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Immunoglobulin and Cytokine Production from Spleen Lymphocytes Is Modulated in C57BL/6J Mice by Dietary Cis-9, Trans-11 and Trans-10, Cis-12 Conjugated Linoleic Acid

Laboratory of Food Chemistry, Division of Applied Biological Chemistry, Department of Bioscience and Biotechnology, Faculty of Agriculture, Kyushu University, Higashi-ku, Fukuoka 812-8581, Japan and ^* Rinoru Oil Mills, Chuo-ku, Tokyo, 103-0027, Japan

¹To whom correspondence should be addressed. E-mail: masawo{at}agr.kyushu-u.ac.jp.

	ABSTRACT

TOP ABSTRACT MATERIALS AND METHODS RESULTS DISCUSSION LITERATURE CITED

 
We evaluated the effect of cis-9, trans-11 (9c,11t) and trans-10, cis-12 (10t,12c) conjugated linoleic acid (CLA) on the immune system in C57BL/6J mice. Mice were fed experimental diets containing 0% CLA (controls), 1% 9c,11t-CLA, 1% 10t,12c-CLA or a 1:1 mixture (0.5% + 0.5%) of these two CLA isomers for 3 wk. Relative spleen weights of all CLA fed mice were greater than the controls. Spleen lymphocytes isolated from the mice fed 10t,12c-CLA produced more immunoglobulin (Ig)A and IgM but not IgG when stimulated with concanavalin A (ConA) compared with controls. IgA production from unstimulated spleen lymphocytes was greater in the 10t, 12c-CLA group than in controls. Conversely, 9c,11t-CLA did not affect the production of any of the Ig subclasses. Lymphocytes isolated from 9c,11t-CLA fed mice produced more tumor necrosis factor- than the control group. The proportion of B cells in the spleen lymphocyte population was significantly lower in the 9c,11t-CLA group, and higher in the 10t,12c-CLA group than in the controls. Compared with the control group, the percentage of CD4⁺ T cells was lower in the 10t,12c-CLA group, and the percentage of CD8⁺ T cells was higher in the 9c,11t-CLA group. Furthermore, the percentage of CD8⁺ T cells was higher in the 1:1 mixture group than in controls. The CD4⁺/CD8⁺ ratio was lower in the 1:1 mixture group than in controls. These results suggest that 9c,11t and 10t,12c-CLA can stimulate different immunological effects and that the simultaneous intake of the two isomers can change the T cell population.

KEY WORDS: • conjugated linoleic acid • immunoglobulin • mice • cytokine

Conjugated linoleic acid (CLA)² is a generic term for the positional and structural isomers of octadecadienoic acid. CLA has been reported to exert various beneficial physiologic effects. In previous reports, the effects of CLA were often evaluated using a mixture of CLA isomers, which contained mainly the cis-9, trans-11 (9c,11t) and the trans-10, cis-12 (10t,12c)-CLA isomers. Recently, the role of each of these CLA isomers has been studied separately. In fact, some differences between the two isomers have been reported, especially the antiobesity effect (1 –4 ). We reported previously that dietary CLA enhanced immunoglobulin (Ig) production in rat spleen and mesenteric lymph node lymphocytes (5 ,6 ). Stimulation of Ig production by dietary CLA was detected at an extremely low dietary level (0.5 g/kg diet) (7 ). However, determining which of the CLA isomers stimulates Ig production and the mechanism of that stimulation remain to be more clearly elucidated. Thus, the objective of this study was to evaluate the individual and simultaneous effect of 9c,11t and 10t,12c-CLA on the production of Ig and cytokines and on the population of B and T cells in spleen lymphocytes from C57BL/6J mice.

	MATERIALS AND METHODS

TOP ABSTRACT MATERIALS AND METHODS RESULTS DISCUSSION LITERATURE CITED

 
Experimental animals and diet.

Semipurified 9c,11t and 10t,12c-CLA and safflower oil were purchased from Rinoru Oil Mills (Nagoya, Japan). Male 4-wk-old C57BL/6J mice (n = 40) (Japan CLEA, Tokyo, Japan) consumed a nonpurified commercial diet and water ad libitum for 2 wk after their arrival. After acclimation, the mice were divided into 4 groups of 10 mice each. They were kept in the Biotron Institute of Kyushu University with a 12-h light:dark cycle (light, 0800–2000 h) in an air conditioned room (20°C and 60% humidity under specific pathogen–free conditions). This experiment was carried out according to the guidelines for animal experiments at the Faculty of Agriculture and the Graduate Course, Kyushu University, and the Law (No. 105) and Notification (No. 6) of the Japanese Government. The experimental diets were manufactured according to the AIN-93G standard (8 ) and contained 0 g/kg CLA (control), 5 g/kg 9c,11t and 5 g/kg 10t,12c-CLA (1:1 mixture), 10 g/kg 9c,11t (9c,11t) or 10t,12c-CLA (10t,12c) (Table 1 ). For the basic dietary fat source, we used high linoleic acid safflower oil as in our previous report (7 ). The fatty acid composition of these diets is shown in Table 2 . At the end of the feeding period, mice were killed by drawing blood from the abdominal aorta under light anesthesia with diethylether. Immediately after excision, each tissue was weighed and the lymphocytes were isolated from the spleen.

Preparation of spleen lymphocytes was performed according to the method described previously (5 ) without removing adhesive cells such as macrophages and mononuclear cells. First, a cell suspension prepared from the spleen was rinsed with RPMI 1640 medium three times (Nissui, Tokyo, Japan). Then, 5 mL of the cell suspension was added to Lympholyte-mouse (Cedarlane, Hornby, Canada) to isolate the lymphocytes and the suspension was again washed three times with RPMI 1640 medium. The lymphocytes, 2.5 x 10⁹ cells/L, were cultured in RPMI 1640 medium containing 10% fetal bovine serum (Intergen, Purchase, NY) with or without 5 mg/L concanavalin A (ConA) (Sigma, St. Louis, MO) and incubated at 37°C for 24 h.

Measurement of immunoglobulin levels.

Measurement of Ig concentration in the cultured medium was performed by a sandwich ELISA. Rabbit anti-mouse IgA (Zymed, San Francisco, CA), goat anti-mouse IgG (H+L) (Zymed), rabbit anti-mouse IgM (µ-chain specific) (Zymed), and anti-mouse IgE from clone LO-ME-3 (heavy-chain specific) (Technopharm Biotechnology, Paris, France) were used to fix each Ig. These antibodies were diluted using 10% Block Ace (Dainihon Pharmaceutical, Osaka, Japan), added to a 96-well plate and incubated for 1 h at 37°C. Then, 300 µL of 10% (25% in IgE) Block Ace was added and kept at 4°C overnight; samples (50 µL) were added to each well for 1 h at 37°C. Each well was treated with a solution of either peroxidase (POD)-conjugated goat anti-mouse IgA (Zymed), POD-conjugated goat anti-mouse IgG (H+L) (Zymed), POD-conjugated rabbit anti-mouse IgM (Zymed), or POD-conjugated goat anti-mouse IgE [GAM/IgE (Fc) PO, Nordic Immunological Laboratory, Tilburg, Netherlands] to detect the respective Ig and incubated for 1 h at 37°C (20 min, 4°C for IgE). The plates were rinsed with PBS containing 0.5 g/L polyethylene sorbitan monolaurate (Nacalai Tesque, Kyoto, Japan) between each step. Then, a 10:9:1 mixture of 1.8 mmol/L H₂O₂ in 0.2 mol/L citrate buffer (pH 4.0), H₂O, and 11.7 mmol/L of 2,2'-azinobis (3-ethylbenzothiazoline sulfonic acid) was added. Finally, absorbance at 415 nm was measured after the addition of 160 mmol/L oxalic acid to stop the coloring reaction.

Measurement of cytokine levels.

We measured the level of interleukin (IL)-2, 4, 5, tumor necrosis factor (TNF)- and interferon (IFN)- in the supernatants of spleen lymphocytes cultured with ConA. IL-2, 4 and 5 were measured using commercial ELISA kits following the manufacturers’ experimental protocols (Mouse IL-2, 4 and 5 ELISA kit, BioSource International, Camarillo, CA). TNF- and IFN- levels were measured by sandwich ELISA as previously reported (9 ). Briefly, rabbit anti-mouse/rat IFN- (BioSource) and anti-mouse TNF- (Endogen, Woburn, MA) (500 times dilution) were used to fix IFN- and TNF- for 1 h at 37°C. Then, blocking was performed using 25% Block Ace at 37°C for 1 h. In the following step, 50 µL of appropriate cultured supernatant was added to each well and incubated for 2 h at 37°C; the plate was then treated with either a diluted solution of biotinylated anti-mouse IFN- (Genzyme, Cambridge, MA) (500 times dilution) or biotinylated anti-mouse/rat TNF- (Genzyme) (250 times dilution) for 1 h at 37°C. After that, streptavidin POD-conjugated (Zymed) diluted by 10% Block Ace was added to each well. Plate washing between each step and the coloring reaction were performed as in the Ig measurement protocol.

B and T cell population analysis.

The cell surface expression of CD45R (as a B cell marker) and CD4 and CD8 (as T cell subpopulation markers) was analyzed by flow cytometry. After the isolation of lymphocytes from the spleen, cells were washed with RPMI-1640 medium three times and treated with PBS containing 3% bovine serum albumin for 1 h at 37°C. Cells were divided into two groups (1.0 x 10⁶ cells each); one was exposed to rat phycoerythrin (PE)-conjugated monoclonal anti-mouse CD45R (clone RA3–6B2, Caltag Laboratories, Burlingame, CA) and the other was double stained with rat fluorescein isothiocyanate–conjugated monoclonal anti-mouse CD4 (clone CT-CD4, Caltag Laboratories) and rat PE-conjugated monoclonal anti-mouse CD8b (clone CT-CD8b, Caltag Laboratories). All of the antibody reactions were performed on ice for 1 h, and cells were washed three times with PBS after the antibody treatment. Samples were subjected to flow cytometry (FACS Calibur, Becton Dickinson, Sunnyvale, CA) and a total of 10⁴ cells were analyzed to determine the percentage of CD45R-, CD4- and CD8-positive lymphocytes.

Statistical Analysis.

At first, data were analyzed using one-way (Tables 3 , 5 and 6) or two-way (Table 4) ANOVA. The latter was used to identify differences due to diet or lymphocyte stimulation status. Fisher’s Protected Least Significant Difference test was used to determine which means differed (P < 0.05). All data are presented as means ± SEM.

View this table: [in this window] [in a new window]   TABLE 5 Tumor necrosis factor- production of spleen lymphocytes isolated from C57BL/6J mice fed 0 g/kg CLA, 5 g/kg 9c, 11t and 5 g/kg 10t, 12c-CLA, 10 g/kg 9c, 11t, or 10t, 12c-CLA diet for 3 wk1

	RESULTS

TOP ABSTRACT MATERIALS AND METHODS RESULTS DISCUSSION LITERATURE CITED

Food intake did not differ among the dietary groups (Table 3 ). Final body weight was higher in the 9c,11t group than in the 1:1 mixture or 10t, 12c groups, but it did not differ significantly from the controls. Relative liver weight was greater in the 1:1 mixture and 10t,12c groups than in the control and 9c,11t groups. Relative spleen weight was significantly higher in all CLA groups than in the control group. Perirenal and epididymal white adipose tissue weights were significantly lower in the 1:1 mixture and 10t,12c compared with the control and 9c,11t groups. Relative lung, heart and kidney weights did not differ among the dietary groups.

Immunoglobulin production from the spleen lymphocytes.

ConA did not affect IgA productivity in spleen lymphocytes (Table 4 ). IgA productivity of 10t,12c-fed mice was approximately twice that (P < 0.05) of the control group in the presence of ConA. IgA productivity also differed between the control and 10t,12c groups without ConA stimulation. IgA production from the lymphocytes in the 9c,11t group was lower than that of the 10t,12c group, irrespective of ConA stimulation. ConA stimulation significantly elevated IgG productivity in spleen lymphocytes. IgM production in spleen lymphocytes was significantly modulated by the type of dietary fat but not by ConA stimulation. IgM production from the lymphocytes in the 10t,12c group was significantly higher than that of the control group, irrespective of ConA stimulation. The 9c,11t and control groups did not differ significantly in IgM productivity. IgM productivity for the 1:1 mixture group was intermediate between the 9c,11t and 10t,12c groups. IgE production by lymphocytes was not affected by diet or ConA stimulation.

Cytokine productivity.

Cytokines were not detected in any of the cultured supernatants from lymphocytes that were not stimulated with ConA. No significant differences were found in IL-2, 4, 5 and IFN- production among any of the dietary groups (data not shown). TNF- production of spleen lymphocytes from mice in the 9c,11t group was significantly higher than that of the 10t,12c and control groups (Table 5 ).

B- and T-cell population.

The percentage of B cells in the 9c,11t group was significantly lower than in the control group, whereas the 10t,12c group was higher (Table 6 ). The value for the 1:1 mixture group was intermediate between the 9c,11t and 10t,12c groups. The percentage of CD4⁺ T cells was significantly lower in the 1:1 mixture and 10t,12c groups than in the control group. The percentage of CD8⁺ T cells was the highest for the 1:1 mixture group followed by the 9c,11t group. The CD4⁺/CD8⁺ ratios in the 9c,11t and 10t,12c groups were lower than in the control group, and the ratio in the 1:1 mixture group was lower than any of the other dietary groups. The CD4⁺/CD8⁺ ratios in the 9c,11t and 10t,12c groups were also significantly lower than the control group.

	DISCUSSION

TOP ABSTRACT MATERIALS AND METHODS RESULTS DISCUSSION LITERATURE CITED

 
The aim of this study was to evaluate the isomer-specific effect of CLA on the function of spleen lymphocytes in C57BL/6J mice. At present, 9c,11t and 10t,12c CLA are recognized for various beneficial physiologic functions, with each CLA isomer having both individual and synergistic roles in carrying out those functions such as a body fat–reducing effect or a growth-promoting effect. For example, 10t,12c-CLA definitely plays a leading part in reducing body fat (1 ,10 ), and we confirmed that this isomer does indeed have a specific body fat–reducing effect in mice (Table 3) . On the other hand, 9c,11t-CLA has been reported to promote mouse growth (10 ), which our present data also support because we found that the final body weight in the 9c,11t group was significantly higher than that in the control group (Table 3) . In addition, dietary CLA significantly increased the spleen weight compared with the control group; however, there was no significant difference among the CLA-fed mice. These data suggest that 9c,11t and 10t,12c-CLA have almost the same ability to increase the spleen mass and that no synergistic effect exists between these isomers. In previous reports, CLA feeding did not increase spleen weight in Sprague-Dawley rats (5 ,7 ). Thus, we speculate that species specificity exists between rats and mice spleens in terms of sensitivity to dietary CLA.

We reported previously that dietary CLA could enhance IgA, IgG and IgM production from the rat mesenteric lymph node and spleen lymphocytes (5 ,6 ). In rat spleen lymphocytes, only a diet containing 0.5 g/kg CLA dramatically promoted IgA, IgG and IgM production (7 ). However, as far as we know, no report has clearly shown which isomer(s) promote Ig production. In the present study, significant enhancement of IgA and IgM production was detected in the 10t,12c group but not in the 1:1 mixture and 9c,11t groups. This result indicates that 10t,12c-CLA plays a leading part in promoting Ig production. To examine the effect of CLA on the B cell ratio in spleen lymphocytes, we measured the amount of B lymphocyte–specific surface marker CD45R present to determine the positive cell population percentage. As a result, the B-cell percentage in the 10t,12c group was significantly higher than the control level. On the other hand, a decrease in the B-cell percentage was observed in the 9c,11t group, and the level in the 1:1 mixture group was intermediate to the 9c,11t and 10t,12c groups. Judging from these results, the elevation of the B-cell percentage in spleen lymphocytes by 10t,12c-CLA might be counteracted by 9c,11t-CLA. Taken together, it is likely that elevation of the B-cell ratio by 10t,12c-CLA contributes in part to the promotion of Ig production in spleen lymphocytes. However, we must consider that this augmentation of Ig production (x2) may be due in part to increased production per B cell because the increased number of CD45R⁺ cells was not doubled.

Significant enhancement of IgA and IgM production was also detected in the 10t,12c group with or without ConA, which is a T lymphocyte–specific mitogen (Table 4) . However, IgA and IgM production by spleen lymphocytes did not increase with ConA stimulation. We reported previously that dietary CLA enhanced Ig production in rat spleen and mesenteric lymph node lymphocytes with LPS stimulation which is a B lymphocyte–specific mitogen (6 ). In a previous report, dietary CLA stimulated IL-2 productivity in mouse spleen lymphocytes or splenocytes (11 ,12 ). In this study, neither dietary 9c,11t nor 10t,12c-CLA could inhibit IL-2 production from spleen lymphocytes. In addition, dietary 9c,11t and 10t,12c-CLA did not affect IL-4 and 5 production from spleen lymphocytes stimulated with ConA, and none of these cytokines were detected without ConA. These data indicate that the enhancement of Ig production in spleen lymphocytes by 10t,12c-CLA was not modulated by the stimulation of T lymphocytes during a 24-h incubation period.

Conversely, 9c,11t-CLA significantly stimulated TNF- production, and this result is consistent with our previous data (9 ). Turek et al. (13 ) reported that dietary CLA reduced TNF- and IL-6 production in rat macrophages, and CLA has been reported to suppress TNF- related cachexia (14 ). Conversely, recent reports showed that CLA did not affect TNF- production in splenocytes isolated from tumor-bearing rats stimulated with Escherechia coli endotoxin (15 ). Unfortunately, there is little information concerning isomer specificity in the regulation of TNF- production. Our present data indicate that 9c,11t and 10t,12c-CLA have quite different effects on the production of TNF- in spleen lymphocytes, but further studies are warranted to elucidate the target immune cells of each CLA isomer.

Most CD8⁺ T cells are major histocompatibility complex class I restricted killer T cells and exert cytotoxic activity when they are activated. CLA has been reported to elevate the CD8⁺ T cell population of porcine peripheral mononuclear cells (16 ,17 ). Conversely, CLA elevated CD4⁺ T cell population in mice and chicks (14 ,18 ), which resulted in an elevation of the CD4⁺/CD8⁺ ratio. In this report, only 10t,12c-CLA decreased the CD4⁺ T cells ratio and only the 9c,11t-CLA elevated the CD8⁺ T cells ratio. Interestingly, the CD4⁺/CD8⁺ ratio was the lowest in the 1:1 mixture group. These results suggest that both CLA isomers cooperatively modulated the T cell subpopulation and also acted individually.

In summary, 10t,12c-CLA increases IgA and IgM production and 9c,11t-CLA increases TNF- production. Moreover, these CLA isomers synergistically reduce the CD4⁺/CD8⁺ T cell population ratio.

	ACKNOWLEDGMENTS

 
We thank Perry Seto for proofreading the manuscript.

	FOOTNOTES

 
² Abbreviations used: ConA, concanavalin A; CLA, conjugated linoleic acid; Ig, immunoglobulin; IFN-, interferon-; IL, interleukin; LPS, lipopolysaccharide; PE, phycoerythrin; POD, peroxidase; TNF-, tumor necrosis factor-.

Manuscript received 24 September 2002. Initial review completed 9 October 2002. Revision accepted 25 November 2002.

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TOP ABSTRACT MATERIALS AND METHODS RESULTS DISCUSSION LITERATURE CITED

 

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