From: Subject: Mechanism of thalidomide to enhance cytotoxicity of temozolomide in U251-MG glioma cells in vitro Date: Sun, 2 Aug 2009 09:17:34 +0200 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0000_01CA1352.1226A910" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.5579 This is a multi-part message in MIME format. ------=_NextPart_000_0000_01CA1352.1226A910 Content-Type: text/html; charset="Windows-1252" Content-Transfer-Encoding: quoted-printable Content-Location: http://www.cmj.org/Periodical/PaperList.asp?id=LW200964465955202403 Mechanism of thalidomide to enhance cytotoxicity of = temozolomide in U251-MG glioma cells in vitro
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GAO Song Department of Neurosurgery, = Tianjin=20 Medical University General Hospital, Tianjin 300052, China; = YANG Xue-jun Department of = Neurosurgery,=20 Tianjin Medical University General Hospital, Tianjin 300052, = China;=20 ZHANG Wen-gao Department of = Neurosurgery,=20 Tianjin Medical University General Hospital, Tianjin 300052, = China;=20 JI Yan-wei Department of = Neurosurgery,=20 Tianjin Medical University General Hospital, Tianjin 300052, = China;=20 PAN Qiang Department of = Neurosurgery,=20 Tianjin Medical University General Hospital, Tianjin 300052, = China=20

Correspondence to: YANG Xue-jun =  Department of=20 Neurosurgery, Tianjin Medical University General Hospital, = Tianjin=20 300052, China  (Tel:86-22-60361043 Fax:86-22- 60362621=20 Email:ydenny@yahoo.com )This study was supported by : National = Natural=20 Science Foundation of China(No. 30571904 and 30772228) = Keywords: gl= ioma=A1=A4<= EM>thalidomide=A1=A4= temozolomide=A1=A4autophagyAbstract:

Background Glioma is the most = common primary=20 brain tumor with poor prognosis. Temozolomide has been used = with=20 thalidomide to treat gliomas. We investigated the = synergistic=20 mechanism of these two drugs in vitro.

Methods Human malignant glioma = cells U251-MG=20 were cultured and assigned to four groups with different = treatments=20 for 3 days: temozolomide group (100 =A6=CCmol/L), = thalidomide group (100=20 =A6=CCg/L), temozolomide (100 =A6=CCmol/L) plus thalidomide = group (100=20 =A6=CCg/L) and control group. MTT assay was applied to = evaluate the cell=20 viability. Cell cycle was analyzed by flow cytometry. The=20 ultra-structural features of autophagosomes were observed = with=20 electron microscope. Acridine orange and = monodansylcadaverine were=20 adopted to label autophagosomes and flow cytometry was = applied for=20 quantification of autophagosomes. The expression of=20 autophagy-associated protein was detected by Western = blotting.

Results Proliferation of tumor = cell was=20 obviously suppressed by temozolomide with thalidomide = treatment than=20 by either drug used alone (P=3D0.000 for each day). = The=20 combination treatment induced cell cycle arrest at G0/G1 = phase.=20 Typical autophagic ultra-structural character was found = after the=20 combined treatment. Thalidomide promoted the autophagy = induced by=20 temozolomide. The autophagy-associated proteins =A3­ = microtubule=20 associated protein 1 light chain 3 (MAP1LC3) and Beclin1 = were more=20 significantly up-regulated by the combined treatment than=20 temozolomide used alone (MAP1LC3, P=3D0.000; Beclin1, = P=3D0.004). The expression level of phosphatase and = tensin=20 homolog deleted on chromosome ten (PTEN), which promoted = autophagy=20 by suppressing PI3K/Akt/mTOR signaling pathway, was elevated = by=20 thalidomide (thalidomide group: P=3D0.000; combined = group:=20 P=3D0.002).

Conclusions Thalidomide enhances = the=20 cytotoxicity of temozolomide by promoting the autophagy = induced by=20 temozolomide. Contributing to the up-regulation of PTEN by=20 thalidomide, the expression of autophagy associated = protein-MAP1LC3=20 and Beclin1 was enhanced, which leads to a reinforced = autophagy in=20 the combined treatment of temozolomide and thalidomide in = vitro.


 2009;122(11):1260-1266
Chinese=20 Medical Journal, 2009, Vol. 122 No. 11 : = 1260-1266
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Mechanism of thalidomide to enhance=20 cytotoxicity of temozolomide in U251-MG glioma cells in=20 vitro
GAO Song, YANG Xue-jun, ZHANG = Wen-gao, JI=20 Yan-wei , PAN Qiang
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Glioma is the most common primary brain = tumor=20 associated with a high degree of morbidity and=20 mortality.1 As the molecular mechanisms of = gliomas have=20 been clarified gradually, improving chemotherapeutic effect = turns=20 out to be the key part in combination treatment.2 = The=20 therapeutic goal of chemotherapy triggers tumor-selective = cell=20 death.3 Compared with necrosis, programmed cell = death=20 (PCD) is more feasible to be induced by chemoagents. Until = now, two=20 types of PCD have been known: apoptosis and autophagic cell = death.=20 Glioma cells are more likely to respond to therapy through = autophagy=20 than through apoptosis.4

Temozolomide (TMZ), is a novel oral=20 chemotherapeutic drug with broad-spectrum antitumor activity = which=20 penetrates into the brain and does not require hepatic = metabolism=20 for activation. Kanzawa et al5 reported TMZ = inhibited the=20 viability of malignant glioma cells by induction of = autophagic cell=20 death. However, Using TMZ alone would induce drug resistance = in=20 tumor and decrease the ratio of efficacy to price. = Fortunately, the=20 favorable safety profile of temozolomide allows it to be=20 co-administered with various agents. Thalidomide (THD) has=20 anti-angiogenic properties.6 A phase II study=20 demonstrated that the combination of TMZ and THD was more = efficient=20 than either agent alone.7 The two drugs act in = different=20 ways to exert the treatment effects: TMZ is related to = cytotoxicity=20 while THD inhibits tumor growth by anti-angiogenesis. It is = poorly=20 documented whether a synergistic mechanism exists or = not.

In our preliminary related study in = vitro,=20 TMZ combined with THD more significantly inhibited tumor = cell growth=20 compared with TMZ used alone. The result implies us that = besides=20 anti-angiogenesis in the treatment of glioma, the = synergistic effect=20 of THD and TMZ may exist.

METHODS

Tumor cell line
Human GBM cell = line=20 U251-MG was obtained from the Institute of Neurology of = Tianjin=20 Medical University. U251-MG is a human brain tumor cell line = that=20 was established from a patient with glioblastoma multiforme = by=20 Ponten, with the characteristics such as low terminal cell = density,=20 strong density dependent inhibition of proliferation, = diploidy and=20 contact inhibition of membrane mobility.8 The = cells were=20 cultured as adherent cells in DMEM (GIBCO BRL Grand Island, = NY, USA)=20 supplemented with 10% fetal bovine serum (GIBCO BRL), 4 = mmol/L=20 glutamine, 100 U/ml penicillin and 100 mg/ml = streptomycin.

Reagents
TMZ was kindly = supplied by Tasly=20 Pharmaceutical Co., Ltd. (Tianjin, China). THD was purchased = from=20 Changzhou Pharmaceutical=20 Factory (Jiangsu, China). TMZ and THD were dissolved in = 10%=20 dimethylsulfoxide (DMSO) (Sigma Chemical Co., St. Louis, MO, = USA) to=20 make a stock solution (TMZ: 100 mg/ml; THD: 100 mg/ml). DMSO = concentration was kept below 0.1% in all the cell cultures = and did=20 not exert any detectable effect on cell growth or cell = death.=20 Acridine orange, 3-(4,5-=20 dimethylthiazol-2-yl)-2,5-diphenyltetrazolium bromide (MTT), = and=20 monodansylcadaverine (MDC) were purchased from Sigma = Chemical Co=20 (St. Louis, MO, USA). Goat anti-human polyclonal antibodies = against=20 Beclin1, MAP1LC3, PTEN and =A6=C2-actin; horseradish=20 peroxidase=A8Cconjugated secondary antibody (rabbit anti = goat) were=20 purchased from Santa Cruz Biotechnology (Santa Cruz, CA, = USA).

Treatments
The IC50 = (concentration=20 resulting in cell viability of 50% of control group) of TMZ = for=20 U251-MG was less than 200 mmol/L. So we fixed the = concentration of=20 TMZ at a clinically achievable dose at 100 = =A6=CCmol/L.9=20 Cultures of U251-MG were assigned to four groups based on = different=20 treatments: TMZ (treated with TMZ 100 =A6=CCmol/L), THD = (treated with=20 THD 100 =A6=CCg/L), TMZ plus THD (treated with TMZ 100 = =A6=CCmol/L plus THD=20 100 =A6=CCg/L) and solvent control (DMSO) group.

Cell viability assay
The = cytotoxicity of=20 TMZ and THD on U251-MG was determined by MTT assay. Tumor = cells were=20 seeded at 2=A1=C1103 cells/well (0.1 ml) in = 96-well flat=20 bottom plates and incubated overnight at 37.1=BAC. After = exposure to=20 the above described treatment for 24, 48, 72, and 96 hours. = MTT (0.5=20 mg) was added to each well every 4 hours before the ending = of=20 incubation, when the MTT reaction was stopped by addition of = 10%=20 SDS-0.1 mol/L HCL. The formazan crystals of the cell were = dissolved=20 in DMSO and then its absorbance at 490 nm was measured by a=20 microplate reader (Bio-Tek ELX800uv, Bio=A8CTek Instrument = Inc.=20 Winooski. VT., USA). All experiments were repeated in = triplicate.=20

Cell cycle analysis
For cell = cycle=20 analysis, after different treatments for 3 days, U251-MG = cells were=20 trypsinized and then fixed in chilled 70% ethanol for at = least 24=20 hours before being stained with 50 =A6=CCg/ml propidium = iodide and 100=20 =A6=CCg/ml RNAse A (Sigma, USA). After being stained for = thirty minutes,=20 the samples were analyzed using flow cytometry (Becton = Dickinson,=20 Franklin Lakes, NJ, USA). The data were analyzed by Modifit = system=20 (Becton Dickinson Company, USA). All experiments were = repeated in=20 triplicate.

Ultra-structural features of treated = cells with=20 electron microscopic observation
To morphologically=20 demonstrate the induction of autophagy in treated tumor = cells, we=20 performed the ultra-structural observation as described=20 previously.10 After different treatments for 3 = days,=20 U251-MG were harvested by trypsinization, washed twice with = PBS and=20 fixed with ice-cold glutaraldehyde (3% in 0.1 mmol/L = cacodylate=20 buffer, pH 7.4) for 30 minutes. After being washed in = PBS=A3=ACthe cells=20 were fixed in OsO4 and embedded in Epon; 0.1 mm thin = sections were=20 stained with uranyl acetate/lead citrate (Fluka) and viewed = in a=20 Philips EM 400 electron microscope.

Detection and quantification of = acidic=20 vesicular organelles (AVOs) stained with acridine=20 orange
Autophagy is characterized by formation and = promotion=20 of AVOs. In acridine orange-stained cells, the cytoplasm and = nucleolus fluoresce bright green, whereas acidic = compartments=20 fluoresce bright red. The volume of the cellular acidic = compartment,=20 which reflects the number of autophagic vacuoles, can be = quantified.=20 After different treatments for 3 days, cells were stained = with=20 acridine orange at a final concentration of 1 =A6=CCg/ml for = 15 minutes,=20 removed from the plate and collected in phenol red-free = growth=20 medium. Green (510=A8C530 nm) and red (>650 nm) = fluorescence=20 emission from 104 cells illuminated with blue = (488 nm)=20 excitation light were measured with flow cytometry and the = data were=20 analyzed with Cell Quest software (Becton Dickinson = company).

Visualization of MDC-labeled = autophagic=20 vacuoles
A fluorescent compound, MDC, has been = proposed as a=20 special tracer for autophagic vacuoles.11 The = U251-MG=20 cells were treated as previously described, respectively. = The=20 autophagic vacuoles were labeled with MDC by incubating cell = growth=20 on cover-slips with 0.05 mmol/L MDC in PBS at 37=A1=E3C for = 1 hour.=20 After incubation, cells were washed twice with PBS and = immediately=20 analyzed by fluorescence microscopy using an inverted = microscope=20 (Olympus IX-71, Japan) equipped with a filter system. Images = were=20 obtained with a CCD camera (Olympus DP30BW, Japan) and = processed=20 using the program Meta View, version 4.5 (Universal Images=20 corporation, USA).

Quantification of MDC labeled = autophagic=20 vacuoles
The cells were stained with the method = mentioned=20 above. Then the samples were analyzed by flow cytometry to = detect=20 the ratio of MDC staining positive cells. Excitation wave = length was=20 530 nm, emission filter was 590 nm. All experiments were = repeated in=20 triplicate.

Western blotting
For Western = blotting,=20 protein was abstracted from the treated cells. After the = protein=20 concentration was determined, equivalent amounts of proteins = was=20 boiled in sample buffers with DL-Dithiothreitol (DTT) and = then=20 concentrifuged the protein at 12000 r/min for 15 minutes at = 4=A1=E3C.=20 The protein was separated by SDS-PAGE (PAGE: 15% gel for = MAP1LC3;=20 10% gel for Beclin1). Separated proteins in gel were = transferred=20 onto PVDF membranes by an electroblot apparatus (Bio-Rad, = USA).=20 Filters were blocked for 2 hours in 5% low fat milk and were = incubated with goat anti-human antibody for = MAP1LC3/Beclin1/PTEN and=20 =A6=C2-actin at 4=A1=E3C for 12 hours. Membranes were then = washed with PBS-T=20 and incubated with HRP-conjugated anti-goat antibody for 1 = hour.=20 Specific signals were detected from quantitative gel and=20 Western blotting imaging system (Becton Dickinson, = Franklin=20 Lakes, NJ, USA).

Statistical = analysis
Statistical analyses=20 were performed using SPSS version 13.0 software (SPSS Inc., = Chicago,=20 IL, USA). Multiple groups were compared using analysis of = variance=20 (ANOVA) followed by post LSD testing where appropriate. Data = were=20 considered statistically significant at P <0.05. = All data=20 are expressed as mean =A1=C0 standard deviation (SD).

RESULTS

Combination of TMZ with THD = exhibited an=20 obvious inhibition on cell viability
As shown in = Figure 1:=20 after different treatments for 3 days, the inhibition of = cell=20 proliferation in TMZ with THD group was more remarkable than = that in=20 TMZ group (P <0.05). Compared with the control = group=20 (P=3D0.63), THD used alone showed no significant = inhibition on=20 cell viability of U251-MG.


view in a new=20 window

Figure 1. The OD value = measured from=20 MTT assay represented cell viability of U251-MG after=20 different treatment at 1, 2, 3, and 4 days = respectively.=20 Results shown are the means of three independent = experiments.=20 *P <0.05 as compared with the = control=20 group, =86P <0.05 as compared = with the TMZ=20 group.

Combination of TMZ with THD induced = cell cycle=20 arrest of G0/G1 phase on U251-MG
Results of cell = cycle=20 analysis shown in Table indicated that after different = treatments=20 for 3 days, TMZ alone increased the population of the G2/M = phase=20 (P <0.01), while TMZ with THD significantly = increased the=20 population of G0/G1 phase (P <0.01). The results = indicated=20 that TMZ alone induced G2/M phase arrest in tumor cells = while TMZ=20 combined with THD advanced the cycle arrest to G0/G1 phase, = which=20 demonstrated a promoted inhibition of cell = proliferation.


view in a new=20 window
Table. Cell cycle = distribution of=20 U251 cells treated with TMZ and THD for 3=20 days

Ultrastructural features of = autophagy were=20 observed after TMZ and THD treatment
As shown in = Figure 2,=20 while untreated tumor cells exhibited few such features = (Figure 2A),=20 autophagosomes and secondary lysosomes were observed in TMZ = treated=20 and combined treatment group (Figure 2B=A8C2E).The sum of=20 autophagosomes in TMZ+THD group was definitely larger than = that of=20 the TMZ group. Most of the autophagosomes contained lamellar = structures (Figure 2D) or residual digested materials = (Figure 2E).=20 These results indicated that U251-MG cells treated with TMZ = for 3=20 days were in the stage of autophagic process. No apoptotic = body was=20 found in either group, which indicated that only autophagy = was=20 induced by TMZ rather than apoptosis.


view in a new=20 window

Figure 2. Ultrastructure = features of=20 U251-MG after different treatments for 3 days with = electron=20 microscopic observation, N indicated nucleus. = A:=20 Untreated cells. B: TMZ (100 =A6=CCmol/L) for 3 = days. C:=20 TMZ (100 =A6=CCmol/L)+THD (100 =A6=CCg/L) for 3 = days. D:=20 Autophagosomes including lamellar structure. E: = Autophagosomes including residual digested material. = The=20 arrows in the pictures indicate autophagosomes in=20 cytoplasma.

Development of AVOs in TMZ with THD = treated=20 U251-MG cells
In the results of AVOs detection with = flow=20 cytometry, the percentage of cells in Q1 and Q2 quadrants = stands for=20 positive-staining rate. As shown in Figure 3, after 3 days=20 treatment, compared with TMZ used alone (10.41%), the = combination of=20 TMZ with THD significantly increased the number of AVOs = (36.05%),=20 which represents the degree of autophagy in cells. THD used = alone=20 did not increase AVOs obviously. The result indicated that = THD may=20 enhance the TMZ induced autophagy, but THD itself did not = do.


view in a new=20 window

Figure 3. Quantification of = AVOs with=20 flow cytometry in U251-MG after different treatments = for 3=20 days. I: control; II: TMZ (100 = =A6=CCmol/L)=20 treatment. III: TMZ (100 =A6=CCmol/L) with THD = (100 =A6=CCg/L)=20 treatment. IV: THD (100 =A6=CCg/L) treatment. = The data=20 shown are the addition percentage from Q1 and Q2=20 quadrants.

MDC accumulation increases after = treatment of=20 TMZ with THD
The fluorescent compound, MDC, could be=20 specifically accumulated in autophagic vacuoles. As shown in = Figure=20 4A, by using fluorescence microscope, autophagic vacuoles = labeled by=20 MDC were observed concentrated in spherical structures = distributed=20 in the cytoplasm and also localized in the perinuclear = region,=20 probably corresponding to the microtubule organizing center. = In the=20 control and THD group, MDC stained U251-MG was hardly found, = nevertheless, there were many MDC stained U251-MG in TMZ and = TMZ=20 with THD group. Quantification of MDC staining positive = cells were=20 determined by flow cytometry. Results were shown in Figure = 4B.The=20 ratio of MDC positive staining cell in TMZ with THD group = was higher=20 than that in the TMZ group, which indicated that more = autophagic=20 vacuoles were formed in the combined treatment group than = TMZ alone=20 group. The result further demonstrated that THD may = reinforce the=20 TMZ induced autophagy.


view in a new=20 window

Figure 4. A: (a) MDC = staining=20 negative U251-MG; (b=A8Cd) MDC staining = positive=20 U251-MG, the arrow heads show MDC labeled autophagic = vacuoles.=20 B: Different ratios of MDC positive staining = cells were=20 shown between treatment groups (%): TMZ group: = 10.04=A1=C02.48;=20 THD group: 1.65=A1=C00.80; TMZ with THD group: = 22.32=A1=C03.06;=20 control group: 1.18=A1=C00.24. Results shown are the = means of=20 three independent experiments, *P = <0.05=20 as compared with the control=20 group.

Combination of TMZ with THD = increased the=20 expression of MAP1LC3, Beclin1 and PTEN = protein
Micro-tubule=20 associated protein 1 light chain 3 (MAP1LC3 or LC3) is an = attractive=20 biomarker for autophagy because it functions at least in = part as a=20 structural component during autophagosome = formation.12=20 Beclin1 is a tumor suppressor gene and induces autophagy = when=20 over-expressed in mammalian cells.13 It may = activate and=20 bind with Type-III PI3K to form the complex attending in the = formation of autophagosome. Phosphatase and tensin homolog = deleted=20 on chromosome ten (PTEN) is a tumor suppressor which = inhibits the=20 class I phosphatidyl-inositol 3-kinase (PI3K)/Akt/mTOR = signaling=20 pathway,14 while the activation of this signaling = pathway=20 may suppress autophagy. We detected LC3, Beclin1 and PTEN = protein=20 expression after different treatments with Western blotting. = An=20 illustration should be made before showing the result. Two = forms of=20 LC3, the cytosolic LC3-I and the membrane-bound LC3-II, are = produced=20 post-translationally. When autophagy is induced, some LC3-I = is=20 converted into LC3-II. The amount of LC3 II or the LC3 II = /LC3 I=20 ratio correlates with the number of autophagosomes. In this = study,=20 the amount of LC3 II was used to evaluate autophagy. = Beta-actin was=20 used as the internal control to calculate the relative = expression=20 level.

The results were shown in Figure 5, after = 3 days=20 treatment, compared with the control group, the protein = expression=20 of LC3II and Beclin1 was up-regulated in TMZ group and TMZ = with THD=20 group but almost stable in THD group (LC3II: = P=3D0.227;=20 Beclin1: P=3D0.229). The up-regulation of LC3II and = Beclin1=20 expression were more significant in TMZ with THD group than = that in=20 TMZ group (P <0.05). While compared with = the=20 control group, the level of PTEN was more obviously elevated = in THD=20 and TMZ with THD group (P <0.05). No = difference was=20 shown between these two groups (P=3D0.716). No = significant=20 alteration of PTEN expression was shown in TMZ group = compared with=20 the control group (P=3D0.635).The result = indicated:compared=20 with TMZ used alone, the combination of THD with TMZ = increased the=20 expression level of LC3II and Beclin1. THD rather than TMZ = promoted=20 the expression of PTEN.


view in a new=20 window

Figure 5. LC3II, Beclin1 and = PTEN=20 protein expression after different treatments for 3 = days in=20 U251-MG. A: Western blotting analysis. = B: The=20 densitometry was applied to qualify the protein = expression of=20 the Western blotting. n=3D3, *P=20 <0.05 compared with the=20 control.

DISCUSSION

This study focused on the synergistic = mechanism of=20 THD to enhance the cytotoxicity of TMZ. In the present = study, we=20 confirmed TMZ may induce autophagy in glioma cells, which = resulted=20 in proliferative inhibition of the tumor cells and cell = cycle arrest=20 at G2/M phase. Meanwhile we found that although THD used = alone was=20 not sufficient to inhibit tumor cell growth, its combination = with=20 TMZ suppressed the cell viability of U251-MG more = significantly than=20 TMZ used alone and advanced the cell cycle arrest to G0/G1 = phase. We=20 also demonstrated the enhancement of tumor growth inhibition = may=20 attribute to the promotion of TMZ induced autophagy by = THD.

Autophagy is a dynamic process of = cytoplasmic=20 protein and organelles degradation, which is observed in = hepatic=20 cells cultured in vitro during nutrient=20 deprivation.15 The process of autophagy begins = with=20 sequestering cytoplasmic protein or organelles in a membrane = vacuole=20 to form autophagosome. Autophagosomes then fuse with = lysosomes,=20 where the materials inside are degraded and recycled. = Normally=20 autophagy is reversible and acts as a self-protection = mechanism.=20 When the process of autophagy turns irreversible, it would = lead to=20 autophagic cell death which is also referred as Type II = programmed=20 cell death. Oncologists announced that different types of = cancer=20 cells undergo autophagy after various anticancer therapies.=20 Autophagy plays a key role in the elimination of cancer = cells by=20 triggering a non-apoptotic cell death program.16 = When=20 autophagic cell death is induced, autophagy functions as a=20 self-destructive mechanism in cancer cells. Besides the = signaling=20 pathway that would be discussed as follows, many other = factors also=20 attend the regulation of autophagy. For example, the Gai3 = protein,=20 as combined with GTP, the GTP-Gai3 is an inhibitor of = autophagy;=20 while GDP-Gai3 turns to be an autophagy activator. As a = member of=20 RGS (regulators of G-protein signaling) family, GAIP (G = alpha=20 interacting protein) promotes autophagy by accelerating the=20 hydrolysis of GTP combined with Gai3.17 AGS3 = (activator=20 of G-protein signaling 3) is another factor that would = enhance=20 autophagy. Amino acid, the product of autophagy, could = provide a=20 negative feedback regulation of autophagy. The removal of = exogenous=20 amino acid would block the mTOR signaling pathway, which = inhibits=20 autophagy, while the endogenous amino acid produced by = autophagy may=20 complement the deficiency of amino acid and that would = resume the=20 mTOR signaling pathway. What=A1=E4s more, hormone, insulin = and calcium=20 all play important roles in regulation of autophagy, but the = mechanisms are not poorly known.

PI3K/Akt/mTOR signaling pathway plays an = important=20 role in cellular proliferation, development and = death.18=20 It has been demonstrated the activation of PI3K/Akt/mTOR = signaling=20 pathway inhibits the process of autophagy.19 = PTEN, a=20 tumor suppressor, is a phosphatase with a variety of = substrate=20 specificities. It functions as a negative regulator of the=20 PI3K/Akt/mTOR signaling pathway.20 Leslie et=20 al21 reported PTEN is often absent or mutated in = many=20 malignant tumors, which leads to an attenuation of = autophagy.=20 Beclin1, another tumor suppressor gene, is essential for = autophagy.=20 It forms complex with class III PI3K and localizes at the=20 trans-Golgi network, suggesting that the Beclin 1-class III = PI3K=20 complex plays a role in sorting putative autophagosomal = components=20 and lysosomal protein.22 Moreover, some small = molecular=20 proteins are also involved in autophagy. LC3 functions as a=20 structural component during autophagosome formation. = Detection of=20 LC3 by immunoblotting or immunofluorescence is a reliable = method for=20 monitoring autophagy and autophagy-related processes, = including=20 autophagic cell death.

Vital cell staining is another dependable = method to=20 investigate autophagy. As described previously, MDC and = acridine=20 orange staining are reliable but simple methods to detect = autophagy.=20 They may be used as qualitative methods as well as = quantitative=20 methods.

In our study, after exposure to TMZ (100 = =A6=CCmol/L)=20 with THD (100 =A6=CCg/L) for 3 days, the acridine orange and = MDC=20 staining positive tumor cells increased significantly which=20 indicated a promotion of autophagy. Considering the enhanced = inhibition of cell proliferation in TMZ with THD group, we = inferred=20 that autophagic cell death was induced. To find out the = mechanism of=20 THD to reinforce the effect of TMZ, we detect the expression = of=20 MAP1LC3 and Beclin1, both of which play crucial roles in the = process=20 of autophagy, at translational level. The result of = immunblotting=20 showed the expression of MAP1LC3 and Beclin1 was = up-regulated in=20 both TMZ group and TMZ with THD group, while the result of = TMZ with=20 THD group was more significant. Interestingly, no = significant=20 difference was shown between THD and negative control group. = So it=20 seems THD may up-regulate the expression of = autophagy-associated=20 protein indirectly. Knobloch et al23 reported = THD-induced=20 signaling protects active PTEN from proteasomal degradation, = resulting in suppression of Akt signaling. Enlightened by = this view,=20 we detected the protein expression level of PTEN in = different=20 treatment groups. The results of THD and THD with TMZ group = were=20 obviously higher than those of TMZ group. So we concluded = the=20 up-regulation of PTEN may facilitate the process of = autophagy rather=20 than induce autophagy. In other words, PTEN is a = facilitative but=20 not crucial factor to promote autophagy. The evidence is = that=20 despite THD up-regulated the expression of PTEN, essential=20 autophagy-associated protein MAP1LC3 and Beclin1 stayed = still in THD=20 alone group; while in TMZ with THD group, both of the = expression=20 level of PTEN and autophagy-associated protein were = up-regulated. It=20 seems that more factors would participate in and regulate = the=20 process of autophgy and further experiments are necessary to = clarify=20 them.

It should be noted that the study of = synergistic=20 mechanism only focused on the translational level. More = small=20 molecules involved in autophagy and PI3K/Akt/mTOR signaling = pathway=20 should be detected. Despite its preliminary character, this = study=20 freshly indicated: besides anti-angiogenesis effect, THD may = also=20 reinforce the effect of TMZ by promoting autophagy. = Furthermore, our=20 study provided the experimental basis for more reasonable = and=20 effective clinical use of combination of TMZ with THD. = Kanzawa et=20 al5 once performed the similar experiment in = U87-MG and=20 A172 cell lines. The result was close to ours, while they = announced=20 inhibition of autophagy at a late stage could increase the=20 therapeutic efficacy of TMZ for malignant glioma cells. It = seems=20 that the combination of TMZ with THD, which promote = autophagy, may=20 not benefit the patient with gliomas at a late stage of = treatment.=20 Moreover, we applied a series of methods to study autophagy = and put=20 forward the idea to combine the research of effectors in=20 PI3K/Akt/mTOR signaling pathway with autophagy regulation. = The=20 methods and idea provided a fresh development in the = research of=20 autophagy.

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