser
"This allows mTOR to bind to its activator RHEB."

sparser
"In response to amino acids, Raptor interacts with the lysosomal Ras-related guanosine 5’-triphosphate (GTP)-binding protein (Rag small GTPase protein complex tethering mTORC1 to the lysosomal membrane, where it encounters another small GTPase, Ras homolog enriched in brain (Rheb) that directly interacts with the mTOR kinase and stimulates the activity of mTORC1 [ xref , xref ]."

sparser
"In the presence of nutrients and growth factors, mTOR interacts with Rheb-GTP on the surface of lysosomes, initiating the phosphorylation of proteins [ xref ]."

sparser
"Consequently, we demonstrated that RHEB binds to the whole mTOR both in the presence or absence of GTPγS, with five-fold weaker affinity in the presence of GTPγS. In addition, RHEB bound to the truncated mTOR fragments of N-heat domain (∆N, aa 60-167) or M-heat domain (∆M, aa 967-1023) with the same affinity in the absence of GTP."

sparser
"Several studies have demonstrated the role of RhebmTOR signaling in axon elongation and synaptic morphogenesis [ xref , xref ]."

sparser
"RhebmTOR signaling is considered an important mechanism for the maintenance of LTP and learning and memory functions [ xref , xref ]."

sparser
"The site of Rheb-MTOR interaction identified by cryo-EM, accompanied by extensive biochemical verification of Rheb-GTP activation of MTORC1, invalidates the earlier conclusion that Rheb binds to the MTOR catalytic domain, which was based on the use of Rheb co-expression with MTOR fragments xref ."

sparser
"As described previously, RhebmTOR signaling is a key regulator of various mechanisms associated with neuronal survival and regeneration."

sparser
"Following inhibition of TSC complex, GTP-bound Rheb binds to the catalytic domain of mTOR to activate mTORC1 ( xref )."

sparser
"Rheb binding to MTOR in vitro exhibits positive cooperativity, and the unique configuration of MTORC1 bound to Rheb as seen in cryo-EM enabled the conclusion that MTORC1 dimers always show either two apo-MTORC1s or two Rheb-MTORC1s."

sparser
"The Rag-Ragulator complex aids in mTOR signal transduction, at least in part, by tethering the mTOR/Raptor complex to the lysosome, where Rheb-GTP can bind and activate mTOR."

sparser
"Although incubation with leucine alone did not increase the association between these two proteins, it prevented the ethanol-induced decrease in mTOR-Rheb association [ xref ]."

sparser
"The half-maximal concentration for Rheb activation of MTORC1 in vitro is about 100 μM, which is consistent with the very low affinity of the Rheb-MTOR interaction identified in early studies."

sparser
"The decrease in mTOR-Rheb binding in the presence of ethanol was associated with an ethanol-induced increase in phosphorylation of AMPK T172 and TSC2 S1387 , and these ethanol-induced changes were prevented by co-incubation with leucine [ xref ]."

sparser
"This finding was interpreted to indicate that MTOR interaction with Rheb-GTP was required to configure MTOR to an active state, a conclusion consistent with genetic data from Drosophila xref , xref and with the recent results of Yang et al . xref ."

sparser
"Activated Rheb protein binds to mTOR complex 1 (mTORC1), promoting the activation of the mTORC1 signaling pathway [ xref ]."

reach
"Rheb binds directly to mTOR, an interaction that appears to be essential for mTORC1 activation."

sparser
"For example, the TSC1/TSC2 complex inhibits mTOR functions by inactivating Rheb; Rheb binds and activates mTOR only when the former is in its GTP-bound form. xref The phosphoinositide-3-kinase (PI3K)-Akt pathway activates mTOR by Akt-mediated phosphorylation of TSC2, followed by disassembly and inhibition of the TSC1/TSC2 complex. xref , xref The extracellular signal-related kinase (ERK) stimulates mTOR activation by either directly phosphorylating TSC2 or by inducing TSC2 phosphorylation via its two downstream effectors, ribosomal S6 kinase (RSK) and DAPK. xref – xref Finally, AMP-activated protein kinase (AMPK) phosphorylates TSC2 and stabilizes the TSC1/TSC2 complex, thereby inhibiting Rheb-mediated activation of mTOR. xref AMPK can also regulate mTOR function in a TSC-independent manner by directly phosphorylating Raptor, an essential component of mTORC1, to inhibit mTOR activity. xref Notably, TSC1 and 2, PI3K, and Akt are all subject to mutation in human tumors."

reach
"The contribution of Rheb 's regulation of PL-D1 and FKBP38 to mTORC1 activation, relative to Rheb 's direct binding to mTOR, remains to be fully defined."

sparser
"It can be argued that the very weak affinity of Rheb for MTOR results in a loss of the Rheb-MTOR interaction upon cell extraction; however, this returns to the question of how the Rheb activation mechanism demonstrated by Yang et al . xref relates to the mechanism operative in the cell and whether a continuing association of Rheb with MTOR is required for the maintenance of MTORC1 activity in the cell."

sparser
"MTOR can bind Rheb regardless of its GDP/GTP content, even though only GTP-Rheb can activate mTORC1 activity [ xref , xref ]."

sparser
"However, elevated cAMP levels disrupt the interaction of PDE4D with Rheb and increase the interaction between Rheb and mTOR."

sparser
"Although it is possible that ethanol reduced mTOR-Rheb binding via AMPK/TSC2/Rheb signaling, it is also possible that ethanol suppressed the ability of mTOR to interact with Rheb due, at least in part, to preventing mTOR from binding the Rag complex at the lysosome."

sparser
"Fluorescence lifetime imaging of overexpressed Rheb, MTOR, and Raptor, each fused to fluorescent proteins, did detect the presence of a Rheb-MTOR complex in both the cytoplasm and the nucleus (the latter devoid of Raptor) xref ; as yet, however, the presence of endogenous Rheb in the nucleus and its localization therein await confirmation."

sparser
"This enhanced Rheb-mTOR interaction induces the activation of mTORC1 and cap-dependent translation, a cellular function of mTORC1."

sparser
"Amino acids, especially BCAA, are believed to activate mTOR and to increase in association with Rheb-mTOR ( Sancak et al., 2010 )."

sparser
"As is known, Rheb interacts with mTOR directly and activates it in the GTP‐bound form. xref According to the association between HOTAIRM1 and the mTOR pathway, we tried to explore whether HOTAIRM1 promotes OS progression by modulating mTOR/c‐MYC‐mediated glycolysis in a Rheb‐dependent manner."

sparser
"RHEB binds to mTOR distally from the kinase active site, yet causes a global conformational change that allosterically realigns active-site residues, accelerating catalysis."

reach
"TSC1 and TSC2 are part of a heterotrimeric complex that functions as a GTPase activating protein for Rheb, which in turn binds and activates mTOR complex 1 (mTORC1)."

sparser
"Song and coworkers have also confirmed that resistance exercise in the human skeletal muscle resulted in the rapid translocation of mTOR toward the lysosome, with the concurrent dissociation of TSC2 from Rheb, which would facilitate the interaction of mTOR and GTP‐Rheb (Song et al.,  xref )."

reach
"TSC2 is a GTPase-activating protein for the small G-protein Rheb, which, in its GTP-form binds to and activates mTOR."

sparser
"This suggests a mode of interaction analogous to the direct association of mTOR with the small GTPase Rheb, in which direct binding of active Rheb to mTOR, potentially through the mTOR kinase domain, allosterically promotes mTORC1 kinase function [ xref , xref ]."

sparser
"Because protein-protein interactions are important in mediating signaling events, the association of Rheb with mTOR was assessed."

reach
"(B) Co-immunoprecipitation experiments showing the interaction between Rheb and mTOR or GAPDH, and between mTOR and Tel2 or Rag C. Protein extracts of liver samples from three controls and three fruct[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"The addition of PA increased the binding of Rheb with mTOR, while EtOH exposure decreased this interaction ( xref )."

sparser
"There, mTOR interacts with Rheb, resulting in the activation of the mTOR kinase."

sparser
"Because TSC2 functions as a GAP for Rheb, inhibition of its activity promotes the binding of Rheb with mTOR, thereby enhancing mTORC1 function."

sparser
"DQ661 in particular has improved lysosomal targeting capability and in addition to inhibiting autophagy, impairs the activity of palmitoyl-protein thioesterase (PPT1) that is required for mTOR interaction with Rheb at the lysosome ( xref )."

reach
"However, elevated cAMP levels disrupt the interaction of PDE4D with Rheb and increase the interaction between Rheb and mTOR."

reach
"How the binding of Rheb to mTOR activates the mTOR kinase remains controversial."

sparser
"GDP inhibits mTOR via weakening the binding of GTP-dependent Rheb to mTOR molecules [ xref ]."

reach
"Inactivation of TSC1 or TSC2 leads to an increase in GTP-bound Rheb, a modulator of mechanistic target of rapamycin (mTOR) activity."

sparser
"The elevation of cellular cAMP levels disrupts phosphodiesterase-Rheb interaction, increases Rheb-mTOR interaction, and consequently leads to mTOR1 activation."

sparser
"MTOR interaction with Rheb is restricted in basal conditions due to the enzymatic activity of the tuberous sclerosis complex proteins TSC1 and TSC2, which maintain Rheb in a GDP-bound state xref ."

sparser
"Previous studies have demonstrated that mTOR interaction with GTP bound Rheb is required to elicit complete mTORC1 activation xref ."

sparser
"In particular, the Rheb-mTOR axis is a pivotal pathway that regulates diverse cellular physiologies such as cell growth, cell cycle progression, axon regeneration, autophagy, nutritional deprivation, [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"The GTP-loaded Rheb interacts with mTOR catalytic domain and robustly stimulates mTORC1 activity [ xref ]."

sparser
"Altogether, these results indicated that FMDV-induced upregulation of amino acids promoted the transfer of mTOR to the lysosome and increased interaction between mTOR and Rheb."

sparser
"This is indeed the case as overexpression of GAP-domain-dead TSC2 mutants ( xref ) efficiently suppressed activity of endogenous or overexpressed Rheb, and perturbed Rheb-mTOR binding, albeit to a lesser extent when compared to the wild-type construct ( xref and xref )."

sparser
"The interaction between PDE4D and Rheb is abolished by cytosolic cAMP and, as a result, Rheb-mTOR interaction is enhanced by releasing PDE4D from Rheb [ 24 ]."

reach
"The activity of mTORC1 in mammalian cells is critically dependent on the small GTPase Rheb, which binds directly to the mTOR catalytic domain and, in its GTP bound state, promotes mTORC1 catalytic activity."

reach
"Using genetic mouse models, we show that Rheb, a small GTPase that binds mTOR, is critical for circadian oscillation of mTORC1 activity in liver."

reach
"RHEB binds to mTOR on the lysosome surface and inactivates downstream signaling ."

sparser
"When PDE4D binds Rheb it inhibits the ability of Rheb to activate mTORC1 and hence it is plausible that PDE4DIP (also called myomegalin) impacts on this relationship, as an increased interaction between Rheb and mTOR should promote growth."

sparser
"Interestingly, the N41A mutation impaired the interaction of mTOR with Rheb, suggesting that mTOR (or a component within the mTOR complex) binds to Asn41 within the switch I region of Rheb."

reach
"Interestingly, the N41A mutation impaired the interaction of mTOR with Rheb, suggesting that mTOR (or a component within the mTOR complex) binds to Asn41 within the switch I region of Rheb."

sparser
"The mitochondrial protein FKBP38 (which belongs to the same immunophilin family as FKBP12) has been shown to interact with mTOR and Rheb (Ras homolog enriched in the brain) in a nutrient-dependent manner."

reach
"Co-immunoprecipitation revealed Rheb as the dhBBR target that bound to mTOR."

sparser
"Under the induction of amino acids and insulin, the formation of the RHEB-mTOR complex in the skeletal muscle of newborn piglets may be related to the activation of mTORC1 that regulates skeletal muscle protein synthesis [ xref ]."

sparser
"FMDV infection promoted the transfer of mTOR to the lysosome and enhanced interaction between mTOR and Rheb."

reach
"We reveal that mTOR binds to Rheb and RhebL1 and this association requires N41 within the putative switch I region."

sparser
"During preparation of this manuscript, work carried out by Long et al. [12,13] also showed that Rheb interacted with mTOR."

sparser
"These findings collectively show that mTOR does not function as a typical small G-protein downstream effector, i.e., mTOR is bound to Rheb regardless of its guanine nucleotide bound status."

sparser
"Whereas the regulators of GTP-loading onto Rheb remain to be established, the interaction between Rheb and mTOR is sufficient both in vitro and in vivo to promote the phosphorylation of mTORC1 substrates [ xref , xref ]."

reach
"Long X, Ortiz-Vega S, Lin Y, Avruch J. Rheb binding to mammalian target of rapamycin (mTOR) is regulated by amino acid sufficiency."

sparser
"To check the effect of Rheb-HSP70 interaction on Rheb interaction with mTOR, we expressed Rheb with HSP70 in HEK293 cells."

sparser
"TSC1/2 possesses Rheb–GAP activity that directly antagonizes RhebmTOR signaling [ xref ]."

sparser
"As shown in Fig. 2 A, Rheb interacted with mTOR and this Rheb-mTOR interaction was almost completely abolished by HSP70 ( Fig. 2 A)."

sparser
"Intriguingly, the interaction of Rheb with mTOR is relatively weak and in a nucleotide-independent manner, which is different from the interactions of classical Ras proteins with their effectors ( xref )."

reach
"TSC1/TSC2 acts as a GTPase-activating protein complex for the small G-protein Rheb, which, in its GTP-form binds to and activates mTOR ( Proud, 2006; Inoki et al., 2005 )."

sparser
"How the binding of Rheb to mTOR activates the mTOR kinase remains controversial."

sparser
"Complex of mTOR and Rheb promote the phosphorylation of downstream molecules, consequently promoting protein synthesis."

sparser
"Suggested leucine mediated mechanisms include promotion of Rheb binding to mTOR, stabilization of the activated mTOR-raptor complex, or mTOR/raptor activation through an alternative class 3 PI3K (hVps34) parallel to that controlled through insulin ( xref ; xref ; xref )."

sparser
"Activation requires interaction between mTOR and the G protein Rheb in its GTP-activated conformation."

reach
"Rheb interacts directly with mTOR and activates mTOR kinase, whereas Rabin8 interacts with Rheb and inhibits the phosphorylation of Ser235 and Ser236 in small ribosomal subunit protein S6."

reach
"Activated (GTP bound) Rheb directly interacts with and activates mTOR (mammalian target of rapamycin)."

sparser
"Studies have shown that FMDV-infected PK15 cells promote the transfer of mTOR to lysosomes to enhance the interaction between mTOR and Rheb and activate the PI3K/AKT/TSC2/Rheb/mTOR/p70S6K1 pathway to promote viral replication [ xref ], which is consistent with the results of RNA‒seq in this study."

sparser
"GTP-bound RHEB directly binds and activates mTOR in the lysosome in response to nutrient levels, resulting in metabolic reprogramming and cell growth xref ."

sparser
"We further investigate the interaction between mTOR and Rheb in the context of viral infection using a coimmunoprecipitation assay."

sparser
"Structural data have shown that Rheb binding to mTOR causes a dramatic conformational change that allows the alignment of residues in the active site, thus enhancing the phosphorylation of S6K and 4E-[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Active Rheb binds mTOR promoting its activation."

sparser
"This is a surprising notion given current favored models of mTORC1 signaling ( xref ), but not unprecedented ( xref ; xref ), that would indicate trans-endomembrane RHEB-mTOR interaction mechanistically underlies mTOR activation ( xref )."

sparser
"Rheb directly binds to the mTOR complex, and the active GTP-bound form of Rheb is required for mTOR activation ( xref ; xref )."

reach
"It is believed that Rheb physically binds with mTOR, leading to its activation [XREF_BIBR]."

reach
"At the same time, by regulating LAT1, enhanced amino acid uptake moves mTOR to the lysosome where it can be activated by GTP bound Rheb."

sparser
"Detailed analysis indicated that FMDV infection promoted the transfer of mTOR to lysosome to enhance interaction between mTOR and Rheb, and activated PI3K/AKT/TSC2/Rheb/mTOR/p70S6K1 pathway to promote viral replication."

sparser
"Association of Rheb to the mTOR Complex 1 appears to require AA, and GTP loading of Rheb has been shown to be blocked by AA depletion in vitro ( xref ; xref )."

sparser
"Follow-up investigations from the same group have established whole-egg consumption results in greater co-localisation of mTOR-Rheb complexes to the lysosome following RET than that of egg whites [ xref ]."

sparser
"The results showed that the interaction between mTOR and Rheb was enhanced in the FMDV-infected cells compared to that in the mock-infected cells ( xref )."

sparser
"Additionally, FMDV-induced interaction between mTOR and Rheb was reduced in the SLC38A8 -/- cells compared to the WT cells ( xref )."

sparser
"In their GTP-bound configurations, Rag GTPases avidly bind raptor, and thus attract the mTORC1 complex to endosomal membranes, facilitating the interaction of mTOR and GTP-bound Rheb."

sparser
"The kinase activity of mTORC1 is stimulated by direct interactions of mTOR with the GTP-bound form of the Rheb GTPase [ xref ], a protein that resides on the cytoplasmic surface of lysosomes [ xref ]."

sparser
"Since it remained possible that FTS may disrupt Rheb interaction with mTOR, we next asked if FTS inhibits Rheb-induced activation of mTOR."

sparser
"Under low-glucose conditions, GAPDH inhibits mTOR by interacting with the mTOR activator Rheb to prevent the binding of Rheb to mTOR in HEK293 cells or mouse embryonic fibroblasts."

reach
"Although direct interaction between Rheb and mTOR has not been ruled out, it appears more likely that Rheb-GTP recruits additional protein (s) to the mTOR complex to modulate its activity.The mTOR pathway plays a central role in nutrient and energy sensing to regulate cell growth and proliferation (reviewed in Ref."

reach
"Consequently, we demonstrated that RHEB binds to the whole mTOR both in the presence or absence of GTPγS, with five-fold weaker affinity in the presence of GTPγS."

reach
"Effects of amino acid starvation on the interaction of Rheb and mTOR were reported [XREF_BIBR]."

sparser
"As described above, amino acid stimulation enhanced interaction between Rheb and mTOR, resulting in the activation of the mTOR/p70S6K1 signaling axis."

reach
"This may agree with the previous observations that the interaction between Rheb and mTOR is rather weak."

reach
"While CHAPS can retain the interaction of Rheb and mTOR, NP40 disrupts the interaction."

sparser
"Altogether, these results indicated that FMDV induced the activation of the PI3K/AKT/TSC2/Rheb pathway, and the upregulation of amino acids enhanced the interaction of mTOR-Rheb to activate the mTOR/p70S6K1 signaling axis to promote protein translation for viral replication."

reach
"While the molecular mechanism is not well understood, these phosphorylation events relieve the inhibitory effects of the TSC1 and TSC2 complex on Rheb and mTOR complex 1 (mTORC1), thereby activating mTORC1 in response to growth factors."

reach
"Rheb directly interacts with mammalian target of rapamycin to increase its activity."

reach
"Structural data have shown that Rheb binding to mTOR causes a dramatic conformational change that allows the alignment of residues in the active site, thus enhancing the phosphorylation of S6K and 4E-[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"SMG9 is oriented to face SMG1’s kinase active site in a manner similar to that seen in mTOR binding by its regulatory factor RHEB [ 53 ]."

sparser
"Proximity ligation assays (PLAs) in WT PPT1 and KO PPT1 cells established a significant loss of physical interaction between the LAMTOR1/p18 subunit of Ragulator and the ATP6V1A subunit of the vacuolar-type H+-ATPase (v-ATPase), which is critical for and results in the loss of mTOR-Rheb physical interactions ( xref ) ( xref , xref )."

sparser
"Our data indicated that FMDV-induced upregulation of amino acids promoted the transfer of mTOR to the lysosome and increased interaction between mTOR and Rheb, resulting in the activation of Rheb/mTOR/p70S6K1/rps6 signaling axis to promote viral replication."

sparser
"On the other hand, the upregulation of amino acids enhanced the interaction between mTOR and Rheb to activate the mTOR/p70S6K1 signaling axis to promote viral replication."

sparser
"In its GTP-bound state, rheb binds to mTOR near its kinase domain and directly stimulates its activity [ xref ]."

reach
"Once activated, Rheb binds and sequesters the endogenous inhibitor of mTOR, FKBP-38, resulting in mTORC1 activation (Bai et al., 2007)."

sparser
"Rheb directly binds to mTOR and activates it in a GTP-dependent manner ( xref )."

reach
"Indeed, over-activation of AKT leads to a disinhibition of Rheb which in turn binds and sequesters the endogenous inhibitor of mTOR, FKBP-38 (Bai et al., 2007)."

sparser
"Another recent study has indicated that amino acids promote the association of Rheb with mTOR, an effect that is probably not caused by increased charging of Rheb with GTP [( Roccio et al., 2006 ); co[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Although it is tempting to conclude that mTOR itself is the amino acid sensor, it is still possible that the effect of amino acids is indirect and that they decrease the concentration of an inhibitor [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Current models suggest that RHEB and the mTOR complex interact on the lysosomes for subsequent mTOR activation."