Blackwell Synergy: J Neurochem, Vol 0, Issue 0, pp. = ???-???: The natural compound n-butylidenephthalide derived from = Angelica sinensis inhibits malignant brain tumor growth in vitro and in = vivo3 (Full Text)

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Journal of=20 Neurochemistry
Online=20 Early
doi:10.1111/j.1471-4159.2006.04151.x

Volume 0 = Issue=20 0

The natural = compound=20 n-butylidenephthalide derived from Angelica=20 sinensis inhibits malignant brain tumor growth = in=20 vitro and in vivo³

Nu-Man = Tsai*^,1,=20 Yi-Lin Chen=86^,1,=20 Chau-Chin Lee=87^,1,=20 Po-Chen Lin=A7,=20 Yeung-Leung Cheng=B6, Wen-Liang = Chang**, = Shinn-Zong=20 Lin=A7^,=86^,2=20 and Horng-Jyh Harn=A7^,==86^,=86=86^,=87=87^,2

Abstract

The naturally-occurring compound,=20 n-butylidenephthalide (BP), which is isolated = from the=20 chloroform extract of Angelica sinensis (AS-C), = has=20 been investigated with respect to the treatment of = angina. In=20 this study, we have examined the anti-tumor effects of = n-butylidenephthalide on glioblastoma = multiforme (GBM)=20 brain tumors both in vitro and in vivo. = In=20 vitro, GBM cells were treated with BP, and the = effects of=20 proliferation, cell cycle and apoptosis were = determined. In=20 vivo, DBTRG-05MG, the human GBM tumor, and RG2, = the rat=20 GBM tumor, were injected subcutaneously or = intracerebrally=20 with BP. The effects on tumor growth were determined = by tumor=20 volumes, magnetic resonance imaging and survival rate. = Here,=20 we report on the potency of BP in suppressing growth = of=20 malignant brain tumor cells without simultaneous = fibroblast=20 cytotocixity. BP up-regulated the expression of Cyclin = Kinase=20 Inhibitor (CKI), including p21 and p27, to decrease=20 phosphorylation of Rb proteins, and down-regulated the = cell-cycle regulators, resulting in cell arrest at the = G₀/G₁ phase for DBTRG-05MG and = RG2=20 cells, respectively. The apoptosis-associated proteins = were=20 dramatically increased and activated by BP in = DBTRG-05MG cells=20 and RG2 cells, but RG2 cells did not express p53 = protein.=20 In vitro results showed that BP triggered both=20 p53-dependent and independent pathways for apoptosis. = In=20 vivo, BP not only suppressed growth of = subcutaneous rat=20 and human brain tumors but also, reduced the volume of = GBM=20 tumors in situ, significantly prolonging = survival rate.=20 These in vitro and in vivo anti-cancer = effects=20 indicate that BP could serve as a new anti-brain tumor = drug.

Glioblastoma multiforme (GBM), the extreme = expression of=20 anaplasia among the glial neoplasmas, accounts for 40% = of all=20 primary intracranial tumors (Mahaley = et al.=20 1989; Graham=20 and Lantos 2002). The diffusely invasive = properties of GBM=20 mean that total resection is almost impossible and = therefore,=20 surgery plus radiotherapy, and eventual chemotherapy, = is the=20 standard treatment (Santarius = et al.=20 1997; Graham=20 and Lantos 2002; Giese = et al.=20 2003). Nonetheless, the clinical course rarely = exceeds=20 18 months from the time of diagnosis, regardless = of=20 therapeutic intervention (Santarius = et al.=20 1997; Graham=20 and Lantos 2002).

Chemotherapy is usually reserved for recurrent = tumors that=20 have already been treated with surgery and = radiotherapy, or=20 for tumors in which surgery was only partial or = infeasible and=20 the effect of radiotherapy was limited (Blacklock = et al.=20 1986; Shapiro=20 and Green 1987). Various chemotherapy schemes have = been=20 used, with most consisting of administering drugs at = high=20 doses (Shapiro and=20 Green 1987). The present chemotherapy regimens are = limited=20 for several reasons, including development of toxicity = and=20 drug resistance, limited success in overcoming the = blood=96brain=20 barrier (BBB) and limited therapeutics (Blacklock = et al.=20 1986; Shapiro=20 and Green 1987; Elliott = et al.=20 1996; Bello=20 et al. 2001; Cheng = et al.=20 2004).

Angelica sinensis (Oliv.) Diels (AS; dong = quai; also=20 called danggui), a traditional Chinese medicine for = menopausal=20 symptoms (Ye=20 et al., 2001a; Ye = et al.,=20 2001b; Yim=20 et al. 2002), has been clinically=20 administrated for several gynecological symptoms in = the US (Abebe = 2002). In=20 our previous study, the chloroform extract of=20 A. sinensis (AS-C) also showed a dramatic=20 anti-tumor effect, causing growth resting and = apoptosis of=20 malignant brain tumors in vitro and in = vivo, and=20 both p53-dependent and -independent pathways of = apoptosis were=20 involved in the cytotoxic mechanisms (Tsai = et al.=20 2005). Thus far six major compounds have been = isolated=20 form A. sinensis: (E)-liguistilide,=20 (Z)-ligustilide, (Z)-n-butylidenephthalide, = palmitic=20 acid, beta-sitosterol and ferulic acid (Wang = et al.=20 1998). n-Butylidenephthalide (BP, BdPh or = K1;=20 molecular weight approximately 188.22) derived from = AS-C was=20 the major component (over 30% of AS-C crude) and was = therefore=20 chosen in this further study to verify its anti-tumor=20 activity.

Several medicinal properties of BP, such as the=20 anti-platelet effect, are due mainly to an inhibitory = effect=20 on cyclo-oxygenase (Teng = et al.=20 1987). It also provides relief from angina (Ko = et al.=20 1998, 2002). In addition, it has been suggested = that the=20 anti-proliferation effect of synthetic BP-42=20 (3-butylidene-4,5-dihydroxyphthalide), which was = modified by=20 adding the hydroxyl molecules in the phthalide group = of BP,=20 could be used as an anti-atherosclerotic treatment (Mimura=20 et al. 1995). However, the anti-tumor=20 activities of BP have not yet been determined. In this = study,=20 the anti-tumor effects of BP on malignant brain tumors = in=20 vitro and in vivo were investigated. The = results=20 revealed that the significant therapeutic anti-tumor = efficacy=20 of BP on GBM tumors involved the induction of = cell-cycle rest=20 and apoptosis.

Materials and = methods

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Top of pageResultsDiscussionAcknowledgementsReferencesAbbreviations = used

Chemicals

n-Butylidenephthalide (BP; molecular weight: = 188.23), purchased from Lancaster Synthesis Ltd = (Newgate,=20 Morecambe, UK) and carmustine (BCNU; Sigma Chemical = Co., St=20 Louis, MO, USA) were dissolved in dimethylsulfoxide = (DMSO) and=20 ethanol, incubated with shaking at 25=B0C for = 1 h, and=20 stored at 4=B0C before each in vitro = experiment.

Cell lines and cell = culture

The DBTRG-05MG line of human GBM cells, RG2 line of = rat GBM=20 cells, SK-N-AS line of human neuroblastoma cells, SVEC = line of=20 mouse vascular endothelial cells and Balb/3T3 line of = mouse=20 fibroblast cells were obtained from the American Type = Culture=20 Collection (Rockville, MD, USA). G5T/VGH human GBM = cells, GBM=20 8401 human GBM cells, GBM 8901 human GBM cells, A549 = human=20 lung adenocarcinoma cells, PA-1 human teratoma cells, = B16/F10=20 mouse melanoma cells, HL-60 human leukemia cells and = N18 mouse=20 neuroblastoma cells were obtained from the = Bioresources=20 Collection and Research Center (BCRC, Hsin Chu, = Taiwan). The=20 J5 line of human hepatocellular carcinoma cells and = BCM-1 line=20 of human breast cancer cells were kindly provided by = Drs M. J.=20 Chou and C. S. Yang (Sherr and Robert = 1995) and=20 Dr D. S. Yu (Reed=20 et al. 1994; Dobashi = et al.=20 2003), respectively. The DBTRG-05MG, GBM8401, = GBM8901,=20 BCM-1, HL-60, A549, PA-1 and J5 cells were maintained = in=20 RPMI-1640 (Sigma) medium with 10% fetal bovine serum, = at 37=B0C=20 in a humidified atmosphere containing 5% = CO₂. The=20 G5T/VGH, RG2, SK-N-AS, N18, B16/F10, SVEC and Balb/3T3 = cells=20 were cultured in Dulbecco's modified Eagle's medium = with 10%=20 fetal bovine serum at 37=B0C in a humidified = atmosphere=20 containing 5% CO₂.

Cytotoxicity analysis

Viable cells were evaluated by a modified=20 3-(4,5-dimethylthiazol-2-yl)-2,5-diphenyltetrazolium = bromide=20 (MTT) assay. Briefly, 5 =D7 10³ = cells were=20 plated in each well of a 96-well plate, incubated = overnight=20 with 100 =B5L growth medium in 96-well plates, = then treated=20 with 100 =B5L BP dissolved in the medium=20 (0=96250 =B5g/mL). The concentration of DMSO=20 was ≤ 0.02% in each preparation. After = 24, 48 or=20 72 h of BP treatment, the medium was replaced = with=20 50 =B5L fresh medium containing MTT = (400 =B5g/mL) for=20 6=968 h. The MTT medium was then removed and = 100 =B5L=20 DMSO added to each well. The solutions were detected = using an=20 MRX Microtiter Plate Luminometer (Dynax Technologies,=20 Chantilly, VA, USA) at 550 nm. Absorbance of = untreated=20 cells was considered as 100%. The 50% inhibitory = concentration=20 (IC₅₀) was the concentration that = caused a=20 50% decrease in the optical density of drug-treated = cells with=20 respect to untreated cells.

Analysis of cell-cycle=20 distribution

The GBM tumor cell lines DBTRG-05MG and RG2 were = cultured=20 with drug diluent for 48 h; DMSO content was = controlled=20 at ≤ 0.02% and BP at 75 =B5g/mL. = Cell-cycle=20 analysis was performed by DNA staining with propidium = iodide=20 (PI) and flow cytometry. Briefly, cells were = harvested,=20 resuspended in 0.8 mL = 1 =D7 phosphate-buffered=20 saline (PBS) and added to 200 =B5L PI solution=20 (50 =B5g/mL PI + 0.05 mg/mL RNase = A; Sigma=20 Chemical Co.). After overnight incubation at 4=B0C, = cells were=20 kept at 25=B0C for 2 h. The cells were then = passed through=20 FACScan (Becton Dickinson Immunocytometry Systems, San = Jose,=20 CA, USA) to measure the DNA content.

Determination of = apoptosis

Apoptosis was assayed using an in Situ Cell = Death=20 Detection Kit, POD (Roche, Mannheim, Germany), = according to=20 the manufacturer's instructions. Briefly, DBTRG-05MG = and RG2=20 cells were cultured in culture dishes and analyzed at = the=20 indicated time points (24, 48, 72 h) after BP=20 (75 =B5g/mL) treatment. In the BP-treated cell = group,=20 suspended cells were collected; in the control group, = adherent=20 cells were collected. Cells were fixed at 25=B0C for = 15 min=20 with 3.7% formaldehyde, smeared on silane-coated glass = slides=20 (Muto Pure Chemicals, Tokyo, Japan), washed once in=20 1 =D7 PBS and incubated in cold = permeabilization=20 solution (0.1% Triton X-100 + 0.1% sodium = citrate)=20 after reducing endogenous peroxidase enzyme activity = with 3%=20 H₂O₂. Then, cells were incubated = with=20 terminal deoxynucleotidyl transferase (TdT)-mediated = dUTP nick=20 end labeling (TUNEL) reaction mixture for 60 min = at 37=B0C,=20 and counterstained with PI for determination of the = cell=20 count. To quantify cell apoptosis, the slides were = viewed=20 under fluorescence microscopy (Nikon, Kawasaki, = Japan).

Western blot analysis

DBTRG-05MG and RG2 cells were seeded in a six-well = plate=20 and later treated with BP (75 =B5g/mL) for = different times=20 (0, 1.5, 3, 6, 12, 24 or 48 h; 0 h as a = vehicle=20 control). Cell pellets were resuspended in lysis = buffer=20 [10 nm Tris-HCl=20 (pH 7.5), 1 mm EGTA,=20 0.5%=20 = 3-[3-(cholamidopropyl)dimethylammonio]-1-propanesulfonate, 10%=20 (v/v) glycerol, 5 mmβ-2-mercaptoethanol and = 0.1 mm = phenylmethylsulfonyl=20 fluoride] and incubated on ice for 30 min. After=20 centrifugation at 13 000 r.p.m. for = 20 min at=20 4=B0C, total cell lysates were collected. The protein=20 concentration of the cell lysates was measured with a=20 bicinchoninic acid (BCA) protein assay kit (Pierce, = Rockford,=20 IL, USA) following the manufacturer's instructions. = Aliquots=20 (20 =B5g) of the cell lysates were separated by = 10=9612%=20 sodium dodecyl sulfate=96polyacrylamide gel = electrophoresis=20 (SDS=96PAGE; Bio-Rad, Hercules, CA, USA) and = transferred to=20 polyvinylidenedifluoride (PVDF) membranes (Amersham=20 Lifesciences, Piscataway, NJ, USA). The membranes were = blocked=20 with 5% skim milk for 1 h at 25=B0C, then = incubated with=20 the respective antibodies. The antibodies included = anti-Fas,=20 anti-Fas-L, anti-caspase 3, anti-caspase 8, = anti-caspase 9,=20 anti-Bax, anti-AIF, anti-p16, anti-p21, anti-p27, = anti-p53,=20 anti-cdk2, anti-cdk4, anti-cdk6, anti-cyclin D1, = anti-cyclin=20 E, anti-actin (1/200 dilution; Santa Cruz = Biotechnology Inc.,=20 Santa Cruz, CA, USA); anti-phospho-p53 (Ser15; 1/2000=20 dilution), and anti-phospho-RB (Ser795; 1/2000 = dilution; Cell=20 Signaling Technology, Beverly, MA, USA).The = immobilized=20 primary antigen-antibody complex was detected with the = respective horseradish peroxidase-conjugated = anti-mouse,=20 anti-rabbit or anti-goat IgG secondary antibodies = (1/1000=20 dilution; Santa Cruz Biotechnology Inc.) for 1 h = at 25=B0C,=20 then visualized with an enhanced chemiluminescence = (ECL) Plus=20 chemiluminescence system (Amersham, Arlington Heights, = IL,=20 USA). The degree of protein expression was calculated = as the=20 expression index. The expression indexes were = calculated as=20 [(sample intensity/sample β-actin = intensity)/(vehicle control=20 intensity/vehicle control β-actin = intensity)].

Animal studies

To examine the anti-tumor effects of BP in = vivo, RG2=20 rat GBM cells were used in male F344 rat = (230=96260 g)=20 experiments, and DBTRG-05MG human GBM cells were used = in male=20 Foxn1 nu/nu mouse (10=9612 weeks) = experiments. The=20 rats and mice were purchased from the National = Laboratory=20 Animal Center (Taipei, Taiwan). All procedures were = performed=20 in compliance with the standard operating procedures = of the=20 Laboratory Animal Center of Tzu Chi University = (Hualien,=20 Taiwan).

In vitro cultured RG2 cells=20 (1 =D7 10⁶) were injected s.c. = into the=20 back of syngeneic F344 rats. Six animals in each = vehicle or BP=20 group were treated on days 3, 6 and 9 after tumor cell = implantation by s.c. injection of BP = (500 mg/kg/day) or=20 vehicle (10 mg/mL Tween-80 and 50 mg/mL = propylene=20 glycol in distilled water; Standard Chem. & = Pharm.,=20 Tainan, Taiwan), far from the inoculated tumor cell = sites=20 (> 1.5 cm). In addition, DBTRG-05MG cells = (2.5 =D7 10⁶) were injected s.c. = into the=20 backs of nude mice (five to six/group), and treated = with=20 vehicle, BP−70, −150, −300, = −500 and −800 mg/kg/day=20 (s.c.) respectively. The injection site was far from = the=20 inoculated tumor sites (>1.5 cm away from = treated=20 site), on days 4 , 5, 6, 7 and 8 after tumor cell=20 implantation. Tumor size was measured with calipers = and the=20 volume was calculated as=20 L =D7 H =D7 W =D7 0.5236 = (Bardon = et al.=20 2002). Animals were killed when the tumor volume = exceeded=20 25 cm³ in rats and = 1500 mm³ in=20 mice; the day on which an animal was killed was = designated as=20 the final survival day for the rats and mice.

The anti-tumor effects of BP on in situ = tumors were=20 determined with RG2 cells. The RG2 cells were injected = i.c.=20 into the striatum of syngenic rats (six/group) and = treated=20 with BP (300 mg/kg/day) or vehicle s.c. on days = 4, 5, 6,=20 7 and 8 after tumor cell implantation. Tumor volume = was=20 measured using 3-T unit magnetic resonance imaging = (MRI;=20 General Electric, Milwaukee, Wisconsin, USA) with = echo-planar=20 imaging capability (Fukuoka = et al.=20 2000) (Signa LX 3.8; General Electric) in Buddhist = Tzu Chi=20 General Hospital (Hualien, Taiwan). Briefly, rats were = anesthetized with chloral hydrate (400 mg/mL,=20 1 mL/100 g). Functional MRI scanning was = conducted=20 with a fast spin echo and an echo-planar acquisition = sequence=20 in which the repetition time was 6000 ms, the = echo time=20 102 ms, the matrix image 256 =D7 256 = pixels, the=20 field of view 5 =D7 5 cm and the = in-plane=20 resolution, 80 =B5m. Twenty slices, each = 1.5 mm thick,=20 were obtained every 19.5 s for a total time of=20 6.5 min/rat.

Immunohistochemical = staining

All tumor tissues (s.c. or i.c. GBM tumors with or = without=20 BP treatments) were harvested and fixed. = Paraffin-embedded=20 sections (7 =B5m/section) were obtained from the = tumors and=20 processed for immunohistochemical staining. A goat = polyclonal=20 anti-Ki-67 antibody (M-19; 1/100 dilution; Santa Cruz=20 Biotechnology Inc.) and a rabbit polyclonal = anti-cleaved=20 caspase 3 (Asp175) antibody (1/1000 dilution; Cell = Signaling=20 Technology.) were used in immunohistochemical studies = with 4=B0C=20 overnight incubation. The immune complexes were = visualized=20 using the horseradish peroxidase-conjugated anti-goat = IgG=20 secondary antibodies (1/1000 dilution; Santa Cruz=20 Biotechnology Inc.) and LSAB2 system (DAKO, Corp.,=20 Carpinteria, CA, USA), respectively, and then = incubated for=20 10 min with 0.5 mg/mL diaminobenzidine and = 0.03%=20 (v/v) H₂O₂ in PBS. Finally, = sections=20 were counterstained with hematoxylin, mounted, = observed under=20 a light microscope at magnifications of 400=D7, and=20 photographed.

Statistics

Data are presented as mean =B1 SD or SE = (standard=20 deviation and standard error, respectively). = Statistical=20 significance was analyzed by Student's t-test. = The=20 survival analysis was performed using the = Kaplan=96Meier method.=20 A p-value of < 0.05 was considered to be=20 statistically significant.

Results

Go to:

Top of pageMaterials and = methodsResults <<DiscussionAcknowledgementsReferencesAbbreviations = used

Cytotoxic effects of BP on = tumor and=20 other cell lines

The anti-proliferative effects of BP on cells from = GBM,=20 neuroblastoma, lung cancer, melanoma, teratoma, = leukemia,=20 breast cancer and hepatocellular carcinoma, as well as = from=20 normal fibroblast and vascular endothelial cells, were = determined. The IC₅₀ values of BP = after a=20 48 h incubation with tumor cell lines=20 (IC₅₀: 15=9667 =B5g/mL) were = significantly=20 lower than the values for normal fibroblast cells=20 (IC₅₀: > 100 =B5g/mL,=20 p < 0.0001) or carmustine = (BCNU)-treated=20 tumor cells (IC₅₀: 40 100 =B5g/mL,=20 p < 0.001; Table 1). In = the two=20 normal cell types, vascular endothelial cells=20 (IC₅₀ =3D=20 25.0 =B1 2.0 =B5g/mL) were more = sensitive to BP=20 than fibroblast cells=20 = (IC₅₀ > 100 =B5g/mL)=20 (p < 0.0001). Briefly, these = experiments=20 showed that BP could induce high cytotoxicity against = brain=20 and other tumor cells but only low, or no cytotoxicity = against=20 normal fibroblast cells.

The morphology of GBM tumor cells gradually changed = with=20 detachment from the bottom of the culture plates, = after BP=20 treatment (data not shown). The TUNEL assay was used = to verify=20 the viability of detached cells and the results showed = that=20 the cells were in the process of apoptosis at = 48 h (Figs 1a and=20 b). Therefore, BP has a cytotoxic ability against = tumor=20 cells.

The effects of BP on cell cycle = of GBM=20 cells

Cell-cycle analysis with GBM cells demonstrated = that=20 75 =B5g/mL BP treatment resulted in cell-cycle = arrest at=20 the G₀/G₁ phase (> 90% = and >=20 70%; Figs 1c=20 and d). BP induced a significant proportion of = cells to=20 arrest at the G₀/G₁ phase, = accompanied=20 by a concurrent decrease of S phase from 12 to = 24 h=20 (p < 0.05). In addition, BP also = led to a=20 decrease in the proportion of the DBTRG-05MG cells = that=20 entered the G₂/M phase, though more cells=20 accumulated at G₂/M phase in RG2 cells. The = results=20 at 48 and 72 h were similar to those at 24 h = (data=20 not shown).

Apoptotic pathways in GBM cells = induced=20 by BP

To investigate apoptotic pathways induced by BP = treatment,=20 phosphorylation of p53 and Rb proteins were first = evaluated by=20 immunoblotting. The DBTRG-05MG and DBTRG 8401 line of = human=20 GBM cells, and RG2 line of rat GBM cells, were used = for=20 comparison. BP caused levels of phosphorylated p53 = protein to=20 increase (9.4-fold) 3 h after treatment (Fig. 2a). = Furthermore,=20 total p53 protein increased (twofold) at 12 h in=20 DBTRG-05MG and DBTRG 8401 (twofold) increased at = 3 h. In=20 RG2 cells, the genes of p53 and p16 are impaired and = exhibit=20 homologous deletion, and these were undetectable in=20 immunoblotting analysis. However, levels of = phosphorylated Rb=20 proteins were decreased after BP treatment in = DBTRG-05MG and=20 DBTRG 8401 cells; in RG2 cells, the levels were = decreased=20 (0.8-fold) at 1.5 h, with phosphorylated proteins = undetectable as early as 3 h after BP treatment = (Fig. 2a).=20 These results indicate that BP can trigger cell-cycle=20 checkpoint machinery. Levels of p16, p21, Bax and AIF = protein=20 in BP-treated GBM cells were therefore measured, and = protein=20 expression levels of all four proteins increased in = DBTRG-05MG=20 and DBTRG 8401 cells; similar results for p27, Bax and = AIF=20 were detected in RG2 cells after treatment with BP (Figs 2b and=20 c). The protein expression of cdk2, cdk4, cdk6, = cyclin=20 D₁ and cyclin E in BP-treated GBM cells was = also=20 measured, and all five proteins were decreased in = DBTRG-05MG=20 and DBTRG 8401 cells; similar results for cdk2, cdk4, = cdk6,=20 cyclin D₁, cyclin E and p21 were detected = in RG2=20 cells after treatment with BP (Fig. 2b).

Expression of the proteins Fas and Fas-L in treated = DBTRG-05MG, DBTRG 8401 and RG2 cells was studied. = Results=20 showed that BP significantly induced Fas expression = (from one=20 to 41.3-fold vs. one to 7.7-fold in DBTRG-05MG and RG2 = cells,=20 respectively) but not Fas-L expression. In addition,=20 activation of death receptor-induced apoptosis-related = caspase=20 8 protein was monitored. Levels of caspase 8 in = DBTRG-05MG and=20 DBTRG 8401 were increased in a dose-dependent manner = from 6 to=20 48 h after BP treatment, whereas the maximum = expression=20 of caspase 8 in RG2 cells at 24 h (Fig. 2d).

Finally, activation of pro-caspase 9 and procaspase = 3 was=20 also determined (Figs 2c and = d). Both=20 procaspase 9 and procaspase 3 were highly activated in = DBTRG-05MG, DBTRG 8401 and RG2 cells after BP = treatment.

Anti-tumor effects of BP on = survival of=20 animals bearing subcutaneous GBM tumor

To investigate BP anti-tumor activity, in = vivo=20 animal experiments were carried out. There was a = significantly=20 higher inhibitory effect on RG2 tumor growth in the = BP-treated=20 group than in the control (vehicle only) group (Fig. 3a;=20 p < 0.005). The average tumor size = at day=20 26 was 20.7 =B1 1.5 cm³ for = the=20 control group, in contrast to a size of only=20 9.6 =B1 0.4 cm³ for the = BP-treated=20 group. Survival of rats in the BP treatment group was=20 significantly prolonged compared with survival in the = control=20 group (30 =B1 2.1 days vs.=20 41.5 =B1 4.2 days; Fig. 3b;=20 p < 0.0001). There were no significant=20 differences between the control and BP-treated groups = with=20 respect to the body weight of the rats after BP = treatment (Fig. 3c).=20 There were remarkable decreases in Ki-67 (a cell = proliferation=20 marker) protein expression in this study, which = indicated that=20 BP had anti-proliferation activity in vivo. = Cleaved=20 caspase 3 is an apoptotic marker, and the results = showed that=20 BP treatment increased the cleavage of caspase 3 = protein to=20 induce apoptosis of tumor cells in vivo (Fig. 3d). = Both the=20 inhibition of tumor cell growth and the induction of = tumor=20 cell apoptosis pathways were mediated by BP treatment = in=20 vitro and in vivo. With s.c. injection of = BP at a=20 dose of 500 mg/kg, no drug-related toxicity was = observed=20 based on the histological analysis of organs in = animals=20 treated with BP (data not shown).

Anti-tumor effects of BP on = in=20 situ GBM tumor of rats

To verify the BP anti-tumor effects on the in = situ=20 GBM tumors in rats, F344 rats were implanted i.c. = (striatum)=20 with 5 =D7 10⁴ RG2 cells and = treated with=20 s.c. BP (300 mg/kg/day) on days 4, 5, 6, 7 and 8. = MRI=20 data revealed that the in situ tumor volumes in = the=20 BP-treated group were smaller than those in the = control group=20 (Fig. 4a). = There were=20 significant declines in tumor volume for the treated = group=20 relative to the untreated group (Fig. 4b;=20 p < 0.05). The mean tumor volumes = at days=20 14 and 16 were = 70.0 =B1 4.8 mm³ and=20 126.4 =B1 11.1 mm³ for the = control=20 group vs. 46.6 =B1 1.8 mm³ = and=20 91.7 =B1 8.3 mm³ for the = BP-treated=20 group. The immunohistochemistry results on day 16 = after BP=20 treatment showed a decrease in Ki-67 protein = expression, an=20 increase in cleaved caspase 3 protein expression and = an=20 increase in apoptosis in tumor cells relative to the = control=20 group in vivo (Fig. 4c); = these=20 results are similar to those shown in the subcutaneous = RG2=20 tumor model. Survival was significantly prolonged for = the=20 BP-treated rats compared with rats of the control = group. At=20 day 19 after BP treatment, BP caused higher survival = rates=20 (50%, 3/6) than the untreated group (16.7%, 1/6)=20 (p =3D 0.0016; data not shown).

Anti-tumor effects of BP on = xenograft=20 tumor growth

To determine whether BP can suppress human GBM = tumor=20 growth, nude mice were inoculated with human = DBTRG-05MG cells=20 and treated with BP (70, 150, 300, 500 and = 800 mg/kg in=20 the BP-70, BP-50, BP-300, BP-500 and BP-800 groups,=20 respectively.) on days 4, 5, 6, 7 and 8. Significant=20 suppression of tumor growth with respect to the = untreated=20 group was observed in the BP-70-, BP-150-, BP-300-, = BP-500-=20 and BP-800-treated groups. At day 200, the tumor = growth rates=20 were 100% (6/6), 80% (4/5), 80% (4/5), 50% (3/6), = 33.3% (2/6)=20 and 16.7% (1/6) in the untreated, BP-70, BP-150-, = BP-300-,=20 BP-500- and BP-800-treated groups, respectively=20 (p < 0.05). The mean tumor sizes = at day 89=20 were > 1000 mm³ in the control = group,=20 605.8 =B1 98.8 mm³ in the=20 BP-70-treated group,=20 504.4 =B1 38.9 mm³ in the=20 BP-150-treated group, = 415 =B1 130 mm³=20 in the BP-300-treated group,=20 365.8 =B1 116.7 mm³ in the=20 BP-500-treated group and 171.6 mm³ in = the=20 BP-800-treated group (Fig. 5a;=20 p < 0.05). Survival was = significantly=20 prolonged for nude mice in the BP-treated with respect = to the=20 control group (Fig. 5b;p < = ;0.001).=20 At day 200, survival rates were 0% (0/6), 20% (1/5), = 40%=20 (2/5), 50% (3/6), 66.7% (4/6) and 83.3% (5/6) in the=20 untreated, BP-70-, BP-150-, BP-300-, BP-500- and=20 BP-800-treated groups, respectively. After tumor = inoculation,=20 tumor masses (average=20 = volume =3D 41.84 =B1 4.45 mm³)=20 were developed on the backs of every mouse, but the = tumors of=20 mice with BP treatment shrank or completely regressed = to be=20 undetectable on day 150. There were no significant = differences=20 between the control and BP-treated groups with respect = to the=20 body weight of the animals (Fig. 5c). = Even with a=20 high BP dose of 800 mg/kg for 5 days of = serial=20 treatment, very low or no drug-related toxicities were = observed in the animals, as evaluated by histological = analyses=20 of various organs (data not shown). The human GBM = tumor=20 tissues with BP treatment in vivo also = displayed=20 decreases in Ki-67 expression, and increases in = cleaved=20 caspase 3 protein expression and tumor cell apoptosis, = relative to the control group in vivo, at day = 10 after=20 treatment (Fig. 5d).

Discussion

Go to:

Top of pageMaterials and = methodsResultsAcknowledgementsReferencesAbbreviations = used

n-Butylidenephthalide (BP) contributes to a = range of=20 biological activities, including diminution of angina, = platelet aggregation, proliferation, non-specific = spasmosis=20 (Ko = 1980; Teng=20 et al. 1987; Mimura = et al.=20 1995; Ko = et al.=20 1998; 2002). In this study, we demonstrated that = BP also=20 acts strongly against GBM tumors in vitro and = in=20 vivo. This anti-tumor effect is encouraging, as = very few=20 drugs currently show efficacy against malignant brain=20 tumors.

To explore what mechanism might account for the = effects of=20 BP on GBM tumor cells, the cell cycles were monitored. = Our=20 results are entirely consistent with an inhibition of = tumor=20 growth by BP due to cell-cycle arrest at the=20 G₀/G₁ phase and the induction of = apoptosis (Fig. 1). The = pattern=20 of increased and decreased protein expression during=20 cell-cycle arrest is best explained by the BP-mediated = regulation of gene expression involving the = cyclin/CDK/CKI=20 cell cycle regulatory system (Figs 2a and = b). The=20 p53 molecule is a negative regulator of cell division. = In=20 response to DNA damage, oncogenic activation of other = proteins=20 and other stresses, p53 levels rise and prevent cells = from=20 entering the S phase of the cell cycle (Reed = et al.=20 1994; Sherr=20 and Robert 1995). The p16 and p21 proteins are cdk = inhibitors that negatively regulate cdk or cyclin/cdk=20 complexes (Sherr=20 et al. 1995). The p16 protein, a = member of the=20 INK4 family, binds to cdk4 or cdk6 to block the kinase = activities at the mid-G₁ phase (Reed = et al.=20 1994). The p21 protein binds to the cyclin/cdk = complex,=20 resulting in the inhibition of the G₁ to S = phase=20 transition (Dobashi = et al.=20 2003) via the abrogation of Rb phosphorylation by = the=20 cyclin/cdk complex. Therefore, the decrease in = phosphorylated=20 Rb should be due to a BP-triggered expression of the = cdk=20 inhibitors, which decreases the activities of the = cyclin/cdk=20 complex. A precedent for this mechanism exists with=20 monoterpene, an essential oil of plants (like BP) that = is a=20 natural anti-cancer compound, which causes = G₁=20 arrest and leads to an increase in p21 expression (Bardon=20 et al. 2002). Another example is = aragusterol=20 A. This compound, isolated from marine sponges, is a = potent=20 anti-cancer marine steroid that causes G₁ = arrest by=20 down-regulation of Rb phosphorylation (Fukuoka = et al.=20 2000).

The immunoblotting results inferred the possible = mechanisms=20 of apoptosis induced by BP in GBM tumor cells. First, = the=20 p53-dependent apoptosis pathway was supposed because = BP=20 treatment could promote the phosphorylation of p53 in = the=20 DBTRG tumor cells and increase p53 expression, which = peaked at=20 12 h (Fig. 2a).=20 Phosphorylation at the N-terminal region of p53 = abolishes Mdm2=20 inhibition, which causes an increase in Bax = transcription (Fig. 2c=20 shows that Bax proteins were increased) and an = inhibition of=20 Bcl-2 transcription. Because of Bax/Bcl-2 imbalance, = Bax can=20 form an active homodimer to trigger cleavage of = procaspase 9=20 (Fig. 2c=20 shows that caspase 9 was increased) and sequentially = results=20 in procaspase 3 activation (Fig. 2d also = shows=20 that caspase 3 was increased) to proceed apoptosis = processes.=20 Second, the Fas=96FasL-induced apoptosis pathway may = also be=20 involved in BP-induced apoptosis. After BP treatment,=20 increasing expression of Fas was observed (Fig. 2d). = Fas is the=20 death receptor responsible for signaling from the cell = membrane, which triggers the activation of procaspase = 8 (Fig. 2d=20 shows that caspase 8 was increased) and subsequently, = promotes=20 procaspase 3 activation resulting in apoptosis. In = order to=20 examine these parameter cause related, different=20 pharmcological blockers had been approach. Briefly, we = pre-treated DBTRG cells with the Fas/FasL antagonist, = 341291=20 (Calbiochem), or the Bax-inhibiting peptide, 196810=20 (Calbiochem), then treated the cells with BP to = determine the=20 effects of these inhibitors. We found that neither = Fas/Fas L=20 nor Bax-inhibiting peptide could reverse BP-induced = growth=20 inhibition. However, BP-induced growth inhibition = effect might=20 be attenuated by the MGMT = (O⁶-methylguanine-DNA=20 methyltransferase) tranfection assay (data not shown). = MGMT is=20 a recognized DNA repair gene (Tano = et al.=20 1997). These results suggest that BP might be an=20 alkylating agent. It could directly injure tumor DNA = and cause=20 tumor cell arrest at the G₀/G₁ = phase in=20 order to repair the damage. Our study using blocking = agents=20 indicated that Fas/Fas L and Bax were not cause tumor=20 apoptosis molecules. The upper regulation of Fas and = Bax=20 protein was an association phenomenon.

RG2 cells have impaired p53 expression and = homozygous=20 deletions at the p16/Cdkn2a/Ink4a gene locus. = Therefore,=20 BP-induced apoptosis should be caused via a = p53-indepedent=20 pathway (Fas-induced apoptosis pathway) but not a = p53-depedent=20 pathway. However, although RG2 cells were impaired for = p53 and=20 p16, p27 and procaspase 9 expression induced by BP = were=20 increased in these cells. Thus, certain mechanisms of=20 p53-independent activation of procaspase 9 and p27 = induction=20 may be involved in BP-induced cell apoptosis and cell = growth=20 inhibition. The molecular profile shown in Fig. (2) = between=20 DBTRG8401 and DBTRG-05 is much similar to RG2. The RG2 = cells=20 had no p53 protein expression. It might be the = possibility of=20 different profile of caspase-9 and p27 activation = between=20 DBTRG and RG2 cells.

Transdermally-applied BP quickly permeates the = peripheral=20 circulation system without accumulating in the skin; = it is=20 then distributed into the lung, liver, bile, brain and = kidney.=20 Total radioactivity, however, is decreased due to its=20 excretion into the urine. In the case of intravenous=20 administration, 80% of the administered BP is excreted = into=20 the urine within 24 h, while only 5% is excreted = into the=20 feces. The metabolite in urine is a cysteine conjugate = (Sekiya=20 et al. 2000). In our experiment, = DBTRG-05MG=20 cells (2.5 =D7 10⁶) were injected = s.c. into=20 the backs of nude mice (five to six/group) and treated = with BP=20 on days 4, 5, 6, 7 and 8 after tumor cell = implantation.=20 Survival of nude mice was significantly prolonged in = the=20 BP-treated groups, relative to the control group (Fig. 5b;=20 p < 0.001). On day 200, the = survival rate=20 in the BP-800 mg/kg-treated group was 83.3% = (5/6).=20 Because BP is excreted into the urine within = 24 h, it is=20 interesting to note that five continuous doses showed=20 significant tumor inhibition.

In conclusion, BP has potent anti-cancer effects on = cell-cycle arrest and apoptosis. The present results = provide=20 new hope for chemotherapy of malignant brain cancer. = This may=20 lead to new therapeutic options in the treatment of = malignant=20 brain cancer, and an improved understanding of the=20 interactions between phytochemicals with the genes = that are=20 critical to the regulation of brain cancer = cells.

Acknowledgements

Go to:

Top of pageMaterials and = methodsResultsDiscussionAcknowledgements = <<ReferencesAbbreviations = used

This work was supported by grant = NSC94-2320-B303-001 from=20 National Science Council, Taiwan, Republic of = China.

References

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Footnotes

¹Nu-Man Tsai, Yi-Lin Chen = and=20 Chau-Chin Lee contributed equally to this = work.

²Shinn-Zong Lin and = Horng-Jyh Harn=20 are the corresponding authors.

³Re-use of this article = is=20 permitted in accordance with the Creative Commons = Deed,=20 Attribution 2.5, which does not permit commercial=20 exploitation.

Journal of = Neurochemistry
Online=20 Early

ASAngelica=20 sinensisAS-CA. sinen= sis=20 chloroform extractBCNUcarmustineBPn-butyliden= ephthalideCKICyclin=20 Kinase InhibitorDMSOdimethylsulfoxid= eMRImagnetic=20 resonance imagingPIpropidium=20 iodide

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Authors:

Nu-Man Tsai

Yi-Lin Chen

Chau-Chin Lee

Po-Chen Lin

Yeung-Leung Cheng

Wen-Liang Chang

Shinn-Zong Lin

Horng-Jyh Harn

Keywords:

Angelica = sinensis

apoptosis

glioblastoma = multiformis

n-butylidenephthalide

Received April 13, 2006; revised manuscript = received July=20 18, 2006; accepted July 25, = 2006.

Accepted article = online 21=20 Aug 2006
Published article online 20 Sep = 2006

Affiliations

*Department of = Applied Life=20 Science, Asia University, Taichung, = Taiwan
=86Neuro-Medical Scientific Center, = =87Department of Radiology, =A7Institute=20 of Medical Sciences, =86=86Department of=20 Pathology and =87=87Department of = Emergency=20 Medicine, Buddhist Tzu Chi General Hospital, Hualien,=20 Taiwan
=B6Division of Thoracic = Surgery,=20 Tri-Service General Hospital, National Defense Medical = Center,=20 Taipei, Taiwan
**School of = Pharmacy,=20 National Defense Medical Center,Taipei,=20 Taiwan

Correspondence

Address correspondence and = reprint=20 requests to Dr Horng-Jyh Harn, Neuro-Medical = Scientific=20 Center, Department of Pathology, Tzu-Chi General = Hospital,=20 707, Section 3, Chung-Yang Road, 970 Hualien, Taiwan. = E-mail:=20 duke@tzuchi.com.= tw

¹Nu-Man Tsai, Yi-Lin Chen = and=20 Chau-Chin Lee contributed equally to this=20 work.

²Shinn-Zong Lin and = Horng-Jyh Harn=20 are the corresponding = authors.

³Re-use of this article = is=20 permitted in accordance with the Creative Commons = Deed,=20 Attribution 2.5, which does not permit commercial=20 exploitation.

Abbreviations

Image Previews

[Full Size]

Fig. 1 BP-induced=20 cell-cycle arrest and apoptosis in GBM cells. = DBTRG-05MG and=20 RG2 cells underwent apo...

[Full Size]

Fig. 2 The=20 expression and activation of apoptosis-associated = molecules in=20 GBM cells treated with BP. = W...

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Fig. 3 BP=20 inhibition of tumor growth with improved survival rate = in a=20 syngenic rat GBM model. RG2 = cell...

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Fig. 4 BP=20 reduction in tumor volume in a syngenic rat GBM tumor = in=20 situ model. RG2 cells = (5 =D7 10⁴) were=20 i...

[Full Size]

Fig. 5 = BP=20 suppression of xenograft tumor growth of human GBM in = nude=20 mice. (a) DBTRG-05MG=20 cells = (2.5 =D7 10...

[Full = Size]

Table 1 The=20 IC₅₀s of BP in different = tumors

To cite = this=20 article
Tsai, Nu-Man, = Chen,=20 Yi-Lin, Lee, Chau-Chin, Lin, Po-Chen, Cheng, = Yeung-Leung,=20 Chang, Wen-Liang, Lin, Shinn-Zong & Harn, = Horng-Jyh=20 (2006)
The natural compound = n-butylidenephthalide=20 derived from Angelica sinensis inhibits = malignant brain=20 tumor growth in vitro and in=20 vivo³.
Journal of=20 = Neurochemistry 0 (0), ???-???.
doi:=20 = 10.1111/
j.1471-4159.2006.04151.x

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+=3D "&orderBeanReset=3D" + aOrderBeanReset;=0A= }=0A= =0A= var link =3D "/servlet/linkout?type=3Drightslink&url=3D" + = escape(location);=0A= var winprops =3D = "location=3Dno,toolbar=3Dno,directories=3Dno,status=3Dno,menubar=3Dno,scr= ollbars=3Dyes,resizable=3Dyes,width=3D650,height=3D550";=0A= PopUp =3D window.open(link, 'Rightslink', winprops);=0A= }=0A= =0A= =0A= /* Function to load images.=0A= * note: make this empty method so that=0A= * 1) mininize code changes; 2) for future integration; 3) easier to = take the feature back=0A= */=0A= function MM_preloadImages() {=0A= }=0A= =0A= function MM_swapImgRestore() {=0A= }=0A= =0A= function MM_findObj(n, d) {=0A= }=0A= =0A= function MM_swapImage() {=0A= }=0A= =0A= function menuMouseOver(type, actualType) {=0A= = document.getElementById('MenuItem_'+type).className=3D'MenuItemOver_'+(ty= pe =3D=3D actualType);=0A= }=0A= function menuMouseOut(type, actualType) {=0A= = document.getElementById('MenuItem_'+type).className=3D'MenuItem_'+(type = =3D=3D actualType);=0A= }=0A= function menuClick(type) {=0A= document.getElementById('MenuLink_'+type).click();=0A= }=0A= =0A= // browser detection=0A= function browserCheck() {=0A= this.ns4 =3D (document.layers)? true:false;=0A= this.ie =3D (document.all&&(!window.opera))? true:false;=0A= this.dom =3D (document.getElementById)? true:false;=0A= this.ns6 =3D (window.sidebar)? true:false;=0A= this.moz =3D = (window.sidebar||navigator.userAgent.indexOf('Gecko')!=3D-1)? true:false;=0A= this.opera =3D (window.opera)? true:false;=0A= this.mac =3D (navigator.userAgent.indexOf('Mac')!=3D-1)? true:false;=0A= }=0A= browser =3D new browserCheck();=0A= var Obj;=0A= =0A= // multibrowser get object by id, in NN4 you cannot access all object = (test it first)=0A= function getObjectByName(nameOfObject){=0A= Obj =3D null;=0A= if (browser.ie) Obj =3D document.all[nameOfObject]=0A= else if (browser.dom) Obj =3D findDOMObject(nameOfObject)=0A= else if (browser.ns4) findLayer(window,nameOfObject);=0A= return (!Obj || ( browser.ns4 && Obj =3D=3D window ) ) ? "Object not = found" : Obj;=0A= }=0A= // helper function=0A= function findDOMObject(nameOfObject) {=0A= for (var i =3D 0; i < document.images.length; i++) {=0A= if (document.images[i].name=3D=3DnameOfObject) return = document.images[i]=0A= }=0A= return document.getElementById(nameOfObject)=0A= }=0A= // helper function=0A= function findLayer(node,nameOfObject) {=0A= if ( node.name =3D=3D nameOfObject ) Obj =3D node;=0A= for ( var counter =3D 0; counter < node.document.images.length; = counter++ ) {=0A= if (node.document.images[counter].name=3D=3DnameOfObject) Obj =3D = node.document.images[counter];=0A= }=0A= for ( var i =3D 0; i < node.document.layers.length; i++ ) {=0A= findLayer(node.document.layers[i],nameOfObject);=0A= }=0A= }=0A= =0A= function toggleVisibility(id) {=0A= obj =3D getObjectByName(id);=0A= if (obj) {=0A= v =3D obj.style.display;=0A= obj.style.display =3D (v =3D=3D 'none') ? 'block' : 'none';=0A= i =3D getObjectByName('hiddenImage');=0A= if(i) i.src =3D '/page/imageHack.jsp?object=3D' + id + = '&display=3D' + obj.style.display + '&dummy=3D' + (new Date()).getTime();=0A= }=0A= }=0A= =0A= /** CSS class support */=0A= /** return true if removed, false if not found */=0A= function removeClass(el, className) {=0A= if (!(el && el.className)) return false;=0A= var cls =3D el.className.split(" ");=0A= var ar =3D new Array();=0A= for (var i =3D cls.length; i > 0;) if (cls[--i] !=3D className) = ar[ar.length] =3D cls[i];=0A= el.className =3D ar.join(" ");=0A= return cls.length !=3D ar.length;=0A= }=0A= /** return true if element has that class */=0A= function hasClass(el, className) {=0A= if (!(el && el.className)) return false;=0A= var cls =3D el.className.split(" ");=0A= for (var i =3D cls.length; i > 0;) if (cls[--i] =3D=3D className) = return true;=0A= return false;=0A= }=0A= /** add class to element, if element already had that class - does = nothink */=0A= function addClass(el, className) {=0A= if (!el || hasClass(el, className)) return;=0A= if (el.className) el.className +=3D " " + className;=0A= else el.className =3D className;=0A= }=0A= /** Add CSS rule as last rule in last stylesheet - override all previous = definitions=0A= removing is harder - not implemented, use it with caution */=0A= function addCssRule(selector, rule) {=0A= if (document.styleSheets) {=0A= var css =3D document.styleSheets[document.styleSheets.length-1]; = // get last css=0A= if (css.addRule) css.addRule(selector, rule); // IE=0A= else if (css.insertRule) css.insertRule(selector + "{" + rule + = "}", css.cssRules.length); // W3C=0A= }=0A= }=0A= /** get CSS style of the document */=0A= function getCssText() {=0A= var imports =3D "";=0A= var cssText =3D "\n";=0A= var styleSheets =3D document.styleSheets;=0A= for (var i =3D 0; i < styleSheets.length; i++) {=0A= var css =3D styleSheets.item(i);=0A= var mediaText =3D typeof css.media =3D=3D "string" ? css.media : = css.media.mediaText;=0A= if (!mediaText || mediaText.indexOf("all") !=3D -1 || = mediaText.indexOf("screen") !=3D -1) {=0A= if (css.imports) {=0A= // IE imports=0A= for (var j =3D 0; j < css.imports.length; j++) {=0A= var href =3D css.imports[j].href;=0A= var lastQuote =3D href.lastIndexOf('"');=0A= if (lastQuote !=3D -1) {=0A= // href contains media - "url" media=0A= mediaText =3D href.substr(lastQuote + 1);=0A= if (mediaText.indexOf("all") =3D=3D -1 && = mediaText.indexOf("screen") =3D=3D -1) continue;=0A= href =3D href.substr(1, lastQuote - 1);=0A= }=0A= imports +=3D '@import url("' + href + '");\n';=0A= }=0A= }=0A= if (css.rules) {=0A= // IE rules=0A= for (var j =3D 0; j < css.rules.length; j++) {=0A= // How to filter media rules ???=0A= cssText +=3D css.rules[j].selectorText + " { " + = css.rules[j].style.cssText + " }\n";=0A= }=0A= } else if (css.cssRules) {=0A= // W3C compliant browser=0A= for (var j =3D 0; j < css.cssRules.length; j++) {=0A= var rule =3D css.cssRules.item(j);=0A= if (rule.type =3D=3D rule.IMPORT_RULE || rule.type =3D=3D = rule.MEDIA_RULE) {=0A= mediaText =3D rule.media.mediaText;=0A= if (mediaText && mediaText.indexOf("all") =3D=3D -1 && = mediaText.indexOf("screen") =3D=3D -1) continue;=0A= if (rule.type =3D=3D rule.IMPORT_RULE) imports +=3D = rule.cssText;=0A= else cssText +=3D rule.cssText + "\n";=0A= } else {=0A= cssText +=3D rule.cssText + "\n";=0A= }=0A= }=0A= }=0A= }=0A= }=0A= return imports + cssText;=0A= }=0A= =0A= /** DOM Event support */=0A= /** addEventListener method may already exist, it's not trivial = implement it and not break it, use other mame 'addListener' is simple */=0A= function addListener(el, evname, func) {=0A= if (el.attachEvent) el.attachEvent("on" + evname, func);=0A= else if(el.addEventListener) el.addEventListener(evname, func, true);=0A= }=0A= /** removeEventListener */=0A= function removeListener(el, evname, func) {=0A= if (el.detachEvent) el.detachEvent("on" + evname, func);=0A= else if (el.removeEventListener) el.removeEventListener(evname, func, = true);=0A= }=0A= =0A= /** Server session access */=0A= /** helper method to access session object */=0A= function _getSession() {=0A= return window.session ? window.session : (window.session =3D new = Array());=0A= }=0A= /** send request to server to preserve some value between different = requests, value is accessible using getSessionAttribute */=0A= function setSessionAttribute(name, value) {=0A= /* just to test callback */=0A= function test(doc) {=0A= // alert("Server returned: " + doc.body.innerHTML);=0A= }=0A= = sendServerMsg("/action/sessionAccess?action=3DsetJavaScriptAttribute&name= =3D" + name + "&value=3D" + value, test);=0A= _getSession()[name] =3D value;=0A= }=0A= /** see setSessionAttribute */=0A= function getSessionAttribute(name) {=0A= return _getSession()[name];=0A= }=0A= =0A= /** send request - url - to the server, response is accessible in = document passed to callback function=0A= DO NOT Send anything before page is loaded!=0A= */=0A= function sendServerMsg(url, callBack) {=0A= /* called when frame is loaded */=0A= function _receiveServerResponse(ev) {=0A= var iFrame =3D window.event ? window.event.srcElement : = ev.currentTarget; // IE : W3C=0A= var doc =3D iFrame.contentDocument ? iFrame.contentDocument : = document.frames(iFrame.id).document; // W3C : IE=0A= if (iFrame.callBack) iFrame.callBack(doc);=0A= document.body.removeChild(document.getElementById(iFrame.id));=0A= }=0A= =0A= var iFrame =3D document.createElement('iframe');=0A= addListener(iFrame, "load", _receiveServerResponse);=0A= iFrame.setAttribute("id", "msgFrame" + new Date().getTime()); // = so IE can retrive document=0A= iFrame.setAttribute("src", url);=0A= iFrame.setAttribute("style", "display:none");=0A= if (callBack) iFrame.callBack =3D callBack;=0A= document.body.appendChild(iFrame);=0A= }=0A= =0A= /**=0A= copy innerHTML of src element to innerHTML of target element=0A= */=0A= function copyInnerHtml(src, target) {=0A= var domSupported =3D document.getElementById ? true : false;=0A= var se =3D (domSupported && typeof src =3D=3D "string") ? = document.getElementById(src) : src;=0A= var te =3D (domSupported && typeof target =3D=3D "string") ? = document.getElementById(target) : target;=0A= if (se.innerHTML && te.innerHTML) te.innerHTML =3D se.innerHTML;=0A= }=0A= /**=0A= filter select options, options that do not contain filter text are = removed=0A= in first use, original select options are backed up as attribute = optionsCopy=0A= */=0A= function filterSelect(selectId, value) {=0A= var select;=0A= if (document.getElementById && (select =3D = document.getElementById(selectId)) && select.options) {=0A= if (!select.optionsCopy) {=0A= select.optionsCopy =3D new Array();=0A= select.optionTexts =3D new Array();=0A= for (var i=3D0; i'+document.title+'');=0A= tmp.writeln(el.innerHTML);=0A= tmp.writeln('');=0A= tmp.close();=0A= }=0A= if (anchor) {=0A= var target =3D (tmp.getElementById) ? = tmp.getElementById(anchor) :=0A= (tmp.all) ? tmp.all[anchor] : false;=0A= if (target) {=0A= if (target.scrollIntoView) target.scrollIntoView();=0A= else if (window.scroll && target.offsetTop) = el.popupWindow.scroll(0, target.offsetTop);=0A= }=0A= }=0A= }=0A= }=0A= =0A= /** marks all check box */=0A= function markAllCheckboxes(aForm, aNamePrefix, aChecked)=0A= {=0A= var elmts =3D aForm.elements;=0A= for (var i=3D0; i 0);=0A= if (addElmt)=0A= url +=3D "&" + elmts[i].name + "=3D" + elmts[i].value;=0A= }=0A= window.location =3D url;=0A= }=0A= }=0A= =0A= /**=0A= * @param aAuthorArray array stores all authors=0A= * @param aKeywordArray array stores all keywords=0A= */=0A= function getAuthorsAndKeywords(aForm, aAuthorArray, aKeywordArray, = aForceAdd)=0A= {=0A= setCheckWhenDefine(aForm.sauthusercbx, true);=0A= setCheckWhenDefine(aForm.keyusercbx, true);=0A= =0A= var elements =3D aForm.elements;=0A= var cntAuthor =3D -1;=0A= var cntKeyword =3D -1;=0A= for (var i=3D0; i0) || (aKeywords.length>0))=0A= {=0A= pmurl =3D = "http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?dopt=3DDocSum&cmd=3DSearch= &db=3DPubMed&term=3D";=0A= =0A= for (var i=3D0; i0) && (aKeywords.length>0)) {=0A= pmurl +=3D "+OR+";=0A= }=0A= =0A= for (var i=3D0; i0) || (aKeywords.length>0))=0A= {=0A= adsurl =3D=0A= "http://adsabs.harvard.edu/cgi-bin/nph-abs_connect?db_key=3DAST" +=0A= "&sort=3DSCORE&ttl_syn=3DYES&version=3D1" +=0A= "&txt_syn=3DYES&txt_logic=3DAND" +=0A= "&aut_syn=3DYES&aut_logic=3DAND";=0A= =0A= if (aAuthors.length > 0) {=0A= adsurl +=3D "&author=3D";=0A= for (var i=3D0; i 0) {=0A= adsurl +=3D "&text=3D";=0A= for (var i=3D0; i0) || (aKeywords.length>0))=0A= {=0A= var array =3D [aAuthors, aKeywords];=0A= for (var i=3D0; i0&&parent.frames.length) {=0A= d=3Dparent.frames[n.substring(p+1)].document; n=3Dn.substring(0,p);}=0A= if(!(x=3Dd[n])&&d.all) x=3Dd.all[n]; for = (i=3D0;!x&&i 0) ? "&" : "?");=0A= x +=3D (aButton + ".x=3D1");=0A= aForm.action =3D x;=0A= }=0A= }=0A= else=0A= {=0A= var sltd =3D x.options[x.selectedIndex];=0A= var pfxJSBTN =3D "jsbtn:";=0A= if (sltd.value.indexOf(pfxJSBTN) =3D=3D -1)=0A= {=0A= var value =3D (pfxJSBTN + aButton + ";" + sltd.value);=0A= //x.options[x.selectedIndex] =3D new Option(sltd.text, value, true, = true);=0A= x.options[x.selectedIndex].value =3D value;=0A= }=0A= }=0A= aForm.submit();=0A= }=0A= }=0A= =0A= =0A= function onChangeClick(aSelect, button)=0A= {=0A= var frm =3D aSelect.form;=0A= submitFormWithButtonClicked(frm, button);=0A= }=0A= =0A= function onChangeClick2(aSelect, button)=0A= {=0A= var frm =3D document.forms[0];=0A= var x =3D frm.action;=0A= x +=3D "?"+button+".x=3D1";=0A= frm.action =3D x;=0A= frm.submit();=0A= }=0A= =0A= function submitArticles(aForm, action) {=0A= submitMultiArticles(aForm, action, true);=0A= }=0A= =0A= /* Function to show next search page */=0A= function searchNextResultPage(aForm, aOffset)=0A= {=0A= var x =3D eval(aForm.startPage.value) + aOffset;=0A= aForm.startPage.value =3D x;=0A= aForm.submit();=0A= }=0A= =0A= /* Function to show help page */=0A= function popupHelp(url)=0A= {=0A= popupHelp550(url);=0A= }=0A= =0A= /* Function to open up a new browser window, without a navigation bar */=0A= function newWindow(url)=0A= {=0A= var new_window;=0A= var windowProperties;=0A= windowProperties =3D = "width=3D750,height=3D700,top=3D30,left=3D230,toolbar=3D0,menubar=3D0,res= izable=3D1,scrollbars=3Dyes";=0A= if(new_window=3D=3Dnull) {=0A= new_window =3D window.open(url,null,windowProperties);=0A= } else {=0A= new_window.document.replace(url);=0A= }=0A= }=0A= =0A= /* Function to open up a new browser window, without a navigation bar */=0A= function newInstitutionWindow(url)=0A= {=0A= var new_window;=0A= var windowProperties;=0A= windowProperties =3D = "width=3D750,height=3D700,top=3D30,left=3D230,toolbar=3D1,menubar=3D1,res= izable=3D1,scrollbars=3Dyes";=0A= if(new_window=3D=3Dnull) {=0A= new_window =3D window.open(url,null,windowProperties);=0A= } else {=0A= new_window.document.replace(url);=0A= }=0A= }=0A= =0A= =0A= /**=0A= * Function to close the current window=0A= */=0A= function close_window() {=0A= window.close();=0A= }=0A= =0A= /**=0A= * get all the form elements and check if the "dbname"=0A= * maps to specifid value.=0A= */=0A= function checkDbName(aForm, aValue)=0A= {=0A= var elements =3D aForm.elements;=0A= for (var i=3D0; i0)) {=0A= alert("Sorry, email alert for journals with \n" +=0A= "full access rights is not available.");=0A= return;=0A= }=0A= =0A= var elmts =3D aForm.elements;=0A= for (var i=3D0; i