摘要
车前子定义为车前（Plantago asiatica L.，又称大粒车前）或平车前（Plantagodepressa Willd.）的干燥成熟种子，是传统中医用药之一。本文以江西吉安产大粒车前子为研究对象，研究车前子多糖体内外消化过程和酵解方式，及其对肠道功能的提升作用。首先通过体外研究，建立人体胃肠道消化酵解模拟系统，研究大粒车前子多糖在口腔及胃肠道中的消化过程和在人体粪便菌群中的酵解方式；模拟和探讨大粒车前子多糖肠道功能。再利用体内实验，研究车前子多糖在小鼠体内的酵解方式及其对结肠功能指标的影响，以及物理加工方式对车前子多糖肠道生理活性的影响；与此同时分析车前子多糖对小鼠体内相关肠道生理指标及菌群的影响，最后进一步通过结肠炎小鼠模型探讨车前子多糖对模型小鼠肠道功能的作用。主要研究结论如下：
1.通过体外模拟口腔唾液，胃部和小肠消化实验研究车前子多糖的消化过程，结果发现唾液淀粉酶对车前子多糖相对分子质量没有影响，而在胃肠道消化过程中多糖受到影响。在模拟的胃、小肠消化体系中，多糖的相对分子质量发生改变，从(1,903.1±93.0) x103降低为(4.7±0.2) x103，还原糖含量从0.157±0.009mM增加为0.622±0.026mM，这表明车前子多糖相对分子质量的降低是由于糖苷键的断裂。同时，在整个模拟的消化过程中，没有检测到游离单糖，表明车前子多糖在模拟胃肠消化过程中没有产生游离单糖。这些结果能为车前子多糖的体外消化提供一些信息，并能其他多糖的消化提供参考。
2.探讨了大粒车前子多糖体外酵解及其碳水化合物对酵解发挥的作用。车前子多糖在体外由人体粪便培养物进行酵解24h。在酵解过程中，粪便培养物的pH由6.1降为5.1，总短链脂肪酸（SCFA）、乙酸、丙酸和正丁酸的含量均显著增加。车前子多糖主要由木糖，阿拉伯糖及半乳糖醛酸组成，因此在酵解过程中，木聚糖酶、阿拉伯呋喃糖酶、木糖苷酶和葡萄糖醛酸酶的活性也都增加。酵解24h后，多糖中47.2±1.6%总碳水化合物被消耗，这其中包括阿拉伯糖（消耗42.9±1.5%）、木糖（消耗53.2±1.6%）和葡萄糖醛酸（消耗76.4±1.2%）。同时，探讨了多糖中碳水化合物的消耗与SCFA的产量间的关系。研究发现，乙酸和正丁酸的增加主要是由于多糖中葡萄糖醛酸和木糖的发酵，而丙酸的增加则主要是由于阿拉伯糖和木糖的酵解。这些结果提示车前子多糖对大肠健康的有益作用，并且其碳水化合物的组成影响其SCFA的产量。
3.体外模拟研究车前子多糖对肠道功能的作用。结果表明，车前子多糖能显著延缓葡萄糖的扩散并对α-淀粉酶的活性有显著抑制作用，这表明车前子多糖的摄入具有潜在降低餐后葡萄糖浓度的能力。车前子多糖能抑制胰脂肪酶的活性，这能有助于降低血清中脂质的水平。车前子多糖对降低了胃蛋白酶的活性，这在一定程度上影响蛋白质的消化率。研究同时发现车前子多糖能结合并绑定胆汁酸，同时能降低可吸收的胆固醇量。这些结果表明车前子多糖对肠道功能有益。
4.体内研究车前子多糖对小鼠结肠的影响。小鼠经口灌胃服用车前子多糖30天（剂量：0.4g/kg体重）后，研究发现车前子多糖处理组小鼠结肠内容物的总SCFA，乙酸，丙酸和正丁酸的浓度均显著高于空白组（p <0.05）。同时，在整个实验周期内多糖处理组小鼠结肠粪便含水量也显著高于空白组（p <0.05），增加结肠粪便的持水力。多糖处理组小鼠的结肠pH在灌胃周期内从7.5±0.1降低为7.2±0.1，而空白组小鼠结肠pH保持7.5±0.1不变。这些结果表明摄入车前子多糖对结肠的健康产生积极影响。
5.体内研究车前子多糖对小鼠营养物质代谢和结肠菌群的影响。小鼠经口灌胃车前子多糖30天（剂量：0.4g/kg体重），研究发现车前子多糖的摄入能降低脂肪的表观吸收率。多糖处理组小鼠的血清总甘油三酯，总胆固醇和动脉硬化指数，以及肝脏胆固醇和总脂水平，均比空白组小鼠相应指标显著性降低（p <0.05）。同时，车前子多糖的摄入能显著缩短食物在胃肠道的通过时间（p<0.05），而这能在一定程度上影响车前子多糖对脂类代谢的作用。此外，多糖处理组的小鼠结肠菌群的多样性高于空白组。小鼠结肠中的Bacteroides sp.，Eubacterium sp.，产丁酸菌Butyrivibrio sp.，益生菌Bifidobacterium bifidum，Lactobacillus fermentum和Lactobacillus reuteri都由于多糖的摄入而增加。这些结果表明车前子多糖对脂类的代谢及结肠菌群有积极的作用。
6.围绕物理处理方式（微波处理及高压均质处理）对车前子多糖肠道功能的作用展开研究。微波处理后，多糖的表观粘度，平均相对分子质量和粒径随着微波的处理而降低。同时，多糖中还原糖含量随着分子量的降低而增加，这表明微波处理过程中多糖的降解伴随有糖苷键的断裂。扫描电镜（ESEM）显示多糖的表观形貌从大薄片状变为小的碎片。FT-IR光谱显示微波处理并没有改变车前子多糖的主要官能团。然而，微波处理多糖后，多糖在体外酵解中的SCFA产量和微生物胞外酶（木聚糖酶，阿拉伯呋喃糖酶，木糖苷酶和葡萄糖醛酸酶）活性显著提高。同时研究发现高压均质能使得多糖粒度的下降，并使得多糖的表观形貌由大的片状结构转变为小的多孔片。FT-IR光谱显示高压均质并没有改变车前子多糖的主要结构特征。然而，高压均质处理多糖后，多糖在小鼠盲肠和结肠中总SCFA，丙酸和正丁酸的产量显著提高。这些结果表明，微波处理及高压均质能成为增加多糖附加价值的方式。
7.研究并探讨摄入车前子多糖对患结肠炎小鼠症状的减缓作用。雌性BALB/c小鼠经饮用水服用葡聚糖硫酸钠（DSS）致结肠炎。随着DSS的摄入或在DSS摄入后，给予小鼠经口每天灌胃不同剂量的车前子多糖（0.2或0.4g/kg小鼠体重）。研究发现车前子多糖能减缓DSS引起的结肠炎症状（体重下降，粪便便血和腹泻）。多糖的摄入能抑制结肠的变短，并增加结肠组织中超氧化物歧化酶的酶活，降低髓过氧化物酶和内皮型一氧化氮合酶的酶活，增加宏观症状状态分数。同时，小鼠因患结肠炎而造成的结肠组织损害，胸腺指数下降，和C-反应蛋白水平上升均由于多糖的摄入而减缓。DSS诱导结肠炎致使的结肠组织中TNF-α，IL-1β，IL-6，MIP-2和LTB4水平的增加也都由于多糖的服用而减轻。同时，所有剂量的多糖能使得患结肠炎小鼠粪便短链脂肪酸下降减缓。荧光定量PCR（QPCR）结果表明，车前子多糖的摄入能减少患结肠炎小鼠结肠菌群中Bifidobacterium，Lactobacilli，Bacteroidaceae和Enterobacteriaceae量的降低。结果表明车前子多糖的摄入具有减缓和治疗结肠炎症状的作用。
8.研究并探讨车前子多糖的提前摄入对患结肠炎小鼠症状的预防作用。雌性BALB/c小鼠经饮用水服用DSS致结肠炎，并在DSS服用前，小鼠经口灌胃车前子多糖（0.2或0.4g/kg小鼠体重）一周。研究发现车前子多糖的提前摄入能减缓DSS引起的结肠炎症状（包括体重下降，粪便便血和腹泻）。多糖的提前摄入能抑制结肠的变短，降低髓过氧化物酶酶活，增加宏观症状状态分数。同时，小鼠因患结肠炎而造成的结肠组织的组织损害，胸腺指数下降，结肠指数上升和C-反应蛋白水平上升均由于多糖的提前摄入而减缓。DSS诱导结肠炎致使的结肠组织促炎症细胞因子TNF-α，IL-1β，IL-6，IL-12水平和巨噬细胞炎症蛋白MIP-2水平的增加，以及抗炎症细胞因子IL-4的下降，也都由于多糖的提前服用而减轻。同时，多糖的提前摄入能使得患结肠炎小鼠粪便短链脂肪酸的下降减缓。QPCR结果表明，车前子多糖的提前摄入能减少患结肠炎小鼠结肠菌群中Bifidobacterium，Lactobacilli，Bacteroidaceae和Enterobacteriaceae量的降低，并能减缓Clostridium量的增加。结果表明车前子多糖的提前摄入具有潜在预防和减缓结肠炎症状的作用。
Plantago, including Plantago asiatica L. and Plantago depressa Willd., waswidely used in traditional Chinese medicine. The seeds of Plantago asiatica L. fromJi’an were subjected for study in this research. Digestion, fermentation and intestinalfunctions of polysaccharide from seeds of P. asiatica L. were investigated in vitroand in vivo. The saliva, gastric and intestinal digestion, and fermentation of thepolysaccharide from P. asiatica L. were simulated and analyzed. Effects of thepolysaccharide on intestinal function were evaluated. Effects of physical treatmentson physiological activities of the polysaccharide in simulated gastrointestinalenvironment were also studied. In addition, mice were given oral administration ofpolysaccharide from P. asiatica L. by gavage to investigate the effects of thepolysaccharide on mouse colon, nutrient metabolism and colon microbiota. Effectsof physical treatments on mouse physiological activities of the polysaccharide werealso studied. Researches were also focus on whether oral administration andpre-administration of the polysaccharide could acute or prevent development andcolon microbiota changes in mice with dextran sodium sulfate (DSS)-inducedexperimental colitis. Main conclusions are summarized as follows:
1. The saliva, gastric and intestinal digestion of polysaccharide from P.asiatica L. seeds was investigated in vitro. It was found that salivary amylase had noeffect on the polysaccharide; however, the polysaccharide was influenced in latergastrointestinal digestion. A steady decrease in molecular weight (Mw) of thepolysaccharide from (1903.1±93.0) x103to (4.7±0.2) x103was observed asdigestion time increased. Meanwhile, the reducing ends were increased from0.157±0.009to0.622±0.026mM, indicating the decrease of Mw may due to thebreakdown of glycosidic bonds. In addition, there was no monosaccharide releasedthroughout the whole digestion period, suggesting that the gastrointestinal digestiondid not result in a production of free monosaccharide. These results may providesome information on the digestion of polysaccharide from P. asiatica L. in vitro, andmay contribute to the methods of studying the digestion of other carbohydrates.
2. In vitro fermentation of the polysaccharide from seeds of P. asiatica L. andthe contribution of its carbohydrates to the fermentation were investigated in thisstudy. The polysaccharide was characterized by high contents of xylose, arabinoseand glucuronic acid, and it was subjected to human fecal cultures to be fermented invitro for24h. During fermentation, pH in fecal cultures decreased from6.1to5.1and the levels of total short-chain fatty acid (SCFA), acetic, propionic and n-butyricacids all significantly increased. Xylanase, arabinofuranosidase, xylosidase andglucuronidase activities were also improved. After24h incubation,47.2±1.6%oftotal carbohydrate in polysaccharide, including42.9±1.5%of arabinose,53.2±1.6%of xylose and76.4±1.2%of glucuronic acid, were consumed. In addition,relationship between carbohydrate consumption of the polysaccharide and SCFAproduction was also evaluated. It was found that the increase of acetic and n-butyricacid productions mainly resulted from the fermentation of glucuronic acid andxylose in polysaccharide, while the increase of propionic acid production wasprimarily due to the fermentation of arabinose and xylose. These results showed thatthe polysaccharide was physiologically active for human large bowel, and itscarbohydrate composition determined its SCFA production.
3. Effects of polysaccharide from the seeds of P. asiatica L. on intestinalfunction were investigated in vitro. Results showed that the polysaccharide hadnotable influence on slowing down glucose diffusion and inhibiting α-amylaseactivity. These might help prolong blood glucose response and hence control thepostprandial glucose concentration. The polysaccharide could also decreasepancreatic lipase and protease activities, which may help lower the levels of serumlipids and modify protein digestibility. In addition, the polysaccharide was able tobind bile acids and may reduce cholesterol level. These results suggested that thepolysaccharide may have potential benefits for human intestinal function and mightbe used as a potential ingredient in functional food applications.
4. Mice were given30days oral administration of polysaccharide from P.asiatica L. at the dose of0.4g/kg body weight by gavage to investigate the effects ofthe polysaccharide on mouse colon. Results showed that the concentrations of totalSCFA, acetic, propionic, and n-butyric acids in mouse colonic content of polysaccharide treated group were all significantly higher than that of control group(water)(p <0.05). In addition, moisture of mouse colonic content of polysaccharidetreated group was also notably higher than that of the control group (p <0.05)indicating the intake of polysaccharide from P. asiatica L. resulted in a strongerwater-holding capacity for colonic content throughout the experimental period.Furthermore, a decreased pH (from7.5±0.1to7.2±0.1) was observed in mousecolon of the polysaccharide treated group compared with the control group (pH from7.5±0.1to7.5±0.1). These results suggested that the intake of the polysaccharidefrom P. asiatica L. might be beneficial for the colon health.
5. Polysaccharide from the seeds of P. asiatica L. was given via oraladministration to mice (0.4g/kg body weight,30days) to observe its effects onmouse nutrient metabolism and colon microbiota. It was found the polysaccharideintake could lower the apparent absorption of lipid. Total triglyceride, cholesterol,and atherogenic index in blood serum with total lipid and cholesterol levels in liverof polysaccharide group mice were all significantly lower than those of the controlgroup (p <0.05). Furthermore, the effect of the polysaccharide intake on mousecolon bacterial communities was investigated. Mice from the polysaccharide groupshowed a higher colon bacterial diversity than the control group. Bacteroides sp.,Eubacterium sp., butyrate-producing bacteria Butyrivibrio sp., and probioticsBifidobacterium bifidum, Lactobacillus fermentum, and Lactobacillus reuteriin inmouse colon were all increased after polysaccharide intake. These indicated that theintake of polysaccharide from P. asiatica L. could be beneficial for lipid metabolismand colon microbiota.
6. Effects of microwave irradiation on microbial SCFA production and theactivities of extracellular enzymes during in vitro fermentation of the polysaccharidefrom P. asiatica L. were investigated. It was found that the apparent viscosity,average molecular weight, and particle size of the polysaccharide decreased aftermicrowave irradiation. Reducing sugar amount increased with molecular weightdecrease, suggesting the degradation may derive from glycosidic bond rupture. Thepolysaccharide surface topography was changed from large flake like structure tosmaller chips. FT-IR showed that microwave irradiation did not alter the primary functional groups in the polysaccharide. However, SCFA productions of thepolysaccharide during in vitro fermentation significantly increased after microwaveirradiation. Activities of microbial extracellular enzymes xylanase,arabinofuranosidase, xylosidase, and glucuronidase in fermentation culturessupplemented with microwave irradiation treated polysaccharide were also generallyhigher than those of untreated polysaccharide. In addition, physiological propertiesof homogenized and non-homogenized polysaccharide from the seeds of P. asiaticaL., such as the SCFA production were compared. High pressure homogenizationdecreased particle size of the polysaccharide, and changed the surface topographyfrom large flake-like structure to smaller porous chips. FT-IR showed that highpressure homogenization did not alter the primary structure of the polysaccharide.However, the production of total SCFA, propionic acid and n-butyric acid in cecaand colons of mice significantly increased after dieting supplementation withhomogenized polysaccharide. These results showed that microwave irradiation andhigh pressure homogenization treatment could be promising approaches for theproduction of value-added polysaccharides in the food industry.
7. Colonic inflammation was induced in female BALB/c mice by feedingdextran sulfate sodium DSS in drinking water. Along with or after DSSadministration, mice were given polysaccharide (0.2or0.4g/kg body weight) bygavage daily. Polysaccharide administration attenuated development of symptoms(body weight loss, fecal occult blood and diarrhea) associated with colitis.Polysaccharide also blocked colon shortening, increased superoxide dismutase butsuppressed myeloperoxidase and endothelial nitric oxide synthase activities, andimproved macroscopic scores. Histological damage of colon, decrease of thymusorgan index, and increase of C-reactive protein level in mice were reduced bypolysaccharide. Increase of colonic tissue levels of TNF-α, IL-1β, IL-6, macrophageinflammatory protein2and leukotriene B4resulting from colitis were significantlydecreased after polysaccharide administration. Colitis induced decrease of fecalshort-chain fatty acid concentration was also attenuated by polysaccharide.Quantitative PCR (QPCR) results showed that polysaccharide administration couldreduce the decrease of Bifidobacterium, Lactobacilli, Bacteroidaceae and Enterobacteriaceae population sizes in colon of mice with colitis. The putativeclinical utility of the polysaccharide suggests its potential application in colitisattenuation.
8. It was investigated whether pre-administration of polysaccharide from P.asiatica L. could prevent development and colon microbiota changes in mice withexperimental colitis. Colitis was induced in female BALB/c mice by feeding DSS indrinking water.7days before DSS administration, mice were given polysaccharidepre-administration (0.2or0.4g/kg body weight) daily. Polysaccharidepre-administration prevented colitis-induced body weight loss, fecal occult blood anddiarrhea. Polysaccharide blocked colon shortening, suppressed myeloperoxidase, andimproved macroscopic scores. Colon histological damage, decrease in thymus organindex, increase in colon organ index and serum CRP level due to colitis were allreduced by polysaccharide. The increase of colon tissue levels of pro-inflammatorycytokines TNF-α, IL-1β, IL-6, and IL-12, and macrophage inflammatory proteinMIP-2resulting from colitis were all significantly decreased, while level ofanti-inflammatory cytokine IL-4was increased. Furthermore, colitis-induced decreaseof fecal SCFA concentration was attenuated by polysaccharide. QPCR results showedpolysaccharide pre-administration could reduce the decrease of Bifidobacterium,Lactobacilli, Bacteroidaceae and Enterobacteriaceae population sizes and theincrease of Clostridium population size in colon of mice with colitis. The putativeclinical utility of the polysaccharide suggests its potential application in colitisprevention.
引文
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