SP和CMP抗PCV2感染的作用和机理的实验研究
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摘要
茯苓多糖是从茯苓[poria cocos(Schw.) Wolf]中提取的,具有很强的抗肿瘤和免疫调节活性,能提高细胞介导的免疫活性的一种活性成分,但由于它的水溶性差,限制了其在临床上的应用。经过硫酸酯化修饰后得到的硫酸酯化茯苓多糖(Sulfation pachymaran, SP)和经羧甲基化得到的羧甲基茯苓多糖(Carboxymethytl pachymaram, CMP)易溶于水,具有抗肿瘤、抗病毒作用以及免疫调节的功能更加显著。而猪圆环病毒2型(porcine circovirus type2, PCV2)是断奶后仔猪多系统衰竭综合征(postweaning mutisystenic wasting syndrome, PMWS)的主要病原。该病在世界各地广泛流行,PMWS患猪淋巴组织中淋巴滤泡内B细胞依赖区域以及滤泡旁T细胞依赖区域的淋巴细胞减少,从而导致免疫缺陷,使宿主长期处于免疫抑制状态。本研究的目的旨在探讨sP和CMP对PcV2引起仔猪免疫细胞产生免疫抑制的调节作用以及对PCV2感染小鼠免疫功能的调节及其机理,为sP和cMP在猪免疫抑制性疾病综合防治中的应用提供理论依据。
1SP和CMP对PCV2感染仔猪脾细胞的增殖和分泌细胞因子的影响
筛选猪繁殖与呼吸综合征(Porcine reproductive and respiratory syndrome, PRRSV)/PCV2抗原和抗体均为阴性的断奶仔猪,无菌分离脾细胞进行体外培养。先加入1000TCID50PCV2病毒悬液孵育2h后,再加入终浓度分别为400μg/mL、200ng/mL、100μg/mL、50μg/mL、25μg/mL和12.5μg/mL的SP/CMP, MTT法测定猪脾T/B淋巴细胞体外增殖；ELISA方法检测脾细胞培养上清液中IL-1β、IL-2、IL-8、IL-10和TNF-α等细胞因子。结果显示,SP和CMP能协同ConA/LPS促进PCV2感染的猪脾T/B淋巴细胞体外增殖,提高PCV2感染的猪脾淋巴细胞的免疫应答能力。同时,sP和cMP能显著促进PcV2感染的猪脾细胞分泌IL-2,浓度为100μg/mL的CMP能下调PCV2感染仔猪脾细胞分泌IL-10。浓度为50～200μg/mL的SP能显著降低PCV2感染的猪脾细胞培养上清液中IL-1p水平,同时可刺激TNF-a的分泌。说明SP和CMP可以通过促进猪脾T/B淋巴细胞增殖活性以及调节细胞因子的分泌,增强细胞免疫应答,发挥免疫调节作用。
2SP和CMP对PCV2感染的猪肺泡巨噬细胞吞噬功能和活性的影响
筛选PRRSV/PCV2抗原和抗体均为阴性的断奶仔猪,通过支气管获取肺泡灌洗液并分离猪肺泡巨噬细胞进行体外培养。首先加入1000TCID50PCV2病毒悬液孵育2h,再加入终浓度分别为400μg/mL、200μg/mL、100μg/mL、50μg/mL、25μg/mL和12.5μg/mL的SP/CMP,分别测定体外培养的猪肺泡巨噬细胞吞噬功能和培养上清液NO含量以及IL-1β、IL-8、IL-10、TNF-a和MCP-1等细胞因子。结果显示,SP和CMP能提高PCV2感染的猪肺泡巨噬细胞吞噬功能。SP在第4h、8h、24h时能下调PCV2感染肺泡巨噬细胞引起NO的过度分泌,CMP在第4h和8h时能下调PCV2感染的猪肺泡巨噬细胞过度分泌的NO水平。同时,SP和CMP能有效地下调PCV2感染的肺泡巨噬细胞过度分泌IL-8和IL-10,并对IL-1p分泌发挥调节作用。结果表明了SP和CMP通过激活猪肺泡巨噬细胞吞噬功能以及调节NO和细胞因子的分泌,提高PCV2感染的猪肺泡巨噬细胞的免疫功能。
3SP和CMP对PCV2感染诱导猪脾细胞氧化应激的调节作用
为了探讨SP和CMP对PCV2感染的猪脾细胞氧化应激的调节作用,从PRRSV/PCV2抗原和抗体均为阴性的断奶仔猪中无菌分离猪脾细胞进行体外培养。首先在培养的脾细胞中加入1000TCID50PCV2病毒悬液,孵育感染2h后加入终浓度分别为200μg/mL、100μg/mL、50μg/mL.25μg/mL的SP/CMP,测定脾细胞内ROS水平以及细胞培养上清液超氧化物岐化酶(T-SOD)和总抗氧化能力(TAOC)。结果显示,SP和CMP能下调PCV2诱导免疫细胞产生过量的ROS；同时能提高感染PCV2的猪脾细胞T-SOD和TAOC活性。结果表明,SP和CMP能通过降低PCV2感染猪脾细胞内ROS,升高T-SOD和TAOC的活性来提高细胞的抗氧化应激能力,从而保护免疫细胞,发挥免疫调节作用。
4CMP对PCV2感染小鼠免疫功能的调节
为了研究CMP对PCV2感染小鼠的免疫调节作用,将体重为18-22g的100只昆明系小鼠随机分成5组(A-E组),A、B、C、D组在第1d、3d、5d、7d通过腹腔注射、滴鼻以及口服三种途径联合感染PCV2病毒悬液(1000TCID50),0.5mL/只,E组给予相同剂量生理盐水。在第8d,9d,10d,A、B、C组小鼠给予腹腔注射终浓度分别为200mg/kg.BW.100mg/kg.BW和50mg/kg.BW的CMP, D和E组给予相同剂量生理盐水。第11d剖杀各组小鼠,测定免疫器官指数、小鼠脾淋巴细胞的体外增殖活性、脾脏和胸腺组织T-SOD、TAOC和GSH含量；ELISA方法检测血清中IL-2、IL-10.IL-6和IFN-a含量；用荧光定量PCR方法检测脾脏和胸腺组织IL-2、IL-10mRNA表达量。结果显示,CMP能显著升高PCV2感染小鼠的脾脏和胸腺指数以及促进T或B淋巴细胞体外增殖,提高脾脏和胸腺组织的TSOD、T-AOC和GSH含量；上调PCV2感染小鼠血清IL-2、IL-6和IFN-a含量以及下调血清IL-10水平,并提高PCV2感染小鼠脾脏和胸腺组织IL-2mRNA表达水平。结果表明,CMP能通过提高PCV2感染小鼠免疫器官的抗氧化能力和免疫功能来调节PCV2引起的免疫抑制作用。
Pachymaran is extracted from Poria cocos (Schw.) Wolf and has strong antitumor and immunomodulatory activities and cell-mediated immune activities.Pachymaran is not easily applied in the clinic because of its low water-solubility. Sulfation pachymaran (SP) is obtained from artificially sulfated modification and Carboxymethylpachymaran (CMP) is obtained from artificially carboxymethylation.The two polysaccharides are water-soluble and exhibit good immune regulation and antitumor activities. Porcine circovirus type2(PCV2) is the primary causative agent of postweaning multisystemic wasting syndrome (PMWS), a worldwide disease that debilitates nursery and fattening pigs with systemic lymphadenopathy and interstitial pneumonia.Microscopic lesions in the lymphoid tissues of pigs severely affected with PMWS exhibits lymphocyte depletion with loss of lymphoid follicles (mainly in the B cell-dependent areas) and reduced cellularity in the parafollicular T cell-dependent parts.So swine infected with PCV2always exhibit immunosuppression in a long-term.The objective of this study is to investigate the function of SP and CMP on porcine immunocytes inoculated with PCV2in vitro and immunological regulation of CMP in mice after infected with PCV2in vivo.
1Effects of SP or CMP on proliferation of splenic lymphocytes and cytokine after inoculated with PCV2
In order to investigate the immunomodulatory mechanism of SP or CMP on the porcine splenic lymphocytes after the cells were inoculated with PCV2.The splenic lymphocytes from pigs which were negative for both antigens and antibodies of PRRSV or PCV2were treated with SP or CMP after inoculation with PCV2for2h.Lymphocyte proliferation was determined using the MTT method. The levels of IL-1β, IL-2, IL-8, IL-10and TNF-a were detected by ELISA methods.The results showed that the two polysaccharides were able to promote the proliferation activities of splenic lymphocytes with or without inoculation of PCV2.The SP at concentrations from50μg/mL to400μg/mL or CMP at concentrations from200μg/mL to400μg/mL was able to stimulate proliferation of splenic B lymphocytes in vitro.The level of IL-1β in splenic lymphocytes inoculated with PCV2decreased after splenic lymphocytes were treated with SP at different concentrations from50μg/mL to200μg/mL.The level of IL-2in splenic lymphocytes inoculated with PCV2increased after splenic lymphocytes were treated with SP or CMP.The level of IL-10in splenic lymphocytes inoculated with PCV2decreased when splenic lymphocytes were treated CMP at with100μg/mL,while the level of TNF-α increased.These results showed that SP and CMP were able to stimulate proliferation of swine splenic T and B lymphocytes with or without inoculation of PCV2in vitro.
2Effects of SP or CMP on activities and phagocytosis immune function of porcine alveolar macrophage after inoculated with PCV2
The effects of SP or CMP on immunological function of swine alveolar macrophage inoculated with porcine circovirus type2(PCV2) were observed.The swine alveolar macrophages were separeated from pigs which were negative for both antigens and antibodies of PRRSV or PCV2and treated with SP or CMP after inoculation of PCV2.The phagocytosis of swine alveolar macrophage was observed by neutral red method,the nitric oxide (NO) level was determined by Griess method and the cytokine levels of IL-1β、IL-8、 TNF-a and MCP-1were measured by ELISA method.The results showed that SP and CMP could stimulate proliferation of swine alveolar macrophages.The content of NO decreased after swine alveolar macrophages were treated with PCV2and SP for4h,8h and24h, while the levels of NO decreased after swine alveolar macrophages were treated with PCV2and SP for4h and8h.The levels of TNF-α increased while the IL-8decreased after swine alveolar macrophages treated with PCV2and SP or CMP.These results suggests that SP and CMP might play an important role in regulating immunological functions by improving phagocytotic activity and promoting secretion of cytokines of swine alveolar macrophages.
3Effect of SP or CMP on oxidative stress in the swine splenic lymphocytes after inocubated with PCV2
The effect of SP or CMP on oxidative stress in the swine splenic lymphocytes inocubated withPCV2were observed.The splenic lymphocytes from pigs which were negative for both antigens and antibodies of PRRSV or PCV2were treated with SP or CMP at different concentrations from25μg/mL to200μg/mL post-inoculation of PCV2. The reactive oxygen species (ROS) level,total superoxidase dismutase (T-SOD) activity and total antioxidant capacity (TAOC) were measured. The results showed that SP and CMP were able to decrease the level of ROS in the swine splenic lymphocytes after inoculated with PCV2,and also could increased the activities of T-SOD and TAOC.Thesr results showed that SP and CMP could enhance the anitoxidantive activities of lymphocytes so as to protect the lymphocyte infection.
4Effect of CMP on immune function of mice after infected with PCV2
To investigate the effect of CMP on the immunological function of mice infected with PCV2,100of Kunming inbred mice were randomly divided into five groups (20mice each).Four groups were inoculated (intranasal and intraperitoneal injection at a dose of0.2mL) four times with PCV2(once daily on Dayl, Day3, Day5and Day7at1000TCID50of PCV2) and labeled the mPCV2group.The mice in the another group were sham-inoculated with a sterile cell culture medium using the same experimental conditions and functioned as the normal control group.Three groups of mPCV2mice were then given intraperitoneal injections of50,100,or200mg/kg body weight CMP once daily on Day8,Day9and Day10post-inoculation (dpi).At the same time,another group of mPCV2mice and the normal control group received the same volume of distilled water under the same conditions.On Day11dpi,the spleen and thymus indices were calculated.Lymphocyte proliferation in vitro was determined using the MTT method.Determination of cytokine levels in murine serum was carried out by using ELISA method.The activities of T-SOD and TAOC and GSH in the spleen and thymus were determined.The results showed that CMP administration significantly improved the spleen or thymus index,promoted the proliferation activities of T or B lymphocytes,and increased the production of glutathione, the superoxidase dismutase capacity and the total antioxidant capacity in the spleen or thymus of PCV2-infected mice.The administration of different CMP doses to PVC2-inoculated mice resulted in the upregulation in the levels of IL-2,IL-6and IFN-a,otherwise the downregulation of IL-10levels in the serum.These findings suggest that CMP has potential application in regulating immune functions to overcome the immunosuppresion caused by PCV2infection in mice.The findings may also prove useful in designing effective therapies against PCV2infection.
1SP和CMP对PCV2感染仔猪脾细胞的增殖和分泌细胞因子的影响
筛选猪繁殖与呼吸综合征(Porcine reproductive and respiratory syndrome, PRRSV)/PCV2抗原和抗体均为阴性的断奶仔猪,无菌分离脾细胞进行体外培养。先加入1000TCID50PCV2病毒悬液孵育2h后,再加入终浓度分别为400μg/mL、200ng/mL、100μg/mL、50μg/mL、25μg/mL和12.5μg/mL的SP/CMP, MTT法测定猪脾T/B淋巴细胞体外增殖；ELISA方法检测脾细胞培养上清液中IL-1β、IL-2、IL-8、IL-10和TNF-α等细胞因子。结果显示,SP和CMP能协同ConA/LPS促进PCV2感染的猪脾T/B淋巴细胞体外增殖,提高PCV2感染的猪脾淋巴细胞的免疫应答能力。同时,sP和cMP能显著促进PcV2感染的猪脾细胞分泌IL-2,浓度为100μg/mL的CMP能下调PCV2感染仔猪脾细胞分泌IL-10。浓度为50～200μg/mL的SP能显著降低PCV2感染的猪脾细胞培养上清液中IL-1p水平,同时可刺激TNF-a的分泌。说明SP和CMP可以通过促进猪脾T/B淋巴细胞增殖活性以及调节细胞因子的分泌,增强细胞免疫应答,发挥免疫调节作用。
2SP和CMP对PCV2感染的猪肺泡巨噬细胞吞噬功能和活性的影响
筛选PRRSV/PCV2抗原和抗体均为阴性的断奶仔猪,通过支气管获取肺泡灌洗液并分离猪肺泡巨噬细胞进行体外培养。首先加入1000TCID50PCV2病毒悬液孵育2h,再加入终浓度分别为400μg/mL、200μg/mL、100μg/mL、50μg/mL、25μg/mL和12.5μg/mL的SP/CMP,分别测定体外培养的猪肺泡巨噬细胞吞噬功能和培养上清液NO含量以及IL-1β、IL-8、IL-10、TNF-a和MCP-1等细胞因子。结果显示,SP和CMP能提高PCV2感染的猪肺泡巨噬细胞吞噬功能。SP在第4h、8h、24h时能下调PCV2感染肺泡巨噬细胞引起NO的过度分泌,CMP在第4h和8h时能下调PCV2感染的猪肺泡巨噬细胞过度分泌的NO水平。同时,SP和CMP能有效地下调PCV2感染的肺泡巨噬细胞过度分泌IL-8和IL-10,并对IL-1p分泌发挥调节作用。结果表明了SP和CMP通过激活猪肺泡巨噬细胞吞噬功能以及调节NO和细胞因子的分泌,提高PCV2感染的猪肺泡巨噬细胞的免疫功能。
3SP和CMP对PCV2感染诱导猪脾细胞氧化应激的调节作用
为了探讨SP和CMP对PCV2感染的猪脾细胞氧化应激的调节作用,从PRRSV/PCV2抗原和抗体均为阴性的断奶仔猪中无菌分离猪脾细胞进行体外培养。首先在培养的脾细胞中加入1000TCID50PCV2病毒悬液,孵育感染2h后加入终浓度分别为200μg/mL、100μg/mL、50μg/mL.25μg/mL的SP/CMP,测定脾细胞内ROS水平以及细胞培养上清液超氧化物岐化酶(T-SOD)和总抗氧化能力(TAOC)。结果显示,SP和CMP能下调PCV2诱导免疫细胞产生过量的ROS；同时能提高感染PCV2的猪脾细胞T-SOD和TAOC活性。结果表明,SP和CMP能通过降低PCV2感染猪脾细胞内ROS,升高T-SOD和TAOC的活性来提高细胞的抗氧化应激能力,从而保护免疫细胞,发挥免疫调节作用。
4CMP对PCV2感染小鼠免疫功能的调节
为了研究CMP对PCV2感染小鼠的免疫调节作用,将体重为18-22g的100只昆明系小鼠随机分成5组(A-E组),A、B、C、D组在第1d、3d、5d、7d通过腹腔注射、滴鼻以及口服三种途径联合感染PCV2病毒悬液(1000TCID50),0.5mL/只,E组给予相同剂量生理盐水。在第8d,9d,10d,A、B、C组小鼠给予腹腔注射终浓度分别为200mg/kg.BW.100mg/kg.BW和50mg/kg.BW的CMP, D和E组给予相同剂量生理盐水。第11d剖杀各组小鼠,测定免疫器官指数、小鼠脾淋巴细胞的体外增殖活性、脾脏和胸腺组织T-SOD、TAOC和GSH含量；ELISA方法检测血清中IL-2、IL-10.IL-6和IFN-a含量；用荧光定量PCR方法检测脾脏和胸腺组织IL-2、IL-10mRNA表达量。结果显示,CMP能显著升高PCV2感染小鼠的脾脏和胸腺指数以及促进T或B淋巴细胞体外增殖,提高脾脏和胸腺组织的TSOD、T-AOC和GSH含量；上调PCV2感染小鼠血清IL-2、IL-6和IFN-a含量以及下调血清IL-10水平,并提高PCV2感染小鼠脾脏和胸腺组织IL-2mRNA表达水平。结果表明,CMP能通过提高PCV2感染小鼠免疫器官的抗氧化能力和免疫功能来调节PCV2引起的免疫抑制作用。
Pachymaran is extracted from Poria cocos (Schw.) Wolf and has strong antitumor and immunomodulatory activities and cell-mediated immune activities.Pachymaran is not easily applied in the clinic because of its low water-solubility. Sulfation pachymaran (SP) is obtained from artificially sulfated modification and Carboxymethylpachymaran (CMP) is obtained from artificially carboxymethylation.The two polysaccharides are water-soluble and exhibit good immune regulation and antitumor activities. Porcine circovirus type2(PCV2) is the primary causative agent of postweaning multisystemic wasting syndrome (PMWS), a worldwide disease that debilitates nursery and fattening pigs with systemic lymphadenopathy and interstitial pneumonia.Microscopic lesions in the lymphoid tissues of pigs severely affected with PMWS exhibits lymphocyte depletion with loss of lymphoid follicles (mainly in the B cell-dependent areas) and reduced cellularity in the parafollicular T cell-dependent parts.So swine infected with PCV2always exhibit immunosuppression in a long-term.The objective of this study is to investigate the function of SP and CMP on porcine immunocytes inoculated with PCV2in vitro and immunological regulation of CMP in mice after infected with PCV2in vivo.
1Effects of SP or CMP on proliferation of splenic lymphocytes and cytokine after inoculated with PCV2
In order to investigate the immunomodulatory mechanism of SP or CMP on the porcine splenic lymphocytes after the cells were inoculated with PCV2.The splenic lymphocytes from pigs which were negative for both antigens and antibodies of PRRSV or PCV2were treated with SP or CMP after inoculation with PCV2for2h.Lymphocyte proliferation was determined using the MTT method. The levels of IL-1β, IL-2, IL-8, IL-10and TNF-a were detected by ELISA methods.The results showed that the two polysaccharides were able to promote the proliferation activities of splenic lymphocytes with or without inoculation of PCV2.The SP at concentrations from50μg/mL to400μg/mL or CMP at concentrations from200μg/mL to400μg/mL was able to stimulate proliferation of splenic B lymphocytes in vitro.The level of IL-1β in splenic lymphocytes inoculated with PCV2decreased after splenic lymphocytes were treated with SP at different concentrations from50μg/mL to200μg/mL.The level of IL-2in splenic lymphocytes inoculated with PCV2increased after splenic lymphocytes were treated with SP or CMP.The level of IL-10in splenic lymphocytes inoculated with PCV2decreased when splenic lymphocytes were treated CMP at with100μg/mL,while the level of TNF-α increased.These results showed that SP and CMP were able to stimulate proliferation of swine splenic T and B lymphocytes with or without inoculation of PCV2in vitro.
2Effects of SP or CMP on activities and phagocytosis immune function of porcine alveolar macrophage after inoculated with PCV2
The effects of SP or CMP on immunological function of swine alveolar macrophage inoculated with porcine circovirus type2(PCV2) were observed.The swine alveolar macrophages were separeated from pigs which were negative for both antigens and antibodies of PRRSV or PCV2and treated with SP or CMP after inoculation of PCV2.The phagocytosis of swine alveolar macrophage was observed by neutral red method,the nitric oxide (NO) level was determined by Griess method and the cytokine levels of IL-1β、IL-8、 TNF-a and MCP-1were measured by ELISA method.The results showed that SP and CMP could stimulate proliferation of swine alveolar macrophages.The content of NO decreased after swine alveolar macrophages were treated with PCV2and SP for4h,8h and24h, while the levels of NO decreased after swine alveolar macrophages were treated with PCV2and SP for4h and8h.The levels of TNF-α increased while the IL-8decreased after swine alveolar macrophages treated with PCV2and SP or CMP.These results suggests that SP and CMP might play an important role in regulating immunological functions by improving phagocytotic activity and promoting secretion of cytokines of swine alveolar macrophages.
3Effect of SP or CMP on oxidative stress in the swine splenic lymphocytes after inocubated with PCV2
The effect of SP or CMP on oxidative stress in the swine splenic lymphocytes inocubated withPCV2were observed.The splenic lymphocytes from pigs which were negative for both antigens and antibodies of PRRSV or PCV2were treated with SP or CMP at different concentrations from25μg/mL to200μg/mL post-inoculation of PCV2. The reactive oxygen species (ROS) level,total superoxidase dismutase (T-SOD) activity and total antioxidant capacity (TAOC) were measured. The results showed that SP and CMP were able to decrease the level of ROS in the swine splenic lymphocytes after inoculated with PCV2,and also could increased the activities of T-SOD and TAOC.Thesr results showed that SP and CMP could enhance the anitoxidantive activities of lymphocytes so as to protect the lymphocyte infection.
4Effect of CMP on immune function of mice after infected with PCV2
To investigate the effect of CMP on the immunological function of mice infected with PCV2,100of Kunming inbred mice were randomly divided into five groups (20mice each).Four groups were inoculated (intranasal and intraperitoneal injection at a dose of0.2mL) four times with PCV2(once daily on Dayl, Day3, Day5and Day7at1000TCID50of PCV2) and labeled the mPCV2group.The mice in the another group were sham-inoculated with a sterile cell culture medium using the same experimental conditions and functioned as the normal control group.Three groups of mPCV2mice were then given intraperitoneal injections of50,100,or200mg/kg body weight CMP once daily on Day8,Day9and Day10post-inoculation (dpi).At the same time,another group of mPCV2mice and the normal control group received the same volume of distilled water under the same conditions.On Day11dpi,the spleen and thymus indices were calculated.Lymphocyte proliferation in vitro was determined using the MTT method.Determination of cytokine levels in murine serum was carried out by using ELISA method.The activities of T-SOD and TAOC and GSH in the spleen and thymus were determined.The results showed that CMP administration significantly improved the spleen or thymus index,promoted the proliferation activities of T or B lymphocytes,and increased the production of glutathione, the superoxidase dismutase capacity and the total antioxidant capacity in the spleen or thymus of PCV2-infected mice.The administration of different CMP doses to PVC2-inoculated mice resulted in the upregulation in the levels of IL-2,IL-6and IFN-a,otherwise the downregulation of IL-10levels in the serum.These findings suggest that CMP has potential application in regulating immune functions to overcome the immunosuppresion caused by PCV2infection in mice.The findings may also prove useful in designing effective therapies against PCV2infection.
引文
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