行政院國家科學委員會補助專題研究計畫成果報告

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1 行政院國家科學委員會補助專題研究計畫成果報告 前胸腺素基因治療併用樹突狀細胞策略治療膀胱癌 可行性之評估 (3 / 3) 計畫類別 : 個別型計畫 整合型計畫計畫編號 :NSC B-6-3-M51 執行期間 : 9 年 8 月 1 日至 91 年 7 月 31 日 計畫主持人 : 吳昭良 共同主持人 : 蕭璦莉 本成果報告包括以下應繳交之附件 : 赴國外出差或研習心得報告一份 赴大陸地區出差或研習心得報告一份 出席國際學術會議心得報告及發表之論文各一份 國際合作研究計畫國外研究報告書一份 執行單位 : 國立成功大學生物化學研究所 中華民國 91 年 1 月 3 日 1

2 行政院國家科學委員會專題研究計畫成果報告前胸腺素基因治療併用樹突狀細胞策略治療膀胱癌可行性之評估 (3/3) Preparation of NSC Project Reports 計畫編號 :NSC B-6-3-M51 執行期限 :9 年 8 月 1 日至 91 年 7 月 31 日主持人 : 吳昭良國立成功大學生物化學研究所共同主持人 : 蕭璦莉國立成功大學微生物免疫學研究所計畫參與人員 : 吳惠如國立成功大學生物化學研究所 一 中文摘要 在病理學上, 約有 2% 膀胱癌屬於侵犯性, 其癒後較差 ; 其他 75~8 % 的膀胱癌是屬於表淺層的移行性上皮細胞癌 ; 後者經手術切除合併卡介苗灌洗膀胱, 雖然可降低復發率, 然而仍有相當多的病人復發, 甚至惡化轉移 ; 部分癌症轉移病人對化學療法有反應, 但是終究治療無效而死亡 由於有效治療膀胱癌的方法仍然不盡理想, 新的治療策略有待發展 利用細胞素基因植入腫瘤細胞, 以提高抗腫瘤的免疫反應的治療策略, 近幾年被廣泛的研究, 其主要運用腫瘤細胞分泌細胞素, 改變腫瘤生長部位的免疫刺激, 以提高具腫瘤專一性免疫細胞的活化, 這種合併基因治療及免疫治療的新興療法, 可以在腫瘤生長部位產生局部高濃度的細胞素, 以引發局部且持續的免疫反應 ; 相較於免疫調節物質的系統性免疫治療, 以免疫調節物質基因治療的策略, 不但可以保有免疫系統對抗病灶的專一性, 也可免除高濃度細胞素造成系統性的毒性 在本項研究中, 我們的策略是利用基因工程構築會表現前胸腺素蛋白之反轉路病毒, 並評估重組病毒對膀胱癌在小鼠模式免疫基因治療的能力 關鍵詞 : 基因治療 膀胱癌 病毒載體 Abstract To explore the potential use of prothymosin α (ProT), a putative thymic hormone, in gene therapy for bladder cancer, we generated a replication-defective recombinant retroviral vector encoding ProT and tested its antitumour effect on the MBT-2 murine bladder cancer. C 3 H/HeN mice injected with MBT-2 cells in conjunction with retroviruses encoding ProT exhibited smaller tumour mass, lower tumour incidence and higher survival rate, as well as higher antitumour cytotoxic activities compared with those injected with control viruses. However, such effects were not observed in severe combined immunodeficiency mice, suggesting that ProT exerts antitumour effects through its immunomodulatory activities. Since ProT enhanced growth and colony formation in soft agar of MBT-2 cells in vitro, a retroviral vector encoding ProT lacking the nuclear localisation signal (NLS) was constructed to circumvent its proliferation-promoting effect on tumour cells. The retroviruses encoding ProT lacking NLS exhibited higher antitumour effects than those expressing wild-type ProT. This is the first report indicating that ProT, in particular NLS-deleted ProT, delivered by retroviral vectors may be further explored for the treatment of bladder cancer. Keywords: prothymosin; retroviral vector; bladder tumour 二 緣由與目的 Conventional therapy often fails in the treatment of solid tumours despite complete surgical resection. With the development of 2

3 effective techniques to deliver genes into target cells, gene therapy becomes a promising alternative in the treatment of cancer. Introduction of cytokine genes with immunostimulatory or antitumour activities have shown promise to inhibit tumour growth in animal models. Because thymic peptides, such as thymosin a1 extracted originally from calf thymus, promote the maturation of T cells, they are used in clinical trials for the treatment of patients with cancer and immunodeficiency. Prothymosin á (ProT), the putative precursor of thymosin á1 that contains 113 amino acid residues with the thymosin ál sequence at its N-terminus, was originally isolated from rat thymus.2 Many data show the behaviour of ProT, like thymosin á1, as a biological response modifier. Furthermore, ProT has been reported to be more effective than thymosin á1 in protection of the mice infected with Candida albicans, suggesting that the biological activity of ProT was not solely determined by its N-terminal region that includes the thymosin á1 fragment. Bladder cancer, especially presenting as superficial disease, is responsive to immunotherapeutic agents such as BCG, and may represent a good candidate for immunological intervention. Indeed, in the murine MBT-2 bladder tumour model, irradiated, interleukin-2 (IL-2) or GM-CSF gene-modified MBT-2 cells were capable of curing about half of the mice bearing wild-type MBT-2 tumours and engendered protective immunological memory in the cured mice. Because ProT induces T-cell maturation and differentiation, and possesses antitumour effects, in this study we investigated the feasibility of retrovirus-mediated ProT gene therapy for murine bladder cancer. Since ProT is implicated recently as a nuclear protein associated with cell proliferation, to circumvent the growth-promoting activity of ProT, we also constructed the retroviral vector encoding ProT deletion mutant lacking the nuclear localisation signal (NLS) and examined its antitumour efficacy. In this report, we showed that retroviruses encoding mouse ProT suppressed MBT-2 tumour growth in syngeneic C3H/HeN mice, but not in severe combined immunodeficient (SCID) mice, suggesting that ProT exerts antitumour effects through its immunomodulatory activities. Moreover, retroviruses encoding ProT lacking NLS exhibited higher antitumour effects than those encoding wild-type ProT. Our results reveal that retrovirus-mediated gene transfer of ProT, in particular ProT lacking NLS, may be further explored for the treatment of bladder cance 三 結果與討論 Characterisation of ProT gene-modified MBT-2 cell clones The expressions of the transgene in ProT gene-modified MBT-2 cell clones were first analysed by reverse transcriptase-polymerase chain reaction (RT-PCR). In MBT-2/ProT clones, mrnas specifying the neo and ProT gene products were detectable, whereas only the neo signal was detected in MBT-2 cells transfected with the vector control, and neither were present in parental MBT-2 cells (data not shown). A bioassay based on thymocyte proliferation for thymic hormones was used to analyse the expression of the ProT protein. The conditioned media from MBT-2/ProT clones and recombinant ProT, serving as the positive control, showed a dose-dependent enhancement of thymocyte proliferation in the presence of suboptimcal concentration of mitogen, whereas those from parental cells had no effect (Figure 1A). Taken together, ProT gene-modified MBT-2 cells via retrovirus-mediated gene transfer secreted bioactive ProT protein. The results also indicate that the RUFProT retrovirus permits highly efficient gene transduction in MBT-2 cells. The proliferations of three ProT-transduced MBT-2 cell clones, one control vector-transduced clone and the parental cells were determined with the [ 3 H]-thymidine incorporation assay. Figure 1B shows that thymidine incorporation was higher in all three ProT gene-transduced MBT-2 clones compared with control vector-transduced or parental MBT-2 cells, whereas no difference was found between the 3

4 control vector-transduced clone and parental cells. The ability of colony formation of these cell clones was further analysed. Colony formation in soft agar was enhanced in all ProT gene-transduced cell clones, particularly in clone #1, when compared with the number of colonies formed by cells transfected with control vector or by the parental cells (Figure 1B). These results indicate that transduction of ProT gene into MBT-2 cells resulted in an enhancement of cell proliferation and colony formation in soft agar. ProT gene-transduced MBT-2 clones #1 and #14 as well as the parental cells were then inoculated subcutaneously (s.c.) into syngeneic C 3 H/HeN mice to measure the effect of ProT production on the tumourigenicity of MBT-2 cells. Whereas tumour grew in all C 3 H/HeN mice inoculated with parental MBT-2 cells, injection of ProT gene-transduced MBT-2 clones #1 and #14 failed to result in tumours in groups of 13 and 12 mice at 3 days postinjection, respectively. Induction of cytotoxic activities in mice inoculated with MBT-2 cells in conjunction with retroviruses encoding ProT To test whether the activity of lymphokine-activated killer (LAK) cells can be enhanced in mice coinjected with MBT-2 cells and retroviruses encoding ProT, spleen and lymph node cells were isolated from treated mice at 48 days postinjection and cultured in the presence of IL-2 for 4 days. The cells were then assayed for cytotoxic activity against MBT-2 using the 51 Cr-release assay. The results indicate that the lymphocytes of either spleens or lymph nodes from tumour-bearing mice exhibited very low cytolytic activities against MBT-2 cells. Coadministration with control retroviruses did not show any enhancement on LAK activities. In contrast, retroviruses expressing ProT significantly enhanced the LAK activities compared with control retroviruses or PBS. The augmentation of LAK activities was more pronounced in lymphocytes from the lymph nodes than those from the spleens (Figure 2). Effects of ProT on tumourigenesis in vivo To test the antitumour efficacy of retroviruses expressing ProT, the tumour incidence in mice inoculated s.c. with MBT-2 cells in conjunction with recombinant retroviruses encoding ProT was compared with those coinoculated with control viruses or PBS in C 3 H/HeN or SCID mice. Tumour grew in 78% of C 3 H/HeN mice inoculated s.c. with 1 6 MBT-2 cells. Coadministration with RUFneo retroviruses had no effect on tumour growth. However, those inoculated with MBT-2 cells in conjunction with RUFProT retroviruses exhibited only 3% of tumour incidence (Table 1). Moreover, as shown in Figure 3A, mice treated with control viruses had detectable tumours at day 9 postinjection and occupied a larger tumour sizes compared to those treated with retroviruses encoding ProT. Throughout the experiment, tumour growth was inhibited in mice treated with RUFProT retroviruses. To evaluate whether the immune response plays any roles in the RUFProT treatment, SCID mice were used for a similar experiment. Contrast to MBT-2 syngeneic C 3 H/HeN mice, the tumour volumes of mice coinjected with MBT-2 cells and RUFProT retroviruses were larger than those treated with control viruses (Figure 3B), suggesting that the antitumour effect of ProT is attributed to its immunomodulatory activities. Taken together, retroviruses encoding ProT exerted antitumour activity only in immunocompetent mice. Effects of the NLS within ProT on cell growth As shown in Figure 3B, the growth of larger tumours in SCID mice treated with MBT-2 in conjunction with retroviruses encoding ProT suggests that the ability of ProT to promote cell growth could be contradictory to its antitumour activity. We next examined whether the proliferation-promoting activity of ProT could be abolished if its NLS located in the C-terminus was deleted. To this end, we generated recombinant retroviruses encoding ProT lacking NLS, designated RUFProTΔ 4

5 NLS, and constructed eukaryotic plasmid vectors for expression of ProT and its NLS-deleted mutant tagged with enhanced green fluorescent protein (EGFP) in MBT-2 cells. Subcellular localisation of the respective proteins was analysed by fluorescent microscopy. EGFP-tagged ProT was localised exclusively in the nucleus, whereas EGFP-tagged ProT Δ NLS distributed in both the cytoplasm and the nucleus (data not shown). Furthermore, the proliferative activity was decreased in the three MBT-2 clones transduced with retroviruses encoding ProTΔNLS compared with cells transduced with wild-type ProT (Figure 4). These results suggest that growth-promoting effect of ProT can be abolished when its NLS sequence is deleted, and, therefore, retroviruses encoding ProTΔ NLS may exert more potent antitumour activity than those encoding wild-type ProT. Antitumour effects of retroviruses encoding ProTΔNLS The final question addressed in this study was whether the retroviruse encoding ProTΔNLS would exert higher antitumour activities in vivo compared with its wild-type counterpart. Mice that concomitantly received MBT-2 cells (5 1 5 ) and 1 5 colony-forming units (CFU) of RUFProTΔ NLS exhibited the lowest incidence of tumour growth compared with those concomitantly received MBT-2 cells and RUFProT or received MBT-2 cells and RUFneo. In terms of tumour size, mice coinoculated with MBT-2 cells and control retroviruses developed tumours at the site of injection and had to be killed due to large tumour burden by day 7. On the other hand, in mice inoculated with MBT-2 cells in conjunction with retroviruses encoding either ProT or ProTΔNLS, tumour growth was slower than those coinjected with control viruses. In mice that received RUFProTΔ NLS, tumour growth was significantly retarded and more than half of the mice survived for over 7 days. In mice coadministered with 1 5 MBT-2 cells and retroviruses (1 5 CFU), both the incidence of tumour growth and the tumour volume were reduced along with the prolongation of survival time in mice treated with RUFProT ΔNLS (Figure 5A). The Kaplan-Meier survival curve is illustrated in Figure 5B. Thus, the survival of RUFProT Δ NLS-treated mice was significantly better than that of the control mice. The essential role of the thymus in the development, maintenance and regulation of the immune system has been well documented. Thymic hormones have been demonstrated to be an effective immunopotentiating agent and can act in lieu on the thymus gland to reconstitute the immune function in immunosuppressed animals and in humans with a number of primary and secondary immunodeficiency. Moreover, they also show promise for treatment of cancer patients. Addition of thymic humoral factor-ã2 as an adjunct to anticancer chemotherapeutic regimes not only strengthens the antitumour activity of each drug, but also repairs tumour/chemotherapy-induced damage to T-cell subpopulations and functions in mice bearing immunogenic tumours or weakly immunogenic tumour such as Lewis lung carcinoma. 5 Furthermore, thymopentin evaluated as an immunotherapeutic agent for the treatment of human melanoma showed encouraging results. Histopathological and immunohistochemical evaluation of regressing metastatic nodules showed the presence of tumour infiltrating lymphocytes, necrosis, sclerosis, intratumoural vascular proliferation and microthrombosis. 6 Salvati et al. 7 demonstrated that chemo-immunotherapy induced a statistically significant enhanced response rate to malignant disease progression compared with chemotherapy alone in patients with advanced non-small-cell lung cancer receiving ifosfamide chemotherapy or chemotherapy followed by thymosin á1, employed together with low-dose interferon-á. It has been shown that ProT was equally effective in stimulating the chemotactic activity of polymorphonuclear cells (PMNs) from tumour patients and healthy donors. PMNs 5

6 from tumour patients, especially those derived from patients with breast tumour, were characterised by a significant enhancement of cytotoxicity against tumour target cells, as compared with healthy donors. ProT may improve some PMN functions of tumour patients, associated with the proposed role in host-tumour interaction. 8 In addition, we have demonstrated that the efficacy of DNA immunisation may be enhanced by the simultaneous expression of ProT serving as a vaccine adjuvant. 9 Collectively, thymic hormones, such as ProT, may be used as candidate genes for gene therapy in non-specific immunological enhancement of the host anti-neoplastic response. Apart from behaving as a biological response modifier, ProT is more recently implicated in cell proliferation and/or differentiation. ProT gene expression occurs ubiquitously in mammalian cells and that the product, apparently without proteolytic processing, is translocated to the nucleus. We have shown previously that overexpression of ProT accelerates cell proliferation by shortening the duration of the G1 phase of cell cycle. 1 The increase of ProT expression during cell proliferation is suggested as an indicator of tumour progression in several tumours In this study, we also demonstrated that ProT gene-transduced MBT-2 bladder cancer cells grew faster and formed more colonies in soft agar than their parental cells in vitro, suggesting that ProT may promote tumour growth, if the finding in vitro can be extrapolated to the in vivo setting. Thus, considering the Janus-faced nature of ProT towards tumour growth, we produced retroviruses carrying NLS-deleted ProT gene and used them to generate MBT-2 expressing ProT lacking the KKQK motif of the NLS. By using indirect immunofluorescence labeling and confocal scanning laser microscopy in relation to nuclear components involved in transcription, translation and splicing, Vareli et al. demonstrated that ProT exhibits a punctuated nuclear distribution and are excluded by nucleoli. The distribution of ProT in the nucleus is related to that of transcription sites and implies that ProT is involved in transcription. 14 Our results from the distribution of the ProT-EGFP fusion protein confirm the nuclear localisation of ProT. Furthermore, the exclusive nuclear localisation of ProT can be disturbed by deleting its NLS. In terms of cell growth, the cell proliferation of ProT Δ NLS gene-modified MBT-2 cells was decreased in vitro compared with that of ProT gene-modified MBT-2 cells. Thus, the proliferation-promoting effect of ProT, which may contribute, in part, to tumour growth, can be circumvented by using the NLS-deleted ProT mutant. In this study, the enhancement of the antitumour effects on murine bladder tumour by in vivo administration of retroviruses encoding ProT ΔNLS compared to those carrying the wild-type ProT gene may be due in part to its diminished effect on growth promotion. Although the biological role of ProT has been controversial, the potential therapeutic use of ProT for patients with immunodeficiency and cancer shows promise. In the work described here, the feasibility of ProT gene therapy for cancer was tested in the murine MBT-2 bladder cancer model. ProT gene-modified MBT-2 cells secreted ProT, which exhibited biological activity of promoting thymocyte proliferation. C 3 H/HeN mice inoculated with MBT-2 cells in conjunction with RUFProT retroviruses induced higher levels of LAK activities compared to those treated with MBT-2 cells along with control viruses, and the enhanced induction of these cytotoxic activities may contribute, in part, to the antitumour responses found. In our tumour model where ex vivo ProT-gene-transduced MBT-2 cells were implanted into C 3 H/HeN mice, loss of tumourigenicity was more significant than the model with in vivo coadministration of parental MBT-2 cells with RUFProT retroviruses. This is in agreement with the notion that the efficiency of retroviral gene transfer in vivo is low compared with ex vivo gene transfer into tumour cells. Nevertheless, we have shown previously that retroviral vectors carrying the IFN-γ gene transduced MBT-2 cells in vivo, which resulted in enhancing local antitumour activities. 15 We 6

7 presumed that in vivo transduction of tumour cells with ProT gene may have occurred in mice coadministered with tumour cells and retroviral vectors carrying the ProT gene. However, the mechanism of ProT-induced inhibition of tumour growth has not been completely elucidated, which requires further investigation. The SCID mice, Prkdc scid, are defective in the gene encoding the catalytic subunits of DNA-dependent protein kinase. Mice having homozygous deficiency of Prkdc have no detectable IgM, IgG1, IgG2a, IgG2b, IgG3, or IgA, as well as one-tenth or less in the size of the lymphoid organs compared with their normal counterpart. Thymus, lymph nodes and splenic follicles are virtually devoid of lymphocytes and deficient in both B and T cell functions. 16 In this study, the homozygous Prkdc scid mice were used to investigate the antitumour effect of ProT. ProT administered to immunocompetnet, syngeneic C 3 H/HeN mice bearing MBT-2 tumours exerted antitumour effects. However, such effects were not observed in SCID mice, suggesting that the antitumour activity of ProT in vivo could be attributable to its immunomodulatory properties. It was noted that SCID mice receiving ProT gene-modified MBT-2 cells even had larger tumour burden compared with those receiving parental MBT-2 cells, which might be due to the growth-promoting activity mediated by ProT. In the work described here, direct administration of retroviruses carrying mouse ProT gene suppressed tumour growth and prolonged survival in mice with primary MBT-2 tumours. Moreover, retroviruses encoding ProT ΔNLS exhibited higher antitumour activity than those encoding wild-type ProT. In conclusion, our results indicate that gene therapy by retrovirus-mediated ProT gene transfer may be an effective approach for the treatment of bladder cancer. This is the first report to exploit ProT gene therapy as an antitumour strategy. Alterations in local appearance of thymic hormones, produced by in vivo viral vector-mediated gene transfer, may therefore be useful in the immunotherapy of cancer. 四 計畫成果自評 In the work described here, direct administration of retroviruses carrying mouse ProT gene suppressed tumour growth and prolonged survival in mice with primary MBT-2 tumours. Moreover, retroviruses encoding ProTÄNLS exhibited higher antitumour activity than those encoding wild-type ProT. In conclusion, our results indicate that gene therapy by retrovirus-mediated ProT gene transfer may be an effective approach for the treatment of bladder cancer. This is the first report to exploit ProT gene therapy as an antitumour strategy and it has been published in Gene Therapy 8: titled as Shiau, A.L., Lin, P.R., Chang, M.Y., and Wu, C.L. (21) Retrovirus-mediated transfer of prothymosin gene inhibits tumor growth and prolongs survival in murine bladder cancer. 五 參考文獻 1 Cordero OJ, Maurer HR, Nogueira M. Novel approaches to immunotherapy using thymic peptides. Immunol Today 1997; 18: Haritos AA, Goodall GJ, Horecker BL. Prothymosin alpha: isolation and properties of the major immunoreactive form of thymosin alpha 1 in rat thymus. Proc Natl Acad Sci USA 1984; 81: Saito S et al. Immunotherapy of bladder cancer with cytokine gene-modified tumor vaccines. Cancer Res 1994; 54: Connor J et al. Regression of bladder tumors in mice treated with interleukin 2 gene-modified tumor cells. J Exp Med 1993; 177: Ophir R et al. Thymic humoral factor-gamma 2 (THF-gamma 2) immunotherapy reduces the metastatic load and restores immunocompetence in 3LL tumor-bearing mice receiving anticancer chemotherapy. Immunopharmacol Immunotoxicol 1996; 18:

8 6 Cascinelli N et al. Evaluation of clinical efficacy and tolerability of intravenous high dose thymopentin in advanced melanoma patients. Melanoma Res 1998; 8: Salvati F et al. Combined treatment with thymosin-alpha1 and low-dose interferon-alpha after ifosfamide in non-small cell lung cancer: a phase-ii controlled trial. Anticancer Res 1996; 16: Heidecke H, Eckert K, Schulze-Forster K, Maurer HR. Prothymosin alpha 1 effects in vitro on chemotaxis, cytotoxicity and oxidative response of neutrophils from melanoma, colorectal and breast tumor patients. Int J Immunopharmacol 1997; 19: Shiau AL, Chu CY, Su WC, Wu CL. Vaccination with the glycoprotein D gene of pseudorabies virus delivered by nonpathogenic Escherichia coli elicits protective immune responses. Vaccine 21; 19: Wu CL, Shiau AL, Lin CS. Prothymosin alpha promotes cell proliferation in NIH3T3 cells. Life Sci 1997; 61: Bodey B, Bodey B, Jr., Siegel SE, Kaiser HE. Review of thymic hormones in cancer diagnosis and treatment. Int J Immunopharmacol 2; 22: Magdalena C, Dominguez F, Loidi L, Puente JL. Tumour prothymosin alpha content, a potential prognostic marker for primary breast cancer. Br J Cancer 2; 82: Mitani M, et al. Significance of plasma thymosin alpha1 measurements in gastric cancer patients. World J Surg 2; 24: Vareli K et al. Nuclear distribution of prothymosin alpha and parathymosin: evidence that prothymosin alpha is associated with RNA synthesis processing and parathymosin with early DNA replication. Exp Cell Res 2; 257: Shiau AL, Lin CY, Tzai TS, Wu CL. Postoperative immuno-gene therapy of murine bladder tumor by in vivo administration of retroviruses expressing mouse interferon-gamma. Cancer Gene Ther 21; 8: Jhappan C et al. DNA-PKcs: a T-cell tumour suppressor encoded at the mouse scid locus. Nat Genet 1997; 17: Danos O, Mulligan RC. Safe and efficient generation of recombinant retroviruses with amphotropic and ecotropic host ranges. Proc Natl Acad Sci U S A 1988; 85: Shiau AL, Yang HM, Wu P, Wu CL. Provision of positive and negative selections in retroviral vectors containing the cytosine deaminase gene. Gene Ther 1998; 5: Evstafieva AG et al. Overproduction in Escherichia coli, purification and properties of human prothymosin alpha. Eur J Biochem 1995; 231: Table 1. Incidence of tumour growth in C 3 H/HeN mice inoculated with MBT-2 cells in the presence or absence of recombinant retroviruses. Treatment Tumour incidenc Percentage MBT-2 7/9 78% MBT-2 + RUFneo 8/9 89% MBT-2 + RUFProT 3/1 3% On day, MBT-2 cells (1 6 ) admixed with 1.5 x 1 5 CFU of retroviruses encoding ProT or neo, or with PBS were injected s.c. into groups of 7-week old C 3 H/HeN mice. The incidence rate of tumour growth at 25 days postinjection is represented as: (numbers of tumour-bearing mice)/(total numbers in each group). 8

9 Colony number Stimulation index A B 1:2 1:8 1:32 1:128 ProT MBT-2 RUF Colony number Growth Treatment MBT-2 RUF Clone [ 3 H]-thymidine incorporation (cpm) Figure 1. The biological behaviours of ProT gene-modified MBT-2 cells. (A) The biological activities of ProT in the conditioned media of gene-modified MBT-2 clones were measured by thymocyte proliferation assay. Thymocytes (5 1 5 ) were incubated with various dilutions of the culture supernatants from various clones or with purified recombinant ProT (7 ng) in the presence of.125 ìg/ml of Con A for 4 days. Cells were pulsed with [ 3 H]-thymidine (.5 ìci/well) for 16 h before harvest and cell proliferation was determined by [ 3 H]-thymidine incorporation. (B) Comparisons of the numbers of colony formation in soft agar and the growth rate among gene-modified MBT-2 clones. The colony numbers were calculated after 14 days of incubation in.33% soft agar. Each clone was determined in quadruplicate. The growth rates of MBT-2 derivatives were measured by [ 3 H]-thymidine incorporation after 3-day incubation. Error bars represent the standard error calculated from hexaplicate measurements. Data are analysed by Student s test. Cytotoxicity (%) Cytotoxicity (%) A B 2 : 1 1 : 1 5 : 1 25 : 1 2 : 1 1 : 1 5 : 1 25 : 1 E/T ratio PBS RUFneo RUFProT Figure 2. Induction of LAK activity against MBT-2 cells by (A) the splenocytes, and (B) the lymphocytes from lymph nodes near the tumour site in mice coadministered with MBT-2 cells and RUFProT or RUFneo retroviruses, or PBS. C 3 H/HeN mice were injected with MBT-2 cells along with retroviruses or PBS. Forty-eight days later, lymphocytes from spleens and lymph nodes were assayed for cytotoxic activities against MBT-2 cells with the 51 Cr-release assay. E/T ratio denotes effector cell to target cell ratio. 9

10 Tumor volume (mm 3 ) A B MBT-2 + RUFneo MBT-2 + RUFProT Days after inoculation Figure 3. Effect of ProT on tumour growth in mice. MBT-2 cells (1 6 ) admixed with retroviruses encoding ProT or with control viruses ( CFU) were inoculated s.c. into groups of (A) C 3 H/HeN mice or (B) SCID Prkdc mice. The tumour volume in each mouse is shown and the curves denote the mean tumour volume in each group. [ 3 H]-thymidine incorporation (cpm) MBT-2 Vector ProT NLS1 NLS2 NLS3 Figure 4. Growth rates of ProT gene- or ProT NLS gene-modified MBT-2 clones. Various MBT-2 clones transduced with ProT or ProT NLS gene were plated in 96-well plates (2 cells/well). After serum starvation for 24 h, cells were cultured in 1% FCS for 48 h. Cells were then pulsed with [ 3 H]-thymidine (.5 ìci/well) for 6 h before harvest and cell proliferation was determined by [ 3 H]-thymidine incorporation. Error bars represent the standard error calculated from hexaplicate measurements. Data are compared by Student s test. Mean tumor volume ( mm 3 ) Survival ( % ) A B MBT-2 + RUFneo MBT-2 + RUF/ProT MBT-2 + RUF/ProT NLS Days after inoculation Figure 5. Antitumour effects of retroviruses encoding ProT or ProT NLS in the MBT-2 tumour model. Groups of C 3 H/HeN mice were inoculated s.c. with MBT-2 cells (5 1 5 ) along with 1 5 CFU of RUFneo, RUFProT, or RUFProT NLS. (A) The individual tumour volume is measured as (length of tumour) (width of tumour) 2.45 (mm 3 ). The mean tumour volume is represented as the sum of the individual tumour volume in tumour-bearing mice divided by total mouse number in each group. Error bars represent standard error. (B) The survival curves of mice in each group are shown. The survival analysis was performed using the Kaplan-Meier survival curve and the log-rank test (p =.87). ** 1

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