Glioblastoma stem cells resistant to = temozolomide-induced autophagy

From: Subject: Glioblastoma stem cells resistant to temozolomide-induced autophagy Date: Sat, 1 Aug 2009 18:18:43 +0200 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0000_01CA12D4.80C03F90" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.5579 This is a multi-part message in MIME format. ------=_NextPart_000_0000_01CA12D4.80C03F90 Content-Type: text/html; charset="Windows-1252" Content-Transfer-Encoding: quoted-printable Content-Location: http://www.cmj.org/Periodical/paperlist.asp?id=LW200964460632407790&linkintype=pubmed Glioblastoma stem cells resistant to = temozolomide-induced autophagy

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FU Jun State Key Laboratory for = Cancer Research=20 in Southern China and Department of = Neurosurgery/Neuro-oncology,=20 Cancer Center, Sun Yat-sen University, Guangzhou, Guangdong = 510060,=20 China; LIU Zhi-gang State Key = Laboratory=20 for Cancer Research in Southern China and Department of=20 Neurosurgery/Neuro-oncology, Cancer Center, Sun Yat-sen = University,=20 Guangzhou, Guangdong 510060, China; LIU=20 Xiao-mei State Key Laboratory for Cancer Research in = Southern=20 China and Department of Neurosurgery/Neuro-oncology, Cancer = Center,=20 Sun Yat-sen University, Guangzhou, Guangdong 510060, China; = CHEN Fu-rong State Key Laboratory for = Cancer=20 Research in Southern China and Department of=20 Neurosurgery/Neuro-oncology, Cancer Center, Sun Yat-sen = University,=20 Guangzhou, Guangdong 510060, China; SHI=20 Hong-liu State Key Laboratory for Cancer Research in = Southern=20 China and Department of Neurosurgery/Neuro-oncology, Cancer = Center,=20 Sun Yat-sen University, Guangzhou, Guangdong 510060, China; = PANG Jesse Chung-sean Department of = Anatomical=20 & Cellular Pathology, Chinese University of Hong Kong, = Shatin,=20 N.T., Hong Kong, China; NG = Ho-keung=20 Department of Anatomical & Cellular Pathology, Chinese=20 University of Hong Kong, Shatin, N.T., Hong Kong, China; = CHEN Zhong-ping State Key Laboratory = for Cancer=20 Research in Southern China and Department of=20 Neurosurgery/Neuro-oncology, Cancer Center, Sun Yat-sen = University,=20 Guangzhou, Guangdong 510060, China

Correspondence = to: CHEN=20 Zhong-ping Department of = Neurosurgery/Neuro-oncology,=20 Cancer Center, Sun Yat-sen University, Guangzhou, Guangdong = 510060,=20 China (Tel:86-20-87343310. = Email:chenzp57@mail.sysu.edu.cn=20 )This study was supported by : National = Natural=20 Science Foundation of China(No. 30772551) Keywords: = glioblastoma=A1=A4neoplastic=20 stem cells=A1=A4chemoresistance=A1=A4= temozolomideAbstract:

Background Recent studies have = demonstrated=20 the existence of a small fraction of cells with features of=20 primitive neural progenitor cells and tumor-initiating = function in=20 brain tumors. These cells might represent primary = therapeutic target=20 for complete eradication of the tumors. This study aimed to=20 determine the resistant phenotype of glioblastoma stem cells = (GSCs)=20 to temozolomide (TMZ) and to explore the possible molecular=20 mechanisms underlying TMZ resistance.

Methods Freshly resected = glioblastoma=20 specimen was collected and magnetic isolation of GSCs was = carried=20 out using the Miltenyi Biotec CD133 Cell Isolation kit. The=20 cytotoxic effect of TMZ on CD133⁺and = CD133^=A8C=20 glioblastoma cells was determined by using the=20 3-(4,5-dimethylthiazol-2-yl)-2,5-diphenyltetrazolium bromide = (MTT)=20 assay. Autophagy-related proteins (Beclin-1, LC3 and Atg5) = and=20 cleaved caspase-3 (p17) were analyzed by Western blotting.=20 Immunofluorescent staining was used to detect Atg5, glial = fibrillary=20 acidic protein (GFAP) and CD133 expression in glioblastoma = cells.=20 Statistical analysis was carried out using SPSS 10.0 = software. For=20 all tests, the level of statistical significance was set at = P=20 <0.05.

Results CD133⁺ = glioblastoma cells=20 exhibited neurosphere-like growth in vitro and high=20 expression of CD133 stem cell marker. The growth-inhibiting = rate in=20 CD133^=A8C glioblastoma cells treated with 5 or 50 = =B5mol/L=20 TMZ was significantly higher than that in CD133⁺=20 glioblastoma cells ((14.36=A1=C03.75)% vs (2.54=A1=C01.36)% = or=20 (25.95=A1=C05.25)% vs (2.72=A1=C01.84)%, respectively, = P <0.05).=20 Atg5, LC3-II and Beclin-1 levels were significantly lower in = CD133⁺ glioblastoma cells than those in = autologous=20 CD133^=A8C cells after TMZ treatment (P = <0.05).=20 Caspase-3 was mildly activated only in CD133^=A8C=20 glioblastoma cells after exposure to TMZ (P = <0.05).=20 Immunofluorescent staining revealed elevated expression of = Atg5 in=20 GFAP⁺ cells following TMZ treatment.

Conclusions The GSCs display = strong=20 capability of tumor=A1=E4s resistance to TMZ. This = resistance is=20 probably contributed by the CD133⁺ cells with=20 down-regulation of autophagy-related proteins. Future = treatment=20 should target this small population of cancer stem cells in = tumors=20 to improve survival of patients.

2009;122(11):1255-1259

Chinese=20 Medical Journal, 2009, Vol. 122 No. 11 : = 1255-1259

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Glioblastoma stem cells resistant to = temozolomide-induced autophagy

FU Jun, LIU Zhi-gang, LIU = Xiao-mei, CHEN=20 Fu-rong, SHI Hong-liu, PANG Jesse Chung-sean, NG Ho-keung , = CHEN=20 Zhong-ping

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Despite progress in study on the = molecular aspects=20 of malignant gliomas, the prognosis of these brain tumors = continues=20 to be dismal. The median survival of patients with = glioblastoma=20 multiforme (GBM), the most aggressive glioma in adults, has = remained=20 from 9 to 12 months for decades.¹ Temozolomide = (TMZ) was=20 the first new chemotherapeutic agent approved for treatment = of=20 high-grade malignant gliomas in more than 20 years. Despite = the fact=20 that TMZ contributes significant therapeutic benefits in GBM = patients, the addition of TMZ to radiotherapy only resulted = in a=20 slightly longer median survival time in newly diagnosed GBM=20 patients, 14.6 versus 12.1 months, and the best 5-year = survival of=20 9.8%.² Recent studies propose the existence of a = small=20 fraction of cells with features of primitive neural = progenitor cells=20 and tumor-initiating function in brain tumors. These cells, = known as=20 cancer stem cells (CSCs) with stem-like properties, = constitute a=20 reservoir of self-sustaining cells with the exclusive = ability to=20 self-renew and to cause the heterogeneous lineages of cancer = cells=20 that comprise the tumor.³

Activation of cell death program has been = shown to=20 be responsible for chemotherapy-induced cytotoxicity in = tumor cells,=20 while alterations in the death machinery have been related = to=20 chemoresistance in gliomas.⁴ However, = conventional=20 approaches targeting the overall population of tumor cells = may well=20 spare CSCs owing to their idiosyncratic = properties.⁵=20 Several studies indicated that altered expression of=20 apoptosis-related proteins and DNA repair protein=20 O6-methylguanine-DNA-methyltransferase (MGMT) may render=20 glioblastoma stem cells (GSCs) strongly resistant to = apoptosis=20 induced by various therapeutic drugs including = TMZ.^5,6=20 Autophagy, also know as programmed cell death type II, = represents an=20 alternative tumor-suppressing mechanism to overcome, at = least=20 partly, the dramatic resistance of many cancers to = proapoptotic=20 chemotherapy.⁷ Autophagy, rather than apoptosis, = has been=20 associated with TMZ-induced cytotoxicity in glioma=20 cells.⁸ Does this machinery also work on GSCs? = This study=20 aimed to explore autophagy-related molecular mechanisms in = GSCs in=20 response to TMZ. Our finding suggests that down-regulation = of=20 autophagy proteins might be important mechanisms for GSCs to = evade=20 TMZ-induced cytotoxicity.

METHODS

Reagents
TMZ was kindly = provided by=20 Tasly Pharmaceutical Co. Ltd (China). The following = antibodies were=20 used: glial fibrillary acidic protein (GFAP), = =A6=C2-tubulin, cleaved=20 caspase-3 (p17) (Santa Cruz, USA); CD133, Atg5, Beclin-1, = LC3B (Cell=20 Signaling, USA); fluorescein isothiocyanate (FITC)- or=20 Cy3-conjugated secondary antibodies (Jackson Laboratory, = USA).

Brain tumors specimens, cell sorting = and culture=20
Primary glioblastoma, named GBM-04, was freshly = obtained=20 from the operating room following approved protocols and = verified by=20 pathologists. CD133⁺ glioblastoma cells were = isolated and=20 cultured as described previously by Singh et al.³ = Briefly, specimens or xenografts were chopped manually, = dissociated=20 with collagenase/dispase, and cultured in stem cell medium = (DMEM/F12=20 medium supplemented with 2% B27 minus vitamin A (Invitrogen, = USA),=20 25 ng/ml of epidermal growth factor (EGF) and 20 ng/ml of = basic=20 fibroblast growth factor (bFGF). Within 3 days of primary = culture,=20 cells were centrifuged, triturated with a fire-narrowed = Pasteur=20 pipette, and resuspended in phosphate buffered saline (PBS) = with=20 0.5% bovine serum albumin (BSA) and 2 mmol/L=20 ethylenediaminetetraacetic acid (EDTA). Magnetic isolation = of GSCs=20 was carried out using the Miltenyi Biotec CD133 Cell = Isolation kit=20 (Miltenyi Biotec GmbH, Germany). CD133⁺ and=20 CD133^=A8C sorted cell populations were = resuspended in stem=20 cell medium.

Cytotoxicity assay
The = cytotoxic effect=20 of drugs on glioma cells was determined by using a=20 3-(4,5-dimethylthiazol-2-yl)- 2,5-diphenyltetrazolium = bromide (MTT)=20 assay as described previously.⁹ Briefly,=20 1=A1=C110³ cells were plated in 100 =A6=CCl of = medium in 96-well=20 microtiter plates pre-coated with poly-ornithine and = incubated for=20 24 hours. Drugs were added and the cells incubated for a = further 72=20 hours, dimethyl sulfoxide was used as solvent control. MTT = solution=20 was added to each well and incubated at 37=A1=E3C for 2 = hours. The=20 reaction was stopped by adding dimethyl sulfoxide. The = amount of MTT=20 formazan product was determined by measuring absorbance with = a=20 microplate reader at a test wavelength of 570 nm and a = reference=20 wavelength of 630 nm. Optical density (OD) value was used to = express=20 the absorbance of the dual wavelengths. The = growth-inhibiting rate=20 of tumor cells equals (1-(the OD values of treated = groups/the OD=20 values of control group))=A1=C1100%. Data were the mean of = triplicate=20 experiments.

Immunofluorescence staining and = microscopy=20
Cells were cultured on cover slips pre-coated with=20 poly-lysine. After various treatments, cells were fixed in = 4%=20 paraformaldehyde, permeated with 0.25% Triton X-100, blocked = with 3%=20 normal goat serum, stained with the first antibody = overnight, and=20 labeled with a goat anti-mouse or goat anti-rabbit IgG = conjugated=20 with fluorescein isothiocyanate (FITC) or Cy3. Cells were=20 counterstained with anti-fade sealant containing=20 4=A1=E46-diamidino-2-phenylindole (DAPI) (Vectashield, USA) = and examined=20 under fluorescence microscope BX61 (Olympus, Japan). = Pictures were=20 captured with DP71 CCD digital camera (Olympus).

Western blotting
Cell lysates = were=20 prepared with cell lysis buffer (Cell signaling, USA). After = sonication, centrifugation, and protein assay (Pierce = protein assay=20 kit, USA), 50 =A6=CCg protein and an equal volume of 2 = =A1=C1 sample buffer=20 (62.5 mmol/L Tris-HCl pH6.8, 2% (w/v) sodium dodecyl = sulfate, 10%=20 glycerol, 50 mmol/L dithiothreitol, 0.01% (w/v) bromophenol = blue)=20 were heated at 94=BAC for 5 minutes. Proteins were separated = on a 10%=20 sodium dodecyl sulfate-polyacrylamide gel and transblotted = onto a=20 polyvinylidene difluoride (PVDF) transfer membrane (Bio-Rad, = USA).=20 The blot was blocked in PBS containing 0.1% Tween-20 and 5% = skim=20 milk at 37=BAC for 1 hour. The membrane was then incubated = in primary=20 antibody (1:200) at 4=BAC overnight, followed by treatment = with=20 secondary antibody conjugated with horseradish peroxidase = (1:1000).=20 Proteins were visualized using the ECL system (Amersham = Biosciences,=20 USA) and quantified using the Image J software version 1.37 = from=20 National Institute of Health (MA, USA). Data were the mean = of=20 triplicate experiments.

Statistical analysis =
Statistical=20 evaluations were carried out using SPSS 10.0 software (SPSS = Inc,=20 USA). For all tests, the level of statistical significance = was set=20 at P <0.05. The experimental data were expressed = as mean=20 =A1=C0 standard deviation (SD). Unless otherwise specified, = Student=A1=E4s=20 t test was used.

RESULTS

Characterization of GSCs
As = shown in=20 Figure 1A, HE staining validated GBM04 primary tumor to be=20 glioblastoma multiforme. After CD133 magnetic sorting, = sorted GSCs=20 were cultured in stem cell medium. Neurosphere-like = gliomaspheres=20 appeared after 3-day culture (Figure 1B). GSCs can also grow = as an=20 adherent monolayer in poly-ornithine-coated plate (Figure = 1C).=20 Immunofluorescence staining showed highly expressed CD133 in = GSCs=20 (Figure 1D).

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Figure 1. Characterization = of=20 CD133⁺ GSCs. A: HE staining of GBM04 = primary=20 tumor (Original magnification =A1=C1400). B: = After sorting,=20 CD133⁺ GSCs grow as neurosphere-like = gliomaspheres=20 in stem cell medium (Original magnification =A1=C140). = C:=20 CD133⁺ GSCs grow adherent to = poly-lysine-coated=20 bottom (Original magnification =A1=C1200). D: = Sorted GSCs=20 exhibit highly expressed CD133 (red), DAPI staining = (blue) was=20 performed to identify cells (Original magnification=20 =A1=C1200).

Cytotoxic effects of TMZ on glioma = cells=20
The cytotoxic effects of TMZ on CD133⁺ = and=20 CD133^=A8Cglioma cells are shown in Table 1. The=20 growth-inhibiting rate of CD133^=A8C cells after = TMZ (5 or=20 50 =B5mol/L) treatment was higher than that of = CD133⁺ cells=20 (P <0.05).

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Table 1. Cytotoxic effects = of=20 temozolomide on glioma=20 cells

Expression of autophagy-related = proteins=20 following TMZ treatment
The expressions of = autophagy-related=20 proteins Beclin-1, Atg5 and LC3 were identified by Western = blotting=20 in groups treated with/without TMZ (50 =B5mol/L, 72 hours). = The=20 results are shown in Figure 2, and the relative OD values = determined=20 by Image J software are displayed in Table 2. TMZ treatment = caused=20 increased Beclin-1, Atg5, LC3-II and cleaved caspase-3 = levels in=20 CD133^=A8C cells, but not in CD133⁺ = cells=20 (P <0.05). There were no statistical differences = of=20 Beclin-1, Atg5, LC3-II and cleaved caspase-3 expression = found=20 between the control and TMZ-treated group in = CD133⁺ cells=20 (P >0.05). All results suggest CD133⁺ = cells are=20 less responsive to TMZ-induced autophagy.

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Figure 2. Western blotting = analysis=20 for expression of autophagy-related proteins following = TMZ=20 treatment. The pictures are scanned and analyzed by = Image J=20 software. =A6=C2-tubulin was used as a loading=20 control.

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Table 2. Quantification of = autophagy-related proteins following TMZ=20 treatment

Expression of Atg5 in glioblastoma=20 cells
As shown in Figure 3, expression of Atg5 was = detected=20 by immunofluorescence staining. Enormous punctate Atg5 = localization=20 emerged after TMZ treatment (50 =B5mol/L, 72 hours). = GFAP⁺=20 glioblastoma cells displayed high level of Atg5 in both the = control=20 and TMZ-treated cells, as compared with that of = CD133⁺=20 cells.

view in a new=20 window

Figure 3. Differential = expression of=20 Atg5 in glioblastoma cells following TMZ treatment. = GFAP and=20 CD133 were shown as red, while Atg5 as green. Cell = nucleus=20 (blue) was counterstained by DAPI. White arrowhead,=20 Atg5-localized autophagosomes (Original magnification=20 =A1=C11000).

DISCUSSION

In 2004, Singh and colleagues³ = successfully isolated CSCs from different types of brain = tumors.=20 CSCs were found exclusively in the fraction of cancer cells=20 expressing CD133. An injection of only 100 CD133⁺ = cells=20 into the NOD-SCID mouse brain led to the growth of a tumor = that=20 could be serially transplanted and was histologically = identical to=20 the tumor harbored by the patient from whom these cells were = derived. In contrast, the CD133^=A8C tumor cells = failed to=20 form tumors, even when 1000-fold more CD133^=A8C = cells were=20 injected into the brains of the mice, suggesting that the = brain=20 tumor stem cells were always in the CD133⁺=20 population.³ In this study, we successfully = enriched=20 CD133⁺ GSCs from clinical specimen.

Activation of the cell death program has = been shown=20 to be responsible for chemotherapy-induced cytotoxicity in = tumor=20 cells, while alterations in the death machinery have been = related to=20 chemoresistance in gliomas.⁴ Eramo et = al¹⁰=20 suggested that drug resistance observed in GSCs may depend = on=20 abnormalities of the cell death pathway such as = overexpression of=20 antiapoptotic factors or silencing of key death effectors. A = study=20 by Liu and his colleagues⁶ demonstrated for the = first=20 time that an increased resistance of CD133⁺ brain = tumor=20 stem cells in response to treatment with chemotherapeutic = agents=20 including TMZ, compared with autologous CD133^=A8C = cells.=20 Gene expression studies revealed a higher expression of = multi-drug=20 resistance gene BCRP1 and DNA repair genes such as MGMT, as = well as=20 genes that inhibited apoptosis in the CD133 expressing = CSCs.^5,6=20

Autophagy represents an alternative=20 tumor-suppressing mechanism to overcome, at least partly, = the=20 dramatic resistance of many cancers to radiotherapy and = proapoptotic=20 related chemotherapy.⁷ Unlike apoptosis, = autophagy is a=20 caspase independent process characterized by the = accumulation of=20 autophagic vacuoles in the cytoplasm accompanied by = extensive=20 degradation of the organelles such as mitochondrias, = polyribosomes=20 and the endoplasmic reticulum, which precedes the = destruction of the=20 nucleus.⁷ Autophagy machinery can be activated by = various=20 agents such as DNA damages,¹¹ disruption of = PI3K/AKT/mTOR=20 signaling,¹² deprivation of nutrients or amino=20 acids,¹³ and oxidative stress.^14,15 = The=20 components of the molecular machinery responsible for = autophagy are=20 products of the autophagy-related (Atg) genes.¹⁶ = These=20 genes control a number of aspects of the autophagic process=20 including induction by Beclin-1 (Atg6) and autophagosomal = vesicle=20 formation through Atg12-Atg5 and LC3 (Atg8).¹⁷=20 Additionally, Bcl-2 binds Beclin-1 to inhibit = Beclin-1-dependent=20 autophagy, thereby functioning both as a pro-survival and as = an=20 anti-autophagic regulator.¹⁸

TMZ contributes significant therapeutic = benefits in=20 glioblastoma patient.^2,19 Part of TMZ cytotoxic = activity=20 is exerted through proautophagic processes, at least in = glioblastoma=20 cells, as a result of the formation of O6-methylguanine in = DNA,=20 which mispairs with thymine during the following cycle of = DNA=20 replication.⁸ Glioma cells thus respond to TMZ by = undergoing G2/M arrest, but ultimately die from=20 autophagy.^20,21 However, Our finding suggested = that=20 CD133⁺ GSCs were less susceptible to TMZ-induced=20 autophagy, as compared with CD133^=A8C cell = fractions. We=20 hypothesized that suppression of autophagy-related proteins = might=20 render GSCs strongly resistant to autophagic cell death. Our = further=20 study showed autophagy-related proteins such as Beclin-1, = Atg5 and=20 LC3 were still low in CD133⁺ cells following TMZ=20 treatment, as compared with CD133^=A8C cell = fractions.=20 Therefore, the differential response of CD133⁺ = and=20 CD133^=A8C cells to TMZ-induced autophagy might = contribute=20 to the development of TMZ resistance in GSCs and recurrence = after=20 TMZ treatment.

Although GSCs might not be susceptible to = classical=20 pathways to autophagy. Recently, Jiang et al^{22 =}reported=20 the use of an oncolytic adenovirus, delta-24-RGD, to target = the=20 abnormal p16INK4/Rb pathway in brain tumor stem cells. = Delta-24-RGD=20 induced enormous autophagic cell death both in cell line = models=20 in vitro and xenografts. Their results show for the = first=20 time that brain tumor stem cells are susceptible to=20 adenovirus-mediated cell death via autophagy. Therefore,=20 understanding the mechanisms of autophagy in GSCs may help = to=20 address this issue and might contribute to the development = of new=20 effective pharmaceutical approaches for the treatment of = brain=20 tumors.

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20. Kanzawa T, Bedwell J, Kondo Y, Kondo = S, Germano=20 IM. Inhibition of DNA repair for sensitizing resistant = glioma cells=20 to temozolomide. J Neurosurg 2003; 99: 1047-1052.

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=A1=A4 Prognostic=20 factors influencing clinical outcomes of = glioblastoma=20 multiforme

=A1=A4 Antitumor=20 activity of F90, an epidermal growth factor = receptor=20 tyrosine kinase inhibitor, on glioblastoma cell = line=20 SHG-44

=A1=A4 Microarray=20 analysis of gene expression profile of multidrug = resistance in pancreatic cancer

=A1=A4 = Chromosome=20 14q may harbor multiple tumor suppressor genes = in=20 primary glioblastom a multiformec

=A1=A4 = Genome-wide=20 allelotype study of primary glioblastoma=20 multiforme

=A1=A4 = A=20 genome-wide allelotype study of primary and=20 corresponding recurrent glioblastoma multiforme = in one=20 = patient

=A92005-2008 Chinese Medical Journal, All = Rights=20 Reserved. ICP:05052599

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{form1.submit(); } } function checkinput_reply1() { document.form1.checkinput.value=3D"nopublish"; document.form1.action=3D"replyregister.asp"; document.form1.target=3D"_blank"; if (checkid(form1)!=3Dfalse) {form1.submit(); } } //=C9=E8=D6=C3=D7=D6=CC=E5=B4=F3=D0=A1 function SetFont(SetFontid, size){ document.getElementById(SetFontid).style.fontSize=3Dsize } //=D1=E9=D6=A4=CA=C7=B7=F1=D1=A1=D4=F1=C1=CB=BC=C7=C2=BC function checkid(form) { var blnAlert =3D 0; if (typeof(form.pid.length)=3D=3D"undefined") { =20 if(form.pid.checked=3D=3Dtrue) =20 blnAlert =3D blnAlert+1; } else { for (i=3D0;i"9"){ return false;}}} return true; } //=D1=E9=D6=A4=B8=DF=BC=B6=BC=EC=CB=F7 function CheckAdSearch(form){ if (isfilled(form.pyear)!=3Dfalse){ if (isint(form.pyear)=3D=3Dfalse){ alert("Published year should be a number=A3=A1"); form.pyear.focus(); return false; }} if (isfilled(form.pissue)!=3Dfalse){ if (isint(form.pissue)=3D=3Dfalse){ alert("Published issue should be a number=A3=A1"); form.pissue.focus(); return false; }} } //=D1=E9=D6=A4=BE=AB=C8=B7=BC=EC=CB=F7 function CheckExSearch(form){ if (isint(form.pissue)=3D=3Dfalse){ alert("Published issue should be a number=A3=A1"); form.pissue.focus(); return false; } if (isint(form.pyear)=3D=3Dfalse){ alert("Published year should be a number=A3=A1"); form.pyear.focus(); return false; } if (isfilled(form.firstpage)=3D=3Dfalse){ alert("First page should be a number=A3=A1"); form.firstpage.focus(); return false; } if (isfilled(form.pissue)=3D=3Dfalse){ alert("Published issue should be a number=A3=A1"); form.pissue.focus(); return false; } if (isfilled(form.pyear)=3D=3Dfalse){ alert("Published year should be a number=A3=A1"); form.pyear.focus(); return false; } if (isint(form.firstpage)=3D=3Dfalse){ alert("First page should be a number=A3=A1"); form.firstpage.focus(); return false; } } //=D1=E9=D6=A4=C6=C0=C2=DB function CheckResponse(form){ if (isfilled(form.username)=3D=3Dfalse){ alert("Your name should not be 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