From: Subject: Bone Marrow-generated Dendritic Cells Pulsed with Tumor Extracts or Tumor RNA Induce Antitumor Immunity against Central Nervous System Tumors -- Ashley et al. 186 (7): 1177 -- The Journal of Experimental Medicine Date: Thu, 16 Nov 2006 18:14:04 +0100 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0001_01C709AB.0016CBA0" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.2962 This is a multi-part message in MIME format. ------=_NextPart_000_0001_01C709AB.0016CBA0 Content-Type: text/html; charset="iso-8859-1" Content-Transfer-Encoding: quoted-printable Content-Location: http://www.jem.org/cgi/content/full/186/7/1177?ijkey=ccf7f3a89a765f99277aef129982d855cc6c1846 Bone Marrow-generated Dendritic Cells Pulsed with = Tumor Extracts or Tumor RNA Induce Antitumor Immunity against Central = Nervous System Tumors -- Ashley et al. 186 (7): 1177 -- The Journal of = Experimental Medicine
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J. Exp. Med.
=A9 The Rockefeller University=20 Press
0022-1007/97/10/1177/06 $2.00
Volume 186, Number 7, October = 6, 1997=20 1177-1182

BRIEF DEFINITIVE = REPORT:
Bone=20 Marrow-generated Dendritic Cells Pulsed with Tumor Extracts or Tumor RNA = Induce=20 Antitumor Immunity against Central Nervous System Tumors

By David M. Ashley,* = Brenda=20 Faiola,3D"par=A7 Smita = Nair, Laura P. Hale,3D"Dagger Darell D. Bigner,3D"Dagger and Eli Gilboa3D"par=A7

From the * Department of Pediatrics, 3D"Dagger Department of Pathology, = =A7 Department=20 of Immunology, and 3D"par Department of Surgery, Duke University = Medical=20 Center, Durham, North Carolina 27710

Summary=20
MATERIALS AND METHODS=20
RESULTS=20
DISCUSSION=20
Footnotes=20
Acknowledgements=20
REFERENCES

Summary =20

Recent studies have shown that the brain is not a barrier to = successful=20 active immunotherapy that uses gene-modified autologous tumor = cell=20 vaccines. In this study, we compared the efficacy of two = types of=20 vaccines for the treatment of tumors within the central = nervous=20 system (CNS): dendritic cell (DC)-based vaccines pulsed with = either=20 tumor extract or tumor RNA, and cytokine gene-modified tumor=20 vaccines. Using the B16/F10 murine melanoma (B16) as a model = for CNS=20 tumor, we show that vaccination with bone marrow-generated = DCs,=20 pulsed with either B16 cell extract or B16 total RNA, can = induce=20 specific cytotoxic T lymphocytes against B16 tumor cells. = Both types=20 of DC vaccines were able to protect animals from tumors = located in=20 the CNS. DC-based vaccines also led to prolonged survival in = mice=20 with tumors placed before the initiation of vaccine therapy. = The=20 DC-based vaccines were at least as effective, if not more so, = as=20 vaccines containing B16 tumor cells in which the granulocytic=20 macrophage colony-stimulating factor gene had been modified. = These=20 data support the use of DC-based vaccines for the treatment = of=20 patients with CNS tumors.

The concept that the brain is an immunologically privileged site has = been=20 supported clinically by the failure of central nervous system = (CNS)=20 tumors to respond to immunotherapy protocols that were = successful=20 systemically (1,=20 2).=20 Recently, several groups including ours have described active = immunotherapy protocols using intradermal vaccines of = genetically=20 modified tumor cells as being effective in rodent brain tumor = models=20 (3,=20 4).=20 However, animal studies have shown that immunization with = CNS-derived=20 tumor material can induce fatal experimental allergic = encephalitis=20 (5).=20 Due to the limited ability of reliably obtaining and growing = a=20 high percentage of tumor specimens without contamination by = normal=20 nervous tissue, the application of similar approaches to = human=20 patients with brain tumors may therefore carry the risk of = causing=20 such autoimmune complications (6,=20 7).=20

The dendritic cell (DC) network is a specialized system for = presenting=20 antigen to naive or quiescent T cells, and it plays a central = role in=20 the induction of T and B cell immunity in vivo. Immunizations = using=20 DCs loaded with tumor antigens may, therefore, represent a = powerful=20 method of inducing antitumor immunity. Recent studies have = shown that=20 immunizing mice with DCs pulsed with specific antigens can = prime a=20 CTL response that is tumor-specific and engenders protective = tumor=20 immunity in the treated mice (8).=20 Immunization using defined tumor antigens is, however, = limited at=20 present to a handful of human tumor types in which candidates = for=20 tumor rejection antigens have been identified (11).=20 More recently, effective tumor immunity in mice was induced = using DCs=20 pulsed with unfractionated tumor-derived antigens in the form = of=20 peptides (12,=20 13),=20 cell sonicates (14),=20 or messenger RNA (mRNA; 15). The advantages of vaccinating = with total=20 tumor-derived material are that the identity of the tumor = antigen(s)=20 need not be known and that the presence of multiple tumor = antigens=20 reduces the risk of antigen-negative escape mutants. The = potential=20 benefit of using total tumor antigens in the form of mRNA is = that it=20 can be amplified from a small number of tumor cells. Hence, = DC=20 vaccine treatment may be extended to patients with brain = tumors from=20 which only a small, possibly microscopic, biopsy can be taken = for=20 diagnosis. Furthermore, isolating bona fide tumor cells from = patient=20 specimens by ex vivo purification methods and combing this = with the=20 use of RNA subtractive hybridization techniques may reduce = the=20 concentration of self, potentially autoreactive, antigens in = the=20 vaccine preparation. This would be of crucial importance for=20 vaccinations with CNS tumor-derived antigens, as it may = diminish the=20 risk of severe autoimmune complications.

The studies presented here evaluated and compared the efficacy of = DC-based=20 tumor vaccines pulsed with either tumor extract or = tumor-derived=20 total RNA, with that of tumor vaccines in which the gene for = GM-CSF=20 had been modified. The vaccines were studied in a model of = active=20 immunotherapy for CNS tumors.

MATERIALS AND METHODS

Tumor Cell Lines and Animal Models. The B16/F10 = murine=20 melanoma cell line (B16) derived from a spontaneous melanoma in a = C57BL/6 mouse=20 (H-2b) was provided by I. Fidler (M.D. Anderson Cancer = Center,=20 Houston, TX; reference 16).=20 The SMA 560 cell line was derived from an intracerebral=20 transplant of a spontaneous astrocytoma from a VM/Dk mouse=20 (H-2b) (17).=20 The SMA 560 cell line was chosen as a control for the = B16, since=20 both cell lines are derived from neural crest. EL-4 = (H-2b)=20 murine thymoma cells were obtained from American Type Tissue = Culture=20 Collection (Rockville, MD). Cell lines were grown in zinc = option=20 medium (GIBCO BRL,=20 Gaithersburg, MD) containing 5% (vol/vol) FCS. All cell lines = were=20 shown to be free from Mycoplasma contamination as = previously=20 described (18).=20 All experiments used 6-12-wk-old female C57BL/6 mice (The Jackson Laboratory, = Bar Harbor,=20 ME), which were maintained in a virus-free environment in=20 accordance with the Laboratory Animal Resources Commission=20 standards.

DC Generation from Bone Marrow Cultures. The = procedure used=20 in these studies was the same as previously described (8,=20 19).=20 In brief, bone marrow was flushed from the long bones of the = limbs=20 and depleted of red cells with ammonium chloride. Bone marrow = cells=20 were depleted of lymphocytes, granulocytes, and = Ia+ cells=20 using a mixture of mAbs and complement. The mAbs used were=20 2.43 or 53-6.72 (CD8), GK1.5 (CD4), RA3-3A1/6.1 (CD45R), = B21-2 anti-Ia (Tumor Immunology Bank=20 210, 105, 207, 146, and 229, respectively; = American Type Tissue Culture Collection), and RB6-8C5 = anti-Gr-1=20 (provided by DNAX, Palo Alto, CA). Cells were plated in = 6-well=20 culture plates (106 cells/ml, 3 ml/ well) in RPMI=20 1640 medium supplemented with 5% heat-inactivated FCS,=20 50 mM 2-mercaptoethanol, 10 mM Hepes (pH 7.4), = 2 mM=20 glutamine, 100 U/ml penicillin, 100 mg/ml streptomycin, = and=20 3.3 ng/ml GM-CSF (provided by Amgen, Inc., Thousand Oaks, = CA).=20 On day 3 of culture, floating cells were gently removed, = and=20 fresh medium was added. On day 7 of culture, nonadherent cells = and loosely adherent proliferating DC aggregates were = collected=20 and replated in 100-mm Petri dishes (106 cells/ml,=20 10 ml/dish). At 10 d of culture, nonadherent cells = (DCs)=20 were removed for analysis and immunizations.

The quality = of DC=20 preparation was characterized by cell surface marker analysis, = morphological=20 analysis, and the ability of the preparation to induce = OVA-specific=20 CTLs in immunized mice as previously described (data not = shown;=20 references 19,=20 20).=20

Vaccination with DCs Pulsed with Tumor Extracts. = DCs were=20 washed twice in Opti-MEM medium (GIBCO BRL) and then = resuspended=20 at 5-10 =D7 106 cells/ml in 50 ml = polypropylene tubes=20 (Falcon, Lincoln Park, NJ). The cationic lipid, DOTAP, (Boehringer Mannheim=20 Biochemicals, Indianapolis, IN) was used to deliver tumor = extracts into cells. Tumor extracts were obtained by = sonicating tumor=20 cells in Opti-MEM (107 cells/500 =B5l) using a = Special=20 Ultrasonic Cleaner (Laboratory Supplies Company, Hicksville, = NY) and=20 were used without any further manipulation. Tumor extracts=20 (500 =B5l) and DOTAP (125 =B5g in 500 =B5l Opti-MEM = medium) were=20 mixed in 12 =D7 75-mm polystyrene tubes at room = temperature=20 for 20 min. The complex was added to the DCs and = incubated at=20 37=B0C in a water bath with occasional agitation for = 25 min. The=20 cells were washed and resuspended in PBS (105 extract-pulsed = DCs in=20 500 =B5l PBS/mouse) for intraperitoneal immunizations. =

Vaccination Using RNA-pulsed DCs. Total RNA was = isolated=20 from actively growing tissue culture cells by standard methods as = previously=20 described (15).=20 Pulsing DCs with RNA was performed in serum-free Opti-MEM = medium (GIBCO BRL) = as=20 described for tumor extracts with the following modifications. = RNA=20 (25 =B5g in 250 =B5l Opti-MEM medium) and DOTAP (50 =B5g = in 250=20 =B5l Opti-MEM medium) were mixed in 12 =D7 75 mm = polystyrene=20 tubes at room temperature for 20 min. The complex was = added to=20 the DCs (2-5 =D7 106 cells/ml) and = incubated at=20 37=B0C in a water bath with occasional agitation for = 25 min. The=20 cells were washed twice and resuspended in PBS = (105=20 RNA-pulsed DCs in 500 =B5l PBS/mouse) for intraperitoneal = immunizations.=20

PBS, B16 extract from 105 cells in PBS, or DCs = prepared as=20 described above were injected intraperitoneally in a volume of 500=20 =B5l.=20

Vaccination with GM-CSF-secreting Tumor Cells. = Crip cells=20 genetically engineered to produce replication-incompetent recombinant = retrovirus=20 with an amphotropic host range and with the ability to encode = the=20 cDNA for the murine GM-CSF gene were used as previously = described (4,=20 21).=20 B16 cells were infected by exposure to viral supernatants = from these=20 cells in the presence of polybrene (Sigma Chemical Co., St. = Louis,=20 MO). GM-CSF production from B16 cells exposed to virus was=20 confirmed by ELISA (Endogen, Cambridge, MA) and standard = bioassays=20 (21).=20

B16 parent cells with the modified GM-CSF gene and the = untransfected=20 cells were harvested, washed once in serum-containing medium, = and=20 washed twice in PBS. Cell pellets were resuspended in PBS at=20 105/500 =B5l, irradiated (3,500 centiGray), and injected = subcutaneously. The subcutaneous route of administration was = chosen=20 because we previously demonstrated that this is the preferred = method=20 for administering tumor vaccines (data not shown). =

In Vitro Cytotoxicity Assay. In vitro = cell-mediated=20 cytotoxicity assays were performed using standard procedures as we = previously=20 described (22).=20 In this study, splenocytes obtained from immunized animals = and=20 controls were restimulated in vitro for 5 d on = monolayers of=20 irradiated and mitomycin C-treated B16 cells or SMA = 560 cells.=20 Target cells included SMA 560 and B16 cells. =

Implantation of Brain Tumors. B16 cells were = harvested by=20 trypsinization, washed twice in Dulbecco's PBS, and mixed with an equal = volume=20 of 10% methylcellulose in zinc option medium. The cells=20 (500 cells in a volume of 5 =B5l) were then = implanted into=20 the right caudate nucleus of the brain of C57BL/6 mice by=20 stereotactic injection as previously described (3,=20 4).=20

Statistical Analysis. Survival estimates and = median=20 survivals were determined using the method of Kaplan and Meier (23).=20 Survival data was compared using Wilcoxon's log-rank test. = Student's=20 t test was used for calculating the significance of = other=20 data. Statistical significance was determined at the <0.05 = level.

RESULTS =20

Vaccination with Bone Marrow-derived DCs Pulsed with Tumor = Extract or=20 Tumor RNA Induces Tumor-specific CTLs.=20

To test whether bone marrow-derived DCs pulsed with tumor extracts or = tumor=20 RNA are capable of inducing tumor-specific CTLs, we first = immunized=20 C57BL/6 mice with three intraperitoneal injections of DCs = spaced=20 1 wk apart. Standard cytotoxicity assays were performed = using=20 splenocytes harvested from immunized animals 7 d after the = third=20 immunization and restimulated for 5 d in vitro with irradiated = B16 cells. Cytotoxic activity was tested against B16 cells. = We=20 previously demonstrated that the B16 is a poorly immunogenic = tumor=20 cell line such that no induction of CTLs occurs after = immunization=20 with unmodified parent cells, and such immunization affords = no=20 antitumor immunity in vivo (4).=20

As shown in Fig. 1,=20 immunization using DCs pulsed with either B16 tumor extract or B16 RNA = induced=20 B16-specific CTL responses that were statistically = significant=20 compared with animals immunized using DCs pulsed with either = SMA=20 560 tumor extract or SMA 560 RNA, or with animals = injected using=20 PBS (Fig. 1,=20 A and C). Conversely, splenocytes harvested = from groups=20 of animals immunized with SMA 560-pulsed (extract or RNA) DCs = specifically lysed SMA 560 targets (Fig. 1,=20 B and D), further demonstrating the tumor-specific = nature=20 of these responses. As seen in Fig. 1,=20 high levels of nonspecific lysis are observed when animals = are=20 injected with bone marrow-derived DCs pulsed with control = antigen,=20 but not those injected with PBS. High levels of nonspecific = lysis=20 associated with the use of bone marrow-derived DCs have been=20 previously described and were shown to be dependent upon the = presence=20 of sygeneic MHC class II molecules on the immunizing DCs (8)=20 and can be partially overcome by the adherence depletion of=20 antigen-presenting cells before the restimulation of = effectors (19).=20 =20

Fig. 1. Induction of specific lytic activity against = tumor=20 cells by immunization with DC pulsed with tumor extracts or with=20 tumor-derived total RNA. Triplicate C57BL/6 mice were = immunized three=20 times with either DC pulsed with (open circles) B16 = tumor=20 extract or (filled squares) SMA 560 tumor extract = (A and B, respectively), or with DC pulsed with = (open circles) B16 tumor RNA or (filled = squares) SMA=20 560 RNA (C and D, respectively) or with PBS = (filled=20 triangle). 7 d later, splenocytes were isolated and=20 restimulated for 5 d with irradiated B16 cells (A = and=20 C) or SMA 560 cells (B and D). = Cytotoxic=20 activity was then measured by chromium release assay using = the=20 targets indicated in each panel. *(P <0.05) indicate = points=20 of significant differences in lysis when B16-immunized = animals are=20 compared to SMA 560-immunized animals. Error bars indicate = 1 SD=20 of the mean.
[View Larger = Version of=20 this Image (35K GIF file)]=20

B16 Tumor Challenge in CNS After Immunization = Using DCs=20 Pulsed with Tumor Extracts or RNA.=20

Next, experiments were performed to determine whether vaccination = using=20 B16-pulsed DCs generated specific and protective immunity = against B16=20 tumors within the CNS. Groups of C57BL/6 mice received three=20 intraperitoneal vaccinations spaced 1 wk apart and composed = of=20 PBS, B16 extract alone, or DCs pulsed with either B16- or SMA=20 560-derived extracts or RNA. Mice were then challenged in the = brain 1 wk later with 500 viable B16 cells. =

As shown in Fig. 2,=20 immunizations with PBS, B16 extract alone, or DCs pulsed with SMA=20 560 extract or RNA did not protect against CNS challenge = with=20 B16 cells. All animals in these groups succumbed to tumor; = median=20 survival was between 21 and 24 d (Fig. 2,=20 A and B). In contrast, the median survival of = animals=20 undergoing vaccinations of DCs pulsed with B16 extract was=20 significantly prolonged to >80 d (P = =3D 0.0006), with=20 8 of 13 animals surviving when the experiment was = stopped=20 at 80 d (Fig. 2=20 A). Likewise, animals receiving vaccinations of DCs = pulsed=20 with B16 RNA experienced a significant improvement in median = survival=20 to 31 d (P =3D 0.0001), with 4 of = 10 animals=20 surviving when the experiment was stopped at 80 d. No=20 statistical significance was demonstrated in the difference = between=20 groups immunized using DCs pulsed with either B16 extract or = RNA=20 (P =3D 0.29). =20

Fig. 2. Vaccination with DCs pulsed with tumor extract = or tumor=20 RNA protects against CNS challenge with B16 tumor and is equipotent = to vaccination with GM-CSF-producing B16 cells. Vaccination = of=20 C57BL/6 mice was performed a total of three times before = intracranial=20 tumor challenge with B16 cells as described in Materials and = Methods.=20 Mice were evaluated daily until death. Data are = representative=20 of two experiments performed with similar results. The = results=20 are divided in two panels for clarity. The results for the=20 PBS-immunized group (closed squares) and the GM-CSF = modified=20 B16 cell immunized group (open squares) are = represented in=20 both A and B for ease of comparison. Median = days of=20 survival, range, number of animals, and significance compared = to=20 PBS-immunized animals based on log-rank analysis for each = group are=20 as follows: closed squares, PBS: 21, 13-41, n=20 =3D 10; X, DC pulsed with B16 tumor extract: >80, = 17->80,=20 n =3D 13, P =3D 0.0006; closed = diamonds,=20 DCs pulsed with SMA 560 extract; 24, 19-30, = n=20 =3D 11, P =3D 0.50; closed triangles, B16 = extract:=20 21, 14-21, n =3D 5, P =3D 0.45; = open=20 squares, GM-CSF- modified B16 cells: 36, 30->80, = n=20 =3D 6, P =3D 0.022; open triangles, DC = pulsed with=20 B16 RNA: 31, 19->80, n = =3D 10, P=20 =3D 0.0001; open diamonds, DC pulsed with SMA = 560 RNA:=20 21, 15-31, n =3D 9, P =3D 0.40; = open=20 circles, unmodified B16 cells: 23, 18-45, n=20 =3D 5, P =3D 0.39.
[View Larger = Version of=20 this Image (28K GIF file)]=20

It has been convincingly demonstrated that vaccination with tumor = cells=20 genetically engineered to secrete various cytokines = stimulates a=20 potent immune response against tumors outside the CNS (21,=20 24,=20 25).=20 We recently showed that subcutaneous vaccination with B16 = cells that=20 are genetically engineered to produce GM-CSF stimulates a = potent=20 antitumor immune response against B16 tumors located in the = brain and=20 increases the survival of tumor-bearing C57BL/6 mice. It was, = therefore, of interest to compare the efficacy of DC-based=20 immunotherapy with immunization using tumor cells having the = modified=20 GM-CSF gene. As shown in Fig. 2,=20 A and B, mice vaccinated with GM-CSF = gene-modified B16=20 cells had a median survival of 36 d with two of six mice = surviving beyond 80 d, whereas mice vaccinated with B16 = cells=20 alone did not exhibit a survival advantage. Vaccination using = DCs=20 pulsed with cell extracts (Fig. 2=20 A) or tumor RNA (Fig. 2=20 B) was as at least as effective as vaccination using = the B16=20 cells containing modified GM-CSF. Although the median = survival of the=20 group vaccinated using DCs pulsed with B16 tumor extract was = higher=20 than the group vaccinated with GM-CSF-modified B16 cells, the = level=20 of protection achieved with the DC-based vaccines was not=20 statistically greater than that obtained with vaccines using=20 GM-CSF-modified B16 cells (P =3D 0.069). =

Prolonged=20 Survival of Mice Bearing CNS B16 Tumors and Treated Using DCs Pulsed = with B16=20 Tumor Extract.=20

In considering the clinical application of a tumor vaccination = strategy, it=20 is more realistic to treat animals with tumor present at the = time of=20 vaccination. Thus, in the next experiment, 500 B16 cells = were=20 implanted in the brain of naive mice, and these mice were = treated=20 starting 4 d later, at which point vascularized tumor = can be=20 demonstrated histologically (4).=20 Animals received treatments with three intraperitoneal = vaccinations=20 spaced 1 wk apart with PBS, DCs pulsed with SMA=20 560 extract, or DCs pulsed with B16 extract (Fig. 3).=20 Mice in the first two groups had median survivals of = 16 d. Mice=20 treated with the DCs pulsed with B16 extract had a = significantly=20 longer median survival of 26 d (P =3D 0.037), = with two=20 of seven animals surviving at 80 d when the experiment = was=20 stopped. =20

Fig. 3. Treatment of established CNS B16 tumors with = DCs pulsed=20 with tumor extract prolongs survival. Intracranial tumor challenge=20 with B16 cells was performed first and then, starting 4 d = later,=20 C57BL/6 mice were treated with a total of three immunizations = as described in Materials and Methods. Mice were evaluated = daily=20 until death. Median days of survival, range, number of = animals,=20 and significance compared to PBS-immunized animals based on = log-rank=20 analysis for each group are as follows: filled squares, = PBS:=20 16, 15-18, n =3D 6; X, DC pulsed with B16 = tumor=20 extract: 26, 14- >80, n = =3D 7, P=20 =3D 0.037; (filled diamonds), DC pulsed with SMA=20 560 extract: 16, 15-26, n = =3D 7, P=20 =3D 0.57.
[View Larger = Version of=20 this Image (18K GIF file)]=20

Histologic Characterization of CNS Tumors in = Immunized=20 Animals.=20

CNS tumors from triplicate animals immunized only once with PBS or = DCs pulsed=20 with B16 or control EL-4 tumor extract were examined = histologically.=20 Tumors of the group vaccinated with DCs pulsed with B16 = extract=20 demonstrated large areas of hemorrhage and necrosis with an=20 associated heavy inflammatory infiltrate composed of both = mononuclear=20 cells and polymorphonuclear leukocytes. In comparison, no = such areas=20 were observed in either control group (Fig. 4).=20 Outside the immediate peritumoral regions, the brain = parenchyma=20 appeared histologically normal.


Fig. 4. CNS B16 tumors from animals = vaccinated with DC pulsed with B16 tumor extract demonstrate large areas = of=20 necrosis, hemorrhage, and inflammatory infiltrate (b). = No such=20 changes are seen in CNS B16 tumors from animals vaccinated = with DC=20 pulsed with control tumor extract (a) or PBS (not = shown).=20
[View Larger = Versions=20 of these Images (172 + 165K GIF file)]=20

DISCUSSION

In this study, we showed that immunization with DCs pulsed with = either=20 unfractionated tumor extracts or with total tumor RNA elicits = potent=20 immunity against CNS tumors in mice. We showed that this = therapy can=20 be used for both protection against CNS tumor challenge and = in the=20 treatment of established tumors. Furthermore, when compared = directly,=20 B16-pulsed DCs had potency at least equivalent to = GM-CSF-modified B16=20 vaccines.

A number of barriers exist, in practice, to the treatment of human = brain=20 tumors using genetically modified autologous tumor cell = vaccines.=20 These vaccines require the considerable tasks of ex vivo=20 purification, culture, expansion, and transfection of tumor=20 specimens, a difficult undertaking even for tumors outside = the CNS=20 (6,=20 7).=20 DC-based vaccines may overcome some of these problems. First, = human=20 DCs can be generated from peripheral blood; therefore, = supplies are=20 not limited. Secondly, the ex vivo manipulations required to = produce=20 antigen-pulsed DCs are simpler than those required for = generating=20 autologous tumor cell vaccines.

It has been shown that immunizing nonhuman primates and guinea pigs = with=20 human glioblastoma multiforme tissue can induce allergic=20 encephalomyelitis that is lethal (5).=20 Vaccination with unfractionated tumor-derived antigens, such = as those=20 possibly contained in an autologous tumor cell vaccine = derived from=20 the CNS and modified genetically, may lead to potentially = disastrous=20 consequences such as autoimmune encephalitis. This risk may = limit the=20 use of such vaccines to a minority of patients: those = suffering with=20 brain tumors from which highly purified tumor specimens can = not be=20 guaranteed. One approach that may overcome these drawbacks is = to use,=20 as a source of antigen, mRNA from tumor cells. In the case of = brain tumors, an important advantage is that RNA may allow = the use=20 of subtractive hybridization techniques to reduce the=20 concentration of antigens shared between tumor and normal CNS = tissue,=20 lessening the potential for autoimmunity.

By demonstrating that vaccines based on DCs pulsed with tumor = extracts or RNA=20 are active against CNS tumors and are equipotent to cytokine=20 gene-modified vaccines, these studies establish a basis for = future=20 preclinical studies of human DC-based vaccines for treating = brain=20 tumors.

Footnotes

Address correspondence to Dr. Eli Gilboa, Duke University Medical = Center,=20 Department of Surgery, Box 2601, Medical Sciences Research Bldg. = (MSRB), Rm=20 401, Durham, NC, 27710. Phone: 919-684-6465; FAX: = 919-681-7970.

Received for publication 11 June 1997 and in revised form 21 July = 1997.=20

   The authors acknowledge the expert technical assistance = of both=20 Paula Greer and Jie Li. Ann Tamariz provided editorial assistance.=20

This work was supported by grants CA61227 (L.P. Hale), CA11898, and = NS20023=20 (D.D. Bigner) from the National Institutes of Health, and the = Pediatric Brain Tumor Foundation (D.D. Bigner). David = M. Ashley=20 is the recipient of an Australian National Health and Medical = Research Council Neil Hamilton Fairley fellowship.

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S.=20 Liu, Y. Yu, M. Zhang, W. Wang, and X. Cao
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A.=20 Heiser, M. A. Maurice, D. R. Yancey, D. M. Coleman, P. Dahm, and = J.=20 Vieweg
Human Dendritic Cells Transfected with Renal = Tumor RNA=20 Stimulate Polyclonal T-Cell Responses against Antigens Expressed = by=20 Primaryand Metastatic Tumors
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A. B.=20 Pawlowska, S. Hashino, H. McKenna, B. J. Weigel, P. A. Taylor, and = B. R.=20 Blazar
In vitro tumor-pulsed or in vivo Flt3 = ligand-generated=20 dendritic cells provide protection against acute myelogenous = leukemia in=20 nontransplanted or syngeneic bone marrow-transplanted=20 mice
Blood, March 1, 2001; 97(5): 1474 - = 1482.=20
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J. S.=20 Yu, C. J. Wheeler, P. M. Zeltzer, H. Ying, D. N. Finger, P. K. = Lee, W. H.=20 Yong, F. Incardona, R. C. Thompson, M. S. Riedinger, W. Zhang, R. = M.=20 Prins, and K. L. Black
Vaccination of Malignant Glioma = Patients=20 with Peptide-pulsed Dendritic Cells Elicits Systemic Cytotoxicity = and=20 Intracranial T-cell Infiltration
Cancer Res.,=20 February 1, 2001; 61(3): 842 - 847.
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X.-Y.=20 Wang, L. Kazim, E. A. Repasky, and J. R.=20 Subjeck
Characterization of Heat Shock Protein 110 and=20 Glucose-Regulated Protein 170 as Cancer Vaccines and the Effect of = Fever-Range Hyperthermia on Vaccine Activity
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S.=20 Rosic-Kablar, K. Chan, M. D. Reis, I. D. Dub=E9, and M. R.=20 Hough
Induction of tolerance to immunogenic tumor = antigens=20 associated with lymphomagenesis in HOX11 transgenic = mice
PNAS,=20 November 2, 2000; (2000) 240221297.
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RNA-transfected dendritic cells = in=20 cancer immunotherapy
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In Vitro and In Vivo = Induction of a=20 Th Cell Response Toward Peptides of the Melanoma-Associated = Glycoprotein=20 100 Protein Selected by the TEPITOPE Program
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Monocyte-Derived = CD1a+ and=20 CD1a- Dendritic Cell Subsets Differ in Their Cytokine Production = Profiles,=20 Susceptibilities to Transfection, and Capacities to Direct Th Cell = Differentiation
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P. R.=20 Walker, T. Calzascia, V. Schnuriger, N. Scamuffa, P. Saas, N. de = Tribolet,=20 and P.-Y. Dietrich
The Brain Parenchyma Is Permissive = for Full=20 Antitumor CTL Effector Function, Even in the Absence of CD4 T=20 Cells
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Rapid Generation of Potent and Tumor-specific = Cytotoxic=20 T Lymphocytes by Interleukin 18 Using Dendritic Cells and Natural = Killer=20 Cells
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G.=20 Pirtskhalaishvili, G. V. Shurin, A. Gambotto, C. Esche, M. Wahl, = Z. R.=20 Yurkovetsky, P. D. Robbins, and M. R. = Shurin
Transduction of=20 Dendritic Cells with Bcl-xL Increases Their Resistance to Prostate = Cancer-Induced Apoptosis and Antitumor Effect in = Mice
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T.=20 Kikuchi, M. A. S. Moore, and R. G. Crystal
Dendritic = cells=20 modified to express CD40 ligand elicit therapeutic immunity = against=20 preexisting murine tumors
Blood, = July 1, 2000;=20 96(1): 91 - 99.
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T.=20 Fushimi, A. Kojima, M. A.S. Moore, and R. G. = Crystal
Macrophage=20 inflammatory protein 3{alpha} transgene attracts dendritic cells = to=20 established murine tumors and suppresses tumor = growth
J. Clin.=20 Invest., May 15, 2000; 105(10): 1383 - 1393.
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C.=20 Klein, H. Bueler, and R. C. Mulligan
Comparative = Analysis of=20 Genetically Modified Dendritic Cells and Tumor Cells as = Therapeutic Cancer=20 Vaccines
J. Exp. Med., May 15, 2000; = 191(10): 1699 -=20 1708.
[Abstract]=20 [Full = Text]=20 [PDF] =
=20 		3D""
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A. I.=20 Chapoval, K. Tamada, and L. Chen
In vitro growth = inhibition of=20 a broad spectrum of tumor cell lines by activated human dendritic=20 cells
Blood, April 1, 2000; 95(7): 2346 - = 2351.=20
[Abst= ract]=20 [Full=20 Text] [PDF]=20
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D.=20 Boczkowski, S. K. Nair, J.-H. Nam, H. K. Lyerly, and E.=20 Gilboa
Induction of Tumor Immunity and Cytotoxic T = Lymphocyte=20 Responses Using Dendritic Cells Transfected with Messenger RNA = Amplified=20 from Tumor Cells
Cancer Res., = February 1, 2000;=20 60(4): 1028 - 1034.
[Abstract]=20 [Fu= ll=20 Text]
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Y.=20 Nishioka, M. Hirao, P. D. Robbins, M. T. Lotze, and H.=20 Tahara
Induction of Systemic and Therapeutic Antitumor = Immunity=20 Using Intratumoral Injection of Dendritic Cells Genetically = Modified to=20 Express Interleukin 12
Cancer Res., = August 1, 1999;=20 59(16): 4035 - 4041.
[Abstract]=20 [F= ull=20 Text] [PDF]=20
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F.=20 Henry, O. Boisteau, L. Bretaudeau, B. Lieubeau, K. Meflah, and M.=20 Gregoire
Antigen-presenting Cells That Phagocytose = Apoptotic=20 Tumor-derived Cells Are Potent Tumor Vaccines
Cancer = Res.,=20 July 1, 1999; 59(14): 3329 - 3332.
[Abstract]=20 [F= ull=20 Text] [PDF]=20
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D. C.=20 LaTemple, J. T. Abrams, S. Y. Zhang, and U. = Galili
Increased=20 Immunogenicity of Tumor Vaccines Complexed with Anti-Gal: Studies = in=20 Knockout Mice for = {{alpha}}1,3Galactosyltransferase
Cancer=20 Res., July 1, 1999; 59(14): 3417 - 3423.
[Abstract]=20 [F= ull=20 Text] [PDF]=20
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K.=20 Tarte, X.-G. Zhang, E. Legouffe, C. Hertog, M. Mehtali, J.-F. = Rossi, and=20 B. Klein
Induced Expression of B7-1 on Myeloma Cells = Following=20 Retroviral Gene Transfer Results in Tumor-Specific Recognition by=20 Cytotoxic T Cells
J. Immunol., July 1, 1999; = 163(1):=20 514 - 524.
[Abstract= ]=20 [Full=20 Text] [PDF] =
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V.=20 Bronte, D. B. Chappell, E. Apolloni, A. Cabrelle, M. Wang, P. Hwu, = and N.=20 P. Restifo
Unopposed Production of = Granulocyte-Macrophage=20 Colony-Stimulating Factor by Tumors Inhibits CD8+ T Cell Responses = by=20 Dysregulating Antigen-Presenting Cell Maturation
J. = Immunol.,=20 May 15, 1999; 162(10): 5728 - 5737.
[Abstra= ct]=20 [Full=20 Text] [PDF] =
=20 		3D""
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J. H.=20 Sampson
THE PREUSS FOUNDATION SEMINAR ON VACCINE = THERAPY FOR=20 MALIGNANT PRIMARY BRAIN TUMORS February 15-17, 199 8, La Jolla,=20 Calif.
Neuro-oncol, January 1, 1999; 1(1): = 33 - 42.=20
[PDF]<= /A>=20
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X.=20 Cao, W. Zhang, L. He, Z. Xie, S. Ma, Q. Tao, Y. Yu, H. Hamada, and = J.=20 Wang
Lymphotactin Gene-Modified Bone Marrow Dendritic = Cells Act=20 as More Potent Adjuvants for Peptide Delivery to Induce Specific = Antitumor=20 Immunity
J. Immunol., December 1, 1998; = 161(11):=20 6238 - 6244.
[Abstra= ct]=20 [Full=20 Text] [PDF] =
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J.=20 Wang, S. Saffold, X. Cao, J. Krauss, and W. = Chen
Eliciting T=20 Cell Immunity Against Poorly Immunogenic Tumors by Immunization = with=20 Dendritic Cell-Tumor Fusion Vaccines
J. Immunol.,=20 November 15, 1998; 161(10): 5516 - 5524.
[Abstra= ct]=20 [Full=20 Text] [PDF] =
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B.=20 Ludewig, B. Odermatt, S. Landmann, H. Hengartner, and R. M.=20 Zinkernagel
Dendritic Cells Induce Autoimmune Diabetes = and=20 Maintain Disease via De Novo Formation of Local Lymphoid=20 Tissue
J. Exp. Med., October 19, 1998; = 188(8): 1493=20 - 1501.
[Abstract]=20 [Full = Text]=20 [PDF] =
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J.=20 Gong, D. Avigan, D. Chen, Z. Wu, S. Koido, M. Kashiwaba, and D.=20 Kufe
Activation of antitumor cytotoxic T lymphocytes by = fusions=20 of human dendritic cells and breast carcinoma = cells
PNAS,=20 March 14, 2000; 97(6): 2715 - 2718.
[Abstract]=20 [Full = Text]=20 [PDF] =
=20 		3D""
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S.=20 Rosic-Kablar, K. Chan, M. D. Reis, I. D. Dube, and M. R.=20 Hough
Induction of tolerance to immunogenic tumor = antigens=20 associated with lymphomagenesis in HOX11 transgenic = mice
PNAS,=20 November 21, 2000; 97(24): 13300 - 13305.
[Abstract]<= /A>=20 [Full = Text]=20 [PDF] =
=20 		3D""

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=3D=3D=3D=3D=3D=3D=3D=3D=0A= =0A= Emulate Gecko 'XMLHttpRequest()' functionality in IE and Opera. Opera = requires=0A= the Sun Java Runtime Environment .=0A= =0A= by Andrew Gregory=0A= http://www.scss.com.au/family/andrew/webdesign/xmlhttprequest/=0A= =0A= This work is licensed under the Creative Commons Attribution License. To = view a=0A= copy of this license, visit = http://creativecommons.org/licenses/by-sa/2.5/ or=0A= send a letter to Creative Commons, 559 Nathan Abbott Way, Stanford, = California=0A= 94305, USA.=0A= =0A= Attribution: Leave my name and web address in this script intact.=0A= =0A= Not Supported in Opera=0A= ----------------------=0A= * user/password authentication=0A= * responseXML data member=0A= =0A= Not Fully Supported in Opera=0A= ----------------------------=0A= * async requests=0A= * abort()=0A= * getAllResponseHeaders(), getAllResponseHeader(header)=0A= =0A= */=0A= // IE support=0A= if (window.ActiveXObject && !window.XMLHttpRequest) {=0A= window.XMLHttpRequest =3D function() {=0A= var msxmls =3D new Array(=0A= 'Msxml2.XMLHTTP.5.0',=0A= 'Msxml2.XMLHTTP.4.0',=0A= 'Msxml2.XMLHTTP.3.0',=0A= 'Msxml2.XMLHTTP',=0A= 'Microsoft.XMLHTTP');=0A= for (var i =3D 0; i < msxmls.length; i++) {=0A= try {=0A= return new ActiveXObject(msxmls[i]);=0A= } catch (e) {=0A= }=0A= }=0A= return null;=0A= };=0A= }=0A= // Gecko support=0A= /* ;-) */=0A= // Opera support=0A= if (window.opera && !window.XMLHttpRequest) {=0A= window.XMLHttpRequest =3D function() {=0A= this.readyState =3D 0; // = 0=3Duninitialized,1=3Dloading,2=3Dloaded,3=3Dinteractive,4=3Dcomplete=0A= this.status =3D 0; // HTTP status codes=0A= this.statusText =3D '';=0A= this._headers =3D [];=0A= this._aborted =3D false;=0A= this._async =3D true;=0A= this._defaultCharset =3D 'ISO-8859-1';=0A= this._getCharset =3D function() {=0A= var charset =3D _defaultCharset;=0A= var contentType =3D = this.getResponseHeader('Content-type').toUpperCase();=0A= val =3D contentType.indexOf('CHARSET=3D');=0A= if (val !=3D -1) {=0A= charset =3D contentType.substring(val);=0A= }=0A= val =3D charset.indexOf(';');=0A= if (val !=3D -1) {=0A= charset =3D charset.substring(0, val);=0A= }=0A= val =3D charset.indexOf(',');=0A= if (val !=3D -1) {=0A= charset =3D charset.substring(0, val);=0A= }=0A= return charset;=0A= };=0A= this.abort =3D function() {=0A= this._aborted =3D true;=0A= };=0A= this.getAllResponseHeaders =3D function() {=0A= return this.getAllResponseHeader('*');=0A= };=0A= this.getAllResponseHeader =3D function(header) {=0A= var ret =3D '';=0A= for (var i =3D 0; i < this._headers.length; i++) {=0A= if (header =3D=3D '*' || this._headers[i].h =3D=3D header) {=0A= ret +=3D this._headers[i].h + ': ' + this._headers[i].v + '\n';=0A= }=0A= }=0A= return ret;=0A= };=0A= this.getResponseHeader =3D function(header) {=0A= var ret =3D getAllResponseHeader(header);=0A= var i =3D ret.indexOf('\n');=0A= if (i !=3D -1) {=0A= ret =3D ret.substring(0, i);=0A= }=0A= return ret;=0A= };=0A= this.setRequestHeader =3D function(header, value) {=0A= this._headers[this._headers.length] =3D {h:header, v:value};=0A= };=0A= this.open =3D function(method, url, async, user, password) {=0A= this.method =3D method;=0A= this.url =3D url;=0A= this._async =3D true;=0A= this._aborted =3D false;=0A= this._headers =3D [];=0A= if (arguments.length >=3D 3) {=0A= this._async =3D async;=0A= }=0A= if (arguments.length > 3) {=0A= opera.postError('XMLHttpRequest.open() - user/password not = supported');=0A= }=0A= this.readyState =3D 1;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= };=0A= this.send =3D function(data) {=0A= if (!navigator.javaEnabled()) {=0A= alert("XMLHttpRequest.send() - Java must be installed and = enabled.");=0A= return;=0A= }=0A= if (this._async) {=0A= setTimeout(this._sendasync, 0, this, data);=0A= // this is not really asynchronous and won't execute until the = current=0A= // execution context ends=0A= } else {=0A= this._sendsync(data);=0A= }=0A= }=0A= this._sendasync =3D function(req, data) {=0A= if (!req._aborted) {=0A= req._sendsync(data);=0A= }=0A= };=0A= this._sendsync =3D function(data) {=0A= this.readyState =3D 2;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= // open connection=0A= var url =3D new java.net.URL(new = java.net.URL(window.location.href), this.url);=0A= var conn =3D url.openConnection();=0A= for (var i =3D 0; i < this._headers.length; i++) {=0A= conn.setRequestProperty(this._headers[i].h, this._headers[i].v);=0A= }=0A= this._headers =3D [];=0A= if (this.method =3D=3D 'POST') {=0A= // POST data=0A= conn.setDoOutput(true);=0A= var wr =3D new = java.io.OutputStreamWriter(conn.getOutputStream(), this._getCharset());=0A= wr.write(data);=0A= wr.flush();=0A= wr.close();=0A= }=0A= // read response headers=0A= // NOTE: the getHeaderField() methods always return nulls for me :(=0A= var gotContentEncoding =3D false;=0A= var gotContentLength =3D false;=0A= var gotContentType =3D false;=0A= var gotDate =3D false;=0A= var gotExpiration =3D false;=0A= var gotLastModified =3D false;=0A= for (var i =3D 0; ; i++) {=0A= var hdrName =3D conn.getHeaderFieldKey(i);=0A= var hdrValue =3D conn.getHeaderField(i);=0A= if (hdrName =3D=3D null && hdrValue =3D=3D null) {=0A= break;=0A= }=0A= if (hdrName !=3D null) {=0A= this._headers[this._headers.length] =3D {h:hdrName, = v:hdrValue};=0A= switch (hdrName.toLowerCase()) {=0A= case 'content-encoding': gotContentEncoding =3D true; break;=0A= case 'content-length' : gotContentLength =3D true; break;=0A= case 'content-type' : gotContentType =3D true; break;=0A= case 'date' : gotDate =3D true; break;=0A= case 'expires' : gotExpiration =3D true; break;=0A= case 'last-modified' : gotLastModified =3D true; break;=0A= }=0A= }=0A= }=0A= // try to fill in any missing header information=0A= var val;=0A= val =3D conn.getContentEncoding();=0A= if (val !=3D null && !gotContentEncoding) = this._headers[this._headers.length] =3D {h:'Content-encoding', v:val};=0A= val =3D conn.getContentLength();=0A= if (val !=3D -1 && !gotContentLength) = this._headers[this._headers.length] =3D {h:'Content-length', v:val};=0A= val =3D conn.getContentType();=0A= if (val !=3D null && !gotContentType) = this._headers[this._headers.length] =3D {h:'Content-type', v:val};=0A= val =3D conn.getDate();=0A= if (val !=3D 0 && !gotDate) this._headers[this._headers.length] = =3D {h:'Date', v:(new Date(val)).toUTCString()};=0A= val =3D conn.getExpiration();=0A= if (val !=3D 0 && !gotExpiration) = this._headers[this._headers.length] =3D {h:'Expires', v:(new = Date(val)).toUTCString()};=0A= val =3D conn.getLastModified();=0A= if (val !=3D 0 && !gotLastModified) = this._headers[this._headers.length] =3D {h:'Last-modified', v:(new = Date(val)).toUTCString()};=0A= // read response data=0A= var reqdata =3D '';=0A= var stream =3D conn.getInputStream();=0A= if (stream) {=0A= var reader =3D new java.io.BufferedReader(new = java.io.InputStreamReader(stream, this._getCharset()));=0A= var line;=0A= while ((line =3D reader.readLine()) !=3D null) {=0A= if (this.readyState =3D=3D 2) {=0A= this.readyState =3D 3;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= }=0A= reqdata +=3D line + '\n';=0A= }=0A= reader.close();=0A= this.status =3D 200;=0A= this.statusText =3D 'OK';=0A= this.responseText =3D reqdata;=0A= this.readyState =3D 4;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= if (this.onload) {=0A= this.onload();=0A= }=0A= } else {=0A= // error=0A= this.status =3D 404;=0A= this.statusText =3D 'Not Found';=0A= this.responseText =3D '';=0A= this.readyState =3D 4;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= if (this.onerror) {=0A= this.onerror();=0A= }=0A= }=0A= };=0A= };=0A= }=0A= // ActiveXObject emulation=0A= if (!window.ActiveXObject && window.XMLHttpRequest) {=0A= window.ActiveXObject =3D function(type) {=0A= switch (type.toLowerCase()) {=0A= case 'microsoft.xmlhttp':=0A= case 'msxml2.xmlhttp':=0A= case 'msxml2.xmlhttp.3.0':=0A= case 'msxml2.xmlhttp.4.0':=0A= case 'msxml2.xmlhttp.5.0':=0A= return new XMLHttpRequest();=0A= }=0A= return null;=0A= };=0A= }=0A= ------=_NextPart_000_0001_01C709AB.0016CBA0 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://www.jem.org/javascript/ajax/utility.js /************************************************************************= *****=0A= * javascript/ajax/utility.js=0A= *=0A= * Utility functions for working with XMLHttpRequest data.=0A= *=0A= * Copyright 2006 Board of Trustees of the Leland Stanford Junior = University.=0A= = *************************************************************************= ***/=0A= =0A= /*=0A= * Copy XML nodes into an HTMLElement. This effectively=0A= * clones XML markup which uses XHTML naming conventions=0A= * into an HTML DOM.=0A= */=0A= function copy_xml_to_html(src, dst) {=0A= if (src.nodeType =3D=3D 1) { /* Node.ELEMENT_NODE */=0A= var e =3D document.createElement(src.nodeName);=0A= for (var i =3D 0; i < src.childNodes.length; i++) {=0A= copy_xml_to_html(src.childNodes[i], e);=0A= }=0A= for (var i =3D 0; i < src.attributes.length; i++) {=0A= var n =3D src.attributes[i].name;=0A= var v =3D unescape_xml_string(src.attributes[i].value); =0A= e.setAttribute(n, v);=0A= if (n =3D=3D "class") {=0A= e.className =3D v;=0A= }=0A= else if (n =3D=3D "style") {=0A= set_css_style(v, e, "");=0A= }=0A= }=0A= dst.appendChild(e);=0A= }=0A= else if (src.nodeType =3D=3D 3) { /* Node.TEXT_NODE */=0A= dst.appendChild(document.createTextNode(src.nodeValue));=0A= }=0A= }=0A= =0A= /* =0A= * It is unclear that this is the right thing to be calling=0A= * from copy_xml_to_html, but it appears that Safari decides=0A= * to convert & to the NCR #, and then encodes that=0A= * NCR to &%26%2338;. So, I'm going to treat the DOM Attr=0A= * value as a plain string, and run our XML string input=0A= * through the decoding routine below.=0A= */=0A= function unescape_xml_string(s) {=0A= return s.replace(/'/g, "'")=0A= .replace(/'/g, "'")=0A= .replace(/"/g, "\"")=0A= .replace(/"/g, "\"")=0A= .replace(/>/g, ">")=0A= .replace(/>/g, ">")=0A= .replace(/</g, "<")=0A= .replace(/</g, "<")=0A= .replace(/&/g, "&")=0A= .replace(/&/g, "&");=0A= }=0A= =0A= /*=0A= * Parse set of CSS rules and apply them to an element.=0A= * This is quite horrifying, but I'm unable to determine=0A= * how else to handle this with IE 6. FireFox and other=0A= * sane browsers let you simply set the style attribute=0A= * or use e.style.setProperty(rule, value, priority),=0A= * IE 6 appears to have neither of these capabilities..=0A= */=0A= function set_css_style(css, e, priority) {=0A= var rules =3D css.split(";");=0A= for (var i =3D 0; i < rules.length; i++) {=0A= var nvpair =3D rules[i].split(":");=0A= if (nvpair.length =3D=3D 2) {=0A= try {=0A= var name =3D nvpair[0]; /* style attribute */=0A= var value =3D nvpair[1]; /* attribute value */=0A= =0A= /*=0A= * For each possible style attribute, set the=0A= * appropriate style property in the element.=0A= */=0A= if (name =3D=3D "background") {=0A= e.style.background =3D value;=0A= }=0A= else if (name =3D=3D "background-attachment") {=0A= e.style.backgroundAttachment =3D value;=0A= }=0A= else if (name =3D=3D "background-color") {=0A= e.style.backgroundColor =3D value;=0A= }=0A= else if (name =3D=3D "background-image") {=0A= e.style.backgroundImage =3D value;=0A= }=0A= else if (name =3D=3D "background-position") {=0A= e.style.backgroundPosition =3D value;=0A= }=0A= else if (name =3D=3D "background-position-x") {=0A= e.style.backgroundPositionX =3D value;=0A= }=0A= else if (name =3D=3D "background-position-y") {=0A= e.style.backgroundPositionY =3D value;=0A= }=0A= else if (name =3D=3D "background-repeat") {=0A= e.style.backgroundRepeat =3D value;=0A= }=0A= else if (name =3D=3D "behavior") {=0A= e.style.behavior =3D value;=0A= }=0A= else if (name =3D=3D "border") {=0A= e.style.border =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom") {=0A= e.style.borderBottom =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-color") {=0A= e.style.borderBottomColor =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-style") {=0A= e.style.borderBottomStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-width") {=0A= e.style.borderBottomWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-collapse") {=0A= e.style.borderCollapse =3D value;=0A= }=0A= else if (name =3D=3D "border-color") {=0A= e.style.borderColor =3D value;=0A= }=0A= else if (name =3D=3D "border-left") {=0A= e.style.borderLeft =3D value;=0A= }=0A= else if (name =3D=3D "border-left-color") {=0A= e.style.borderLeftColor =3D value;=0A= }=0A= else if (name =3D=3D "border-left-style") {=0A= e.style.borderLeftStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-left-width") {=0A= e.style.borderLeftWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-right") {=0A= e.style.borderRight =3D value;=0A= }=0A= else if (name =3D=3D "border-right-color") {=0A= e.style.borderRightColor =3D value;=0A= }=0A= else if (name =3D=3D "border-right-style") {=0A= e.style.borderRightStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-right-width") {=0A= e.style.borderRightWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-style") {=0A= e.style.borderStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-top") {=0A= e.style.borderTop =3D value;=0A= }=0A= else if (name =3D=3D "border-top-color") {=0A= e.style.borderTopColor =3D value;=0A= }=0A= else if (name =3D=3D "border-top-style") {=0A= e.style.borderTopStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-top-width") {=0A= e.style.borderTopWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-width") {=0A= e.style.borderWidth =3D value;=0A= }=0A= else if (name =3D=3D "bottom") {=0A= e.style.bottom =3D value;=0A= }=0A= else if (name =3D=3D "clear") {=0A= e.style.clear =3D value;=0A= }=0A= else if (name =3D=3D "clip") {=0A= e.style.clip =3D value;=0A= }=0A= else if (name =3D=3D "color") {=0A= e.style.color =3D value;=0A= }=0A= else if (name =3D=3D "cssText") {=0A= e.style.Sets =3D value;=0A= }=0A= else if (name =3D=3D "cursor") {=0A= e.style.cursor =3D value;=0A= }=0A= else if (name =3D=3D "direction") {=0A= e.style.direction =3D value;=0A= }=0A= else if (name =3D=3D "display") {=0A= e.style.display =3D value;=0A= }=0A= else if (name =3D=3D "font") {=0A= e.style.font =3D value;=0A= }=0A= else if (name =3D=3D "font-family") {=0A= e.style.fontFamily =3D value;=0A= }=0A= else if (name =3D=3D "font-size") {=0A= e.style.fontSize =3D value;=0A= }=0A= else if (name =3D=3D "font-style") {=0A= e.style.fontStyle =3D value;=0A= }=0A= else if (name =3D=3D "font-variant") {=0A= e.style.fontVariant =3D value;=0A= }=0A= else if (name =3D=3D "font-weight") {=0A= e.style.fontWeight =3D value;=0A= }=0A= else if (name =3D=3D "height") {=0A= e.style.height =3D value;=0A= }=0A= else if (name =3D=3D "ime-mode") {=0A= e.style.imeMode =3D value;=0A= }=0A= else if (name =3D=3D "layout-flow") {=0A= e.style.layoutFlow =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid") {=0A= e.style.layoutGrid =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-char") {=0A= e.style.layoutGridChar =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-line") {=0A= e.style.layoutGridLine =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-mode") {=0A= e.style.layoutGridMode =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-type") {=0A= e.style.layoutGridType =3D value;=0A= }=0A= else if (name =3D=3D "left") {=0A= e.style.left =3D value;=0A= }=0A= else if (name =3D=3D "letter-spacing") {=0A= e.style.letterSpacing =3D value;=0A= }=0A= else if (name =3D=3D "line-break") {=0A= e.style.lineBreak =3D value;=0A= }=0A= else if (name =3D=3D "line-height") {=0A= e.style.lineHeight =3D value;=0A= }=0A= else if (name =3D=3D "list-style") {=0A= e.style.listStyle =3D value;=0A= }=0A= else if (name =3D=3D "list-style-image") {=0A= e.style.listStyleImage =3D value;=0A= }=0A= else if (name =3D=3D "list-style-position") {=0A= e.style.listStylePosition =3D value;=0A= }=0A= else if (name =3D=3D "list-style-type") {=0A= e.style.listStyleType =3D value;=0A= }=0A= else if (name =3D=3D "margin") {=0A= e.style.margin =3D value;=0A= }=0A= else if (name =3D=3D "margin-bottom") {=0A= e.style.marginBottom =3D value;=0A= }=0A= else if (name =3D=3D "margin-left") {=0A= e.style.marginLeft =3D value;=0A= }=0A= else if (name =3D=3D "margin-right") {=0A= e.style.marginRight =3D value;=0A= }=0A= else if (name =3D=3D "margin-top") {=0A= e.style.marginTop =3D value;=0A= }=0A= else if (name =3D=3D "min-height") {=0A= e.style.minHeight =3D value;=0A= }=0A= else if (name =3D=3D "overflow") {=0A= e.style.overflow =3D value;=0A= }=0A= else if (name =3D=3D "overflow-x") {=0A= e.style.overflowX =3D value;=0A= }=0A= else if (name =3D=3D "overflow-y") {=0A= e.style.overflowY =3D value;=0A= }=0A= else if (name =3D=3D "padding") {=0A= e.style.padding =3D value;=0A= }=0A= else if (name =3D=3D "padding-bottom") {=0A= e.style.paddingBottom =3D value;=0A= }=0A= else if (name =3D=3D "padding-left") {=0A= e.style.paddingLeft =3D value;=0A= }=0A= else if (name =3D=3D "padding-right") {=0A= e.style.paddingRight =3D value;=0A= }=0A= else if (name =3D=3D "padding-top") {=0A= e.style.paddingTop =3D value;=0A= }=0A= else if (name =3D=3D "page-break-after") {=0A= e.style.pageBreakAfter =3D value;=0A= }=0A= else if (name =3D=3D "page-break-before") {=0A= e.style.pageBreakBefore =3D value;=0A= }=0A= else if (name =3D=3D "pixelBottom") {=0A= e.style.pixelBottom =3D value;=0A= }=0A= else if (name =3D=3D "pixelHeight") {=0A= e.style.pixelHeight =3D value;=0A= }=0A= else if (name =3D=3D "pixelLeft") {=0A= e.style.pixelLeft =3D value;=0A= }=0A= else if (name =3D=3D "pixelRight") {=0A= e.style.pixelRight =3D value;=0A= }=0A= else if (name =3D=3D "pixelTop") {=0A= e.style.pixelTop =3D value;=0A= }=0A= else if (name =3D=3D "pixelWidth") {=0A= e.style.pixelWidth =3D value;=0A= }=0A= else if (name =3D=3D "posBottom") {=0A= e.style.posBottom =3D value;=0A= }=0A= else if (name =3D=3D "posHeight") {=0A= e.style.posHeight =3D value;=0A= }=0A= else if (name =3D=3D "position") {=0A= e.style.position =3D value;=0A= }=0A= else if (name =3D=3D "posLeft") {=0A= e.style.posLeft =3D value;=0A= }=0A= else if (name =3D=3D "posRight") {=0A= e.style.posRight =3D value;=0A= }=0A= else if (name =3D=3D "posTop") {=0A= 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{=0A= e.style.scrollbarHighlightColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-shadow-color") {=0A= e.style.scrollbarShadowColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-track-color") {=0A= e.style.scrollbarTrackColor =3D value;=0A= }=0A= else if (name =3D=3D "float") {=0A= e.style.styleFloat =3D value;=0A= }=0A= else if (name =3D=3D "table-layout") {=0A= e.style.tableLayout =3D value;=0A= }=0A= else if (name =3D=3D "text-align") {=0A= e.style.textAlign =3D value;=0A= }=0A= else if (name =3D=3D "text-align-last") {=0A= e.style.textAlignLast =3D value;=0A= }=0A= else if (name =3D=3D "text-autospace") {=0A= e.style.textAutospace =3D value;=0A= }=0A= else if (name =3D=3D "text-decoration") {=0A= e.style.textDecoration =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationBlink") {=0A= e.style.textDecorationBlink =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationLineThrough") {=0A= e.style.textDecorationLineThrough =3D value;=0A= }=0A= else if (name =3D=3D 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