ࡱ>  M bjbj== 5WW lXXXl8$TxTl;" D'$:::::::$= ?=:AXK  KK=:0~;000K~"X:0K:00'5:{9,rX: Is+l/9 :;0;9RT@/T@:0ll ENDOMORPHIN 1 ACTIVATES NOS 2 ACTIVITY AND DOWNREGULATES NOS 2 mRNA EXPRESSION A. `ARI, T. BALOG*, S. SOBO ANEC AND T. MAROTTI Division of Molecular Medicine, Rudjer Boakovi Institute, Bijeni ka 54, 10000 Zagreb, Croatia *Correspondence to: T. Balog, Division of Molecular Medicine, Rudjer Boakovi Institute, Bijeni ka 54, 10000 Zagreb, Croatia. Tel.: +385-1-4561-172; fax.: +385-1-4561-010. E-mail address: balog@irb.hr Section Editor Abbreviations: 1400W, N-(3-[Aminomethyl]benzyl)acetamidine; EM, endomorphin; FCS, fetal calf serum; ICI-174,864, N,N-diallyl-Tyr-Aib-Aib-Phe-Leu; L-NMMA, NG-Monomethyl-L-arginine acetate salt; LPS, lipopolysaccharide; NO, nitric oxide; NOS, nitric oxide synthase; RT-PCR, reverse transcription-polymerase chain reaction; -FNA, -Funaltrexamine hydrochloride ABSTRACT Endomorphins 1 and 2 are newly discovered opioid tetrapeptides whose structure is more resistant to enzymatic degradation than of other opioid peptides. Endomorphins 1 and 2 are considered as endogenous ligands with a high affinity for receptors. A number of studies have shown that opioid peptides per se can induce release of nitric oxide from rodent and human immune cells. Endomorphins seemed to be involved in the process of vasodilatation by stimulating release of nitric oxide. In our study we stimulated in vitro J774 macrophages with different concentrations of endomorphin 1 or 2 for measuring nitric oxide release and nitric oxide synthase 2 mRNA expression. Results showed that 48 h incubation did not enhance nitric oxide release when measured with the Griess method. On the other hand, using real-time amperometric detection of nitric oxide release shortly after challenge with endomorphins, we showed that only 10-6 M endomorphin 1 was able to stimulate nitric oxide release from a J774 macrophage cell line by activation of NOS 2 izoenzyme. The peak release was 1000-1500 s after stimulation and was in the range of nitric oxide release stimulated with 10 g/ml lipopolysaccharide. In contrast to this, endomorphin 2 failed to induce nitric oxide release in all tested concentrations. Using a specific inhibitor of nitric oxide synthase 2 (1400W) we eliminated the stimulatory effect of endomorphin 1 on nitric oxide release. The expression of mRNA for nitric oxide synthase 2 in J774 macrophages, after 30 min incubation with either lipopolysaccharide or 10-6 M endomorphin 1 was not upregulated. As expected, lipopolysaccharide induced de novo nitric oxide synthase 2 transcription within 4 h. At the same time, in contrast to lipopolysaccharide, mRNA expression of cells treated with endomorphin 1 was downregulated. Since a -opioid receptor specific antagonist -Funaltrexamine hydrochloride inhibited nitric oxide release from endomorphin 1 treated cells, the effect seemed to be -opioid receptor mediated. Key words: nitric oxide, -opioid receptor, amperometric detection, INTRODUCTION Endomorphins 1 and 2 are newly discovered opioid tetrapeptides (Zadina et al., 1997) with a specific structure which is more resistant to enzymatic degradation than other opioid peptides such as enkephalins, endorphins or dynorphines (Peter et al., 1999). In contrast to enkephalins and endorphins which posses low affinity and selectivity to -opioid receptors, endomorphins 1 and 2 are considered as endogenous ligands with a high affinity for receptors (Zadina et al., 1997). While endomorphin 1 is more distributed throughout the brain, endomorphin 2 prevails in the spinal cord (Martin-Shild et al., 1999). Both tetrapeptides are detected by radioimmunoassay and high performance liquid chromatography in cells of the immune system of rats and humans (Jessop et al., 2000). During infection several authors have noticed an increase of opioids in peripheral blood as well as on the site of an inflammatory reaction (Kanjhan, 1995; Stein, 1995). That is to say, under inflammatory conditions various types of immune cells are able to produce opioid peptides in culture (Sharp and Linner, 1993) and in situ (Stein et al., 1990). Immune cells from peripheral blood, splenic lymphocytes and macrophages express mRNA for beta-endorphin and other proopiomelanocortin derived peptides (Peterson et al., 1993a,b; Eisenstein et al., 1996). Endogenous opioids also modulate the effect of cytokines on CNS (Hori et al., 1991; Xin and Blatteis, 1991), while on the other hand cytokines may alter the neural effects of opioids (Jeanjean et al., 1995). A number of studies have shown that opioid peptides, like enkephalins also induce a release of nitric oxide (NO) from rodent and human immune cells (Marotti et al., 1998; Balog et al., 2001; Bengoechea-Alonso et al., 2003; Vuji et al., 2004). Champion et al. (1998) demonstrated that endomorphins cause vasodilatation in rats by stimulating NO release from the endothelium. NO is a molecule involved in the immune, cardiovascular and nervous systems (Moncada et al., 1988; Magazine et al., 1996). NO is produced from aminoacid L-arginine by an enzyme called nitric oxide synthase (NOS). NOS persists in cells in three different forms: the endothelial (NOS 3), neuronal (NOS 1) and inducible (NOS 2) form. The first two forms are constitutively expressed in cells and are Ca2+ dependent as well, while NOS 2 is inducible and Ca2+ independent (Moncada et al., 1991; Moncada and Higgs, 1993). Constitutive forms of NOS (endothelial and neuronal) release NO in nM concentration, while the inducible form (NOS 2) releases NO in M range (Stefano et al., 2000). NO is primarily a vascular, immune and neural signalling molecule, being highly effective as general antibacterial and antiviral agents with the ability to downregulate proinflammatory events (Kubes and Granger, 1992; Niu et al., 1995; Stefano et al., 1998a). So far, several studies have shown that inducible NOS 2 can be induced with various signal molecules and proinflammatory cytokines in vivo (Moncada et al., 1991; Stefano et al., 1998b). The aim of this paper was to demonstrate the ability of the J774 macrophages cell line to release NO upon stimulation with endomorphins 1 and 2, using real-time amperometric detection of NO release in short-term experiments. Using specific inhibitors of various types of NOS and opioid receptor antagonists, we aimed to identify the type of NOS involved, and also the opioid receptor through which NO release was mediated. EXPERIMENTAL PROCEDURES Reagents Endomorphins 1 and 2, lipopolysaccharide (LPS), RPMI 1640 without phenol red, sulphanilamide, N-(1-naphtyl)ethylendiamine dihydrochloride were all purchased from Sigma, St Louis, USA. Phenol red-free medium was supplemented with antibiotic/antimycotic (Sigma St. Louis USA) and used only for Griess assay. For all other assays phenol-red RPMI 1640 medium (Institute of Immunology, Inc., Zagreb, Croatia) was used. Preparation of cell cultures A murine macrophage cell line, J774, was obtained from Pliva, Zagreb, Croatia. Cells were cultured in 75 cm2 plactic flasks in phenol-red RPMI medium with 10% fetal calf serum (FCS, Invitrogen, Life Technologies, Carlsbad, CA, USA ) and passed three times a week until they reached confluency. The cells were maintained in a humidified atmosphere of 95% air 5% CO2 at 37C. Treatment conditions The J774 cell suspension was seeded into either 24-well flat-bottom plates at 1x106 cells/ml/well for both nitrite assay (Griess) and reverse transcription-polymerase chain reaction (RT-PCR) experiments, or 2x106 cells/2 ml/well for real-time amperometric measurements. The J774 cultures were challenged with various concentrations of endomorphin 1, endomorphin 2 (10-6, 10-7, 10-8, 10-9, 1010 and 1011M) or LPS (10 g/ml). The cells incubated in the culture medium alone were used as a control. In addition, the cells were pretreated with either the -selective antagonist -Funaltrexamine hydrochloride (-FNA, 10-4 M, Tocris Bioscience, USA) (Vega and Soto, 2003), the -selective antagonist N,N-diallyl-Tyr-Aib-Aib-Phe-Leu (ICI-174,864, 10-4 M, Research Biochemicals International USA) (Zhang et al., 1992) or with the NOS 2 selective inhibitor N-(3-[Aminomethyl]benzyl)acetamidine (1400W, 10-4 M, Sigma, St. Louis, USA) (Kalinowski et al., 2002; Korhonen et al., 2002) for 30 min before the addition of LPS or endomorphins. Assay for nitrite concentration Following stimulation for 48 h, with endomorphin 1, endomorphin 2 or LPS, the nitrite concentration was measured as an indicator of NO production, according to the Griess reaction (Green et al., 1982). Briefly, 800 l of cell culture supernatant was incubated with an equal volume of Griess reagent (sulfanilamide and N-(1-naphtyl)ethylendiamine dihydrochloride dissolved in 2.5% H3PO4 as 1% and 0.5% solution, respectively, and mixed in the volume ratio 1:1 immediately before use) for 15 min at room temperature, in the dark. The optical density was measured spectrophotometrically at 540 nm. Nitrite concentration was determined using sodium nitrite as standard (10-100 M). Real-time detection of NO A real-time NO release was measured using the NO detection system ISO-NO MK II with 2 mm diameter NO sensor (World Precision Instruments, Inc., Sarasota, USA). NO diffuses through a gas-permeable hydrophobic membrane covering the sensor and is oxidized at the working (Pt) electrode, resulting in an electrical current. This redox current is proportional to the NO concentration in the sample outside the membrane, and is continuously monitored with DUO 18 data recording system connected to a PC. Electrode calibration was performed according to the manufacturers instructions. Different volumes of sodium nitrite standard (NaNO2, 100 M) were used as a generator of NO. A calibration curve was obtained by measurement of the current, generated by the addition of a known amount of NaNO2 to produce a known amount of NO. The relationship between the NO concentration and the output current of the electrode was always linear. Due to the high sensitivity of the method, a noise of maximum 50 pA amplitude was accepted as background. For experiments, the tip of the probe was inserted vertically into the 24-well containing 2x106 adherent cells per 2 ml RPMI medium with 10% FCS. Before cell culture treatment, the sensor probe was allowed to stabilize to achieve NO basal level. Production of NO was assayed immediately after challenging cells with various concentrations of endomorphin 1, endomorphin 2 (10-6 to 10-11 M) or LPS (10 g/ml). For inhibition studies the effects of endomorphins and LPS were evaluated after 30 min pre-incubation with a -selective opioid receptor antagonist -FNA (10-4 M) or NOS inhibitors NG-Monomethyl-L-arginine acetate salt (L-NMMA 10-4 M, Sigma St Louis USA) and 1400W (10-4 M). During the experiment, each culture well was kept on a thermostated block at 37C. RNA isolation At the end of stimulation, the cell monolayer was washed twice with 1 ml of ice-cold phosphate buffered saline to eliminate excess FCS and nonadherent cells. Total RNA was extracted using TRIzol reagent (Invitrogen, Carlsbad, CA, USA) according to the manufacturers instructions. Briefly, the cells were lysed directly in the plate wells using 0.5 ml TRIzol reagent per well. After chloroform extraction, the total RNA was recovered from the aqueous phase and precipitated with an equal volume of isopropanol (Kemika, Zagreb, Croatia). Total RNA pellet was dissolved in 20 l of diethylpyrocarbonate (DEPC)-treated water (Sigma, St. Louis, SAD) after a brief wash with 75% ethanol (Kemika, Zagreb, Croatia). The concentration and integrity of RNA was checked by measuring absorbance at 260 nm followed by electrophoresis on 1% agarose gel (Chomczynski and Sacchi, 1987). RT-PCR Of the total RNA, 1 g was used for reverse transcription reaction in a final volume of 20 l with SuperScriptTM II RNase H Reverse Transcriptase (Invitrogen, Life Technologies, Carlsbad, CA, USA). DNase I pretreatment was used to eliminate contaminating DNA (Invitrogen, Life Technologies, Carlsbad, CA, USA). To amplify specific gene product the following primer sequences (5`3`) were used: NOS 2: 5ATG CCC GAT GGC ACC ATC AGA 3(forward), 5 CAC TTC CTC CAG GAT GTT GTA 3(reverse) and -actin: 5 GTG GGC CGC TCT AGG CAC CAA 3(forward), 5 CTC TTT GAT GTC ACG CAC GAT TTC 3(reverse). These primer sets discriminated between cDNA and genomic DNA due to intronic sequences and yielded PCR products of 372 and 540 bp for NOS-2 and -actin, respectively. Template cDNA (5 l) was amplified in a final volume of 25 l reaction mixture containing 0.5 IU HotMasterTM Taq DNA polymerase (Eppendorf, Hamburg, Germany), 0.25 mM dNTP Mix, 0.2 mM Mg2+ in 10x Hot Master Taq Buffer and 0.25 M concentration of each primer. After initial template denaturation at 94C for 2 min, amplifications were performed in a Mastercycler personal 5332 (Eppendorf, Germany), under the following cycle profile: 94C for 35 s, 60C for 2 min and 72C for 2 min (NOS 2) or 94C for 20 s, 55C for 20 s and 70C for 40 s (-actin). A final step of 10 min at 72C was performed to complete the PCR reaction. For each primer pair, control experiments were performed to determine the range of cycles in which a given amount of cDNA would be amplified in a linear fashion: 29 cycles for NOS 2, or 33 cycles for -actin. PCR products were subjected to electrophoresis on 1.5% agarose gels, stained with ethidium bromide and imaged on a UV transilluminator (Image Master VDF, Pharmacia Biotech). Semiquantitative analysis was performed by measuring the relative optical brightness of each band. The data were normalized to transcript levels for the constitutive expressed -actin gene, employed as an internal control. Statistical analysis Data were analyzed using the statistical package Statgraphic plus 4.0 for Windows, and reported as mean SEM. Statistical comparison between groups was performed using Analysis of Variance (ANOVA). Post-hoc comparisons were performed using LSD test. P- value less than 0.05 was considered significant. RESULTS The effect of endomorphins on nitrite production To investigate the long-term effect of endomorphins on NO production, J774 macrophages were challenged with endomorphins (10-6 to 10-10 M) or LPS (10 g/ml) for 48 h. The accumulated nitrite was measured in the culture media using Griess reagent. Incubation of cells with either endomorphin 1 (Fig. 1A) or endomorphin 2 (Fig. 1B) alone did not change NO release. LPS (10 g/ml) caused a significant increase (p<0.001) in NO production after a 48 h incubation period. The effect of endomorphins on NO production To evaluate the effect of endomorphins on NO production we used the direct real time amperometric detection method. J774 macrophages were exposed to a concentration gradient of endomorphins (10-11 to 10-6 M) (Fig. 2). Endomorphin 2 had no effect on NO production while endomorphin 1 significantly increased (p=0.001) NO only at 10-6 M concentration (Fig. 3). The maximum NO release (39017 nM compared with medium alone 7846 nM) was reached within 1400 sec and did not decline to baseline for at least 1 hour. Endomorphin 1 increased NO release was comparable to LPS (10 g/ml) induced NO release (40556 nM) at the same time (1400 s). The LPS induced NO release was completely eliminated after 30 min preincubation with NOS inhibitor L-NMMA (10-4 M) (Fig. 3) demonstrating that the source of NO was due to enzymatic generation by NOS. In an attempt to identify which type of NOS is responsible for NO production induced by 10-6 M endomorphin 1, we used selective inhibitor for NOS 2 isoform 1400W. As shown in Fig. 4, 1400W caused a complete inhibition of endomorphin 1 induced NO formation, implying that the release of NO upon endomorphin 1 stimulation was due to NOS 2 activity. NOS 2 inhibitor 1400W alone, no effect on NO release from J774 macrophages was noticed (data not shown). Since our results showed NO release within 30 min after stimulation, while NOS 2 synthesis usually occurs 2-4 h after LPS or cytokine treatment, RT-PCR experiments were performed to examine the effect on the transcription of NOS 2. As expected, LPS did not induce NOS 2 transcription 30 min post-LPS (Fig. 5A), but within a 4 h incubation period NOS 2 transcription was significantly upregulated (p<0.05) (Fig. 5B). Surprisingly, a significant amount of NOS 2 mRNA was present in naive unstimulated macrophages after 30 min incubation with medium alone (control). Similarly to LPS, after 30 min incubation of J774 with endomorphin 1 no significant difference in mRNA expression was detected (Fig. 5A). In contrast to LPS, endomorphin 1 downregulated NOS 2 expression after 4 h (p<0.05) (Fig. 5B). A possible involvement of the -opioid receptor in endomorphin 1 mediated NO-release was investigated using -opioid receptor selective antagonist -FNA. NO release from endomorphin 1 treated J774 macrophages was partially blocked (46%) using -FNA (10-4 M) pretreatment (Fig. 6). -FNA (10-4 M) alone had no effect on NO release from J774 macrophages (data not shown). DISCUSSION It is known that opioid peptides affect reactive oxygen species release in different cell systems. Marotti et al. (1990) have shown a donor-dependent upregulation of superoxide anion release from human neutrophils by opioid peptides, while verko et al. (2002) demonstrated met-enkephalin modulated lipid peroxidation in the liver and thymus of mice. Met-enkephalin also modulates NO release from peritoneal macrophages in vivo and ex vivo (Balog et al., 2001; Marotti et al., 1998). LPS and interferon-stimulated NO release from J774 macrophage cell line can be decreased by dynorphin in vitro (Gabrilovac et al., 2003), while Vuji et al. (2004) have demonstrated that met-enkephalin synergizes with LPS for NO release from rat peritoneal macrophages. Recently, Lin et al. (2003) provided evidence that endomorphins can protect human mononuclear blood cells from oxidative damage. In our study, the J774 macrophage cell line stimulated in vitro with different concentrations of endomorphin 1 or endomorphin 2 during 48 hours did not enhance NO release when measured with the Griess method. This might be due to much lower sensitivity of the Griess assay as compared to amperometric measurement of NO (M versus nM). That is to say, since the effects of short-term incubation are most probably related to direct -opioid receptor related effects, a long incubation period might be cytokine release related. These effects of opioid peptides can be observed in the case of in vivo rather than in vitro opioid peptide treatment, as we demonstrated in our study (Balog et al., 2001). Results have shown that only 10-6 M endomorphin 1 stimulated NO release from the J774 macrophage cell line. The peak of release was 1000-1500 s after stimulation and was in the range of NO release stimulated with 10 g/ml LPS. In contrast to this, endomorphin 2 in all tested concentrations failed to induce NO release from J774 mouse macrophages. The difference might be related to the fact that endomorphin 2 inhibits release of TNF alpha from macrophages, a cytokine strongly directed to NO release (Azuma and Ohura, 2002). Another possible explanation of the difference between the effective endomorphin 1 and the non-effective endomorphin 2 is the fact that endomorphin 2 is more susceptible to enzymatic degradation by carboxypeptidase Y than endomorphin 1 (Peter et al., 1999). This is, however, unlikely since according to our results the stimulation of NO release from J774 macrophages by endomorphin 1 seems to be -opioid receptor mediated. Thus, lack of carboxyterminal aminoacid probably would not affect our findings. In contrast to that, more impact on the effect of endomorphin 1 or endomorphin 2 in our study could be caused by the combination of dipeptydil peptidase IV and aminopeptidase N which affects the aminoterminal side of the molecule and splitting of aminothermus tyrosine inactivates opioid peptides (Roques et al., 1993). The specificity of the measured effect of endomorphin 1 on NO release was demonstrated using L-NMMA which completely inhibited NOS activity. According to most of the data in the literature, release of NO from J774 macrophage cell line after short-term stimulation should be attributed to constitutive NOS isoenzyme. However, since we inhibited the stimulatory effect of endomorphin 1 on NO release by 1400W, a specific NOS 2 inhibitor in our study NO release was due to activation of NOS 2 izoenzyme. Also, we detected mRNA for NOS 2 in unstimulated J774 macrophages. Since de novo NOS 2 synthesis usually occurs 2-4 h after LPS or cytokine treatment, RT-PCR experiments were performed in order to exclude the possibility that observed NO release after endomorphin 1 treatment was due to de novo induction of NOS 2 mRNA expression. After 30 min incubation of J774 macrophages with either LPS or 10-6 M endomorphin 1 we did not notice upregulated mRNA expression for NOS 2. As expected, LPS-induced de novo NOS 2 transcription became evident within 4 h, but not 30 min post-LPS. Contrary to LPS, mRNA expression of cells treated with endomorphin 1 for 4 h was downregulated. However, 4 h after stimulation LPS up-regulated mRNA for NO, which was followed by a second peak (5-6 h after stimulation) in NO release measured by amperometer. No second peak observed with endomorphin 1 paralleled down-regulation in mRNA observed after 4 h stimulation with endomorphin 1 (data not shown). Besides, the discrepancy between LPS and endomorphin 1 effect might be the result of a different pathway of macrophage stimulation which, in the case of LPS, involves CD14 activation and for endomorphin 1 -opioid receptor activation. Although NO synthesis (by NOS 2) is mainly regulated at the transcriptional level by the induction of mRNA expression and there ussualy is a strong correlation between mRNA level and enzyme activity (Jacobs and Ignarro, 2001; Korhonen et al., 2001; Cho et al., 2003), there are numerous data in the literature which describe that NOS 2 activity does not follow the up-regulation of NOS 2 mRNA expression and vice versa (Bustamante et al. 1996; Lortie et al., 2000; Nakaya et al. 2005; Sakai et al., 2006). Therefore, changes in activity which do not correspond to changes in mRNA level could be due to post-translational regulation of NO synthesis such as protein stability, dimerization, phosphorylation, cofactor binding and availability of O2 and L-arginine as substrates (Pan et al., 1996; Salh et al., 1998; Aktan et al., 2004). A possible involvement of the -opioid receptor in endomorphin 1 mediated NO release was investigated using -opioid receptor selective antagonist -FNA. The greatest amount of NO release from endomorphin 1 treated J774 macrophages was partially blocked by 10-4 M -FNA (46% inhibition). Thus, the effect of endomorphin 1 on NO release was mostly -opioid receptor mediated. Also, using ICI 174,864 a -opioid receptor selective antagonist together with 10-6 M endomorphin 1 did not inhibit NO release, applying predominantly -opioid receptor involvement. 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(2003) Opioid receptors mediate a postsynaptic facilitation and a presynaptic inhibition at the afferent synapse of axolotl vestibular hair cells. Neuroscience 118:75-85. Vuji V, Stanojevi S, Dimitrijevi M (2004) Methionine-enkephalin stimulates hydrogen peroxide and nitric oxide production in rat peritoneal macrophages: Interaction of , , and  opioid receptors. Neuroimmunomodulation 11:392 403. Xin L, Blatteis CM (1992) Hypothalamic neuronal responses to interleukin-6 in tissue slices: Effects of indomethacin and naloxone. Brain Res Bull 29:27-35. Zadina JE, Hackler L, Ge L-J, Kastin AJ (1997) A potent and selective endogenous agonist for the -opiate receptor. Nature 386:499-502. Zhang L, Gu ZF, Pradhan T, Jensen RT, Maton PN (1992) Characterization of opioid receptors on smooth muscle cells from guinea pig stomach. Am J Physiol Gastrointest Liver Physiol 262: G461-G469. TABLES AND FIGURES  Fig. 1.  Fig. 2.  Fig. 3.  Fig. 4.  Fig. 5.  Fig. 6. FIGURE LEGENDS Fig. 1. Total nitrite concentration in cell culture supernatant accumulated during 48 h incubation of J774 macrophages with different concentrations of endomorphin 1 (EM 1) (A), endomorphin 2 (EM 2) (B) (10-6, 10-8 or 10-10 M), LPS (10 g/ml) or medium alone (control). NO production was measured using Griess reagent. Results are expressed as mean SEM of 3 experiments with individual cultures. Control values are 1.960.35 M for EM 1 and 2.630.53 M for EM 2 experiment. *p=0.0001 as compared with control. Fig. 2. Real time analysis of endomorphin 1 (EM 1) or endomorphin 2 (EM 2) (10-11, 10-10, 10-9, 10-8, 10-7 or 10-6 M) induced NO release by J774 (2x10-6cells/2 ml) macrophages. NO release was evaluated for 1400 s. Data are representative of each independent culture (n=3-4) which was subjected to all treatments examined. Fig. 3. NO release (nM) during J774 macrophage incubation with 10-6 M EM 1 or EM 2. After a baseline was established (10 min), cells were challenged with different drugs and the chart recordings were marked as zero. The NO nanomolar concentrations were calculated from picoamper current readings based on linear regression equation from standard curve using NaNO2 solution as standard (1.2041 nM/pA represents the sensitivity of nitric oxide meter sensor). Data are expressed as mean SEM from 3 to 5 experiments with individual cultures. *p=0.0001 and #p=0.001 as compared with control media treated cells. Inset, original recordings of the time course of NO release from J774 caused by 10-6 M EM 1. Fig. 4. Real time analysis of endomorphin 1 (EM 1, 10-6 M) induced NO release from J774 macrophages inhibited by a selective NOS 2 inhibitor 1400W in J774 cells. 1400W (10-4 M) was administered 30 min before EM 1. Data are expressed as mean SEM from 2 to 3 experiments using individual cultures. *p=0.001 compared to control media treated cells. Inset, original recordings of the time course of NO release. Fig. 5. Effect of LPS (10 g/ml) or 10-6 M EM 1 on NOS 2 mRNA expression in J774 macrophages. Cells were exposed to EM 1 or LPS (positive control) for 30 min (A) or 4 h (B). A basal condition of media treated cells served as control. Total RNA was extracted for RT-PCR amplifications of NOS 2 and -actin mRNA. Top; Electrophoresis of the RT-PCR products is shown via an image of the ethidium bromide stained agarose gel visualized under UV-transilluminator. The top image shows the representative results using NOS 2 specific primers, and the bottom image shows the results for -actin. Bottom; The mRNA bands on the gel photograph were quantified by densitometric analysis and expressed as NOS 2/-actin ratio. Results are representatives from 4 independent cultures for each group. *p=0.005 and #p=0.034 as compared with control. Fig. 6. Real time analysis of attenuated endomorphin 1 (EM 1, 10-6 M) induced NO release by a selective -opioid receptor antagonist -FNA in J774 macrophages. -FNA (10-4 M) was administered 30 min before EM 1. Data are expressed as mean SEM from 2 to 3 experiments. *p=0.001 compared with control media treated cells. Inset, original recordings of the time course of NO release. 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