肉类是膳食的重要组成部分，植物基仿制肉作为肉类的替代品备受关注，其与肉类的营养功能差异尚不明确。食物的消化和营养物质的吸收是决定食物营养价值的重要因素。鉴于此，本研究在体外和小鼠体内系统探究了猪肉、牛肉和对应的植物基仿制肉的消化吸收特性及其对小鼠胃肠消化功能、摄食行为、脂质代谢和结肠稳态的影响，拟从消化、吸收和代谢角度揭示肉和植物基仿制肉对机体健康的影响及相关的营养代谢机理，为肉类和植物基仿制肉类的合理摄入提供理论参考和科学依据。主要的研究结果如下：

1. 猪肉、牛肉及其植物基仿制肉蛋白的体外消化特性研究

本研究首先通过体外模拟胃肠消化，从消化率和消化产物等方面评估猪肉、牛肉、植物基仿制猪肉和植物基仿制牛肉中蛋白质的消化差异，进一步从消化体系的理化特性和蛋白结构的角度，探究肉和植物基仿制肉蛋白消化差异的原因。结果显示：胃阶段消化后，猪肉与植物基仿制猪肉蛋白消化率无显著差异，而牛肉蛋白消化率显著低于植物基仿制牛肉（P < 0.01）。肠阶段消化后，肉蛋白消化率均显著高于相应的植物基仿制肉（P < 0.01）。而且与植物基仿制肉相比，肉消化释放的总肽段、小分子量多肽和潜在的生物活性肽均更多。进一步探究发现，肉和植物基仿制肉消化体系的电荷和离子分布明显不同，尤其是在胃消化阶段它们表现出相反的电位状态（P < 0.01）。肉组消化体系的表观黏度明显低于植物基仿制肉组（P < 0.01）。此外，肉蛋白的总巯基含量和表面疏水性均显著高于植物基仿制肉蛋白（P < 0.01）。且与植物基仿制肉蛋白相比，肉蛋白的α-螺旋含量均较高，但β-折叠、β-转角和无规则卷曲含量较低。这些结果表明，与植物基仿制肉相比，肉消化体系中较低的表观粘度等理化性质以及肉蛋白较高的表面疏水性和较少的β-折叠结构等特性易于消化酶和食物蛋白的更充分作用，从而更容易被消化。

2. 猪肉、牛肉及其植物基仿制肉对小鼠胃肠消化功能的影响和在体内的消化吸收规律

进一步以小鼠为模型，探究这4种膳食的摄入对胃肠消化功能的影响及长期摄入后其在体内的消化吸收规律。结果显示：与植物基仿制肉相比，肉的长期摄入（68天）均显著增加了胃壁细胞的数量，细胞外胃泌素、乙酰胆碱及相应受体以及细胞内Ca2+、钙/钙调蛋白依赖性蛋白激酶II、蛋白激酶C和蛋白激酶A的水平（P < 0.05）。同时，与植物基仿制肉相比，肉的长期摄入增加了H+, K+-ATP酶和胃蛋白酶的活性、十二指肠绒毛高度以及绒毛高度与隐窝深度的比值（P < 0.05）。进一步探究发现，与植物基仿制肉相比，肉在体内消化过程中释放了更多的肽，上调了胃肠道氮营养素感应体（GPR92、GPRC6A、T1R1、T1R2或T1R3）的表达（P < 0.05）。同时，与植物基仿制肉相比，肉的摄入通过降低宿主胃肠道蛋白的分泌和细胞的脱落改变了宿主源和微生物源多肽的组成。此外，猪肉组和植物基仿制牛肉组小鼠空肠中可供吸收的氨基酸含量显著高于植物基仿制猪肉组和牛肉组，但牛肉组中血清氨基酸含量更高（P < 0.05）。这些结果表明，与植物基仿制肉相比，肉中蛋白质在小鼠体内的消化吸收性能更好。长期摄入后，肉释放的更多肽和氨基酸通过上调氮营养素感应体的表达，增强了小鼠的胃肠消化功能，改变了胃肠道内宿主源和微生物源多肽的组成。

3. 猪肉、牛肉及其植物基仿制肉对小鼠摄食行为的影响和相关机制研究

基于以上的研究结果，从胃肠运动和食欲调节两个方面，进一步探究具有不同消化吸收特性的这4种膳食对小鼠摄食行为的影响，并从食物的感官气味特性和消化特性明确其差异原因。结果显示：在胃肠运动方面，与植物基仿制肉相比，肉的长期摄入均显著降低了小鼠十二指肠和空肠的肌肉层厚度以及通过下调胰岛素样生长因子1及其受体（IGF-1和IGF-1R）、干细胞因子（SCF）和肌肉原癌基因受体酪氨酸激酶（c-kit）的表达抑制了胃肠间质细胞的活动（P < 0.01）。同时，与植物基仿制肉相比，肉的摄入通过下调五羟色胺合成酶（TPH1）以及上调其转运体（SERT）和分解代谢酶（Maoa）的表达显著减少了血清中五羟色胺的含量，改变了肠道中五羟色胺受体2A、2B、3A、3B 、4或 7的表达（P < 0.05）。在食欲调节方面，与植物基仿制肉相比，肉的摄入显著降低了外周或下丘脑中促食欲因子（Ghrelin、NPY和Y5R）的水平，但增加了抑食欲因子（PYY、Y2R、GLP-1、GLP-1R、POMC和MC4R）的水平（P < 0.05）。肉摄入引起的这两方面变化增强了饱腹感，进而减少了小鼠的摄食量（P < 0.01）。进一步探究发现，膳食中一些挥发性化合物与食欲调节因子显著相关（P < 0.05）。其中，芳樟醇等挥发性化合物的含量在肉类膳食中显著低于植物基仿制肉（P < 0.05）。KEGG富集分析表明胃和十二指肠内容物中的肽主要通过神经活性配体-受体相互作用通路发挥激素样作用或影响内分泌细胞分泌，进而调节摄食行为。这些结果表明，与植物基仿制肉相比，肉特定的感官气味特性和消化特性通过减缓胃肠排空和改变食欲调节激素间的平衡，增强了饱腹感，进而减少了小鼠的摄食量。

4. 猪肉、牛肉及其植物基仿制肉对小鼠脂质代谢的影响和相关机制研究

为了进一步探究这4种膳食消化吸收后对脂质积累的影响，测定了血清代谢物的组成和脂质代谢相关基因的表达。结果显示：与植物基仿制肉相比，肉的长期摄入均显著降低了脂肪细胞的大小和肝脏组织中的脂质积累。同时，与植物基仿制肉相比，肉的摄入显著降低了血清或脂肪组织中促炎因子（瘦素、抵抗素、F4/80、SAA3、MCP-1、IL6、IL-1β和TNFα）的水平，增加了抗炎因子（脂联素）的水平（P < 0.05）。进一步探究发现，与植物基仿制肉相比，肉的摄入均显著改变了小鼠血清代谢物的组成，下调了肝脏组织中脂质摄取（CD36、FABP4、FATP2或FATP5）和合成（SCD-1、SREBP-1、ACCα或FAS）相关基因的表达，上调了脂酸β氧化（ACSL1、CPT1α、PPARα或PGC-1α）相关基因的表达（P < 0.05）。同时，与植物基仿制肉相比，牛肉的摄入下调了脂肪组织中与脂质合成、分解（ATGL或HSL）和脂肪细胞分化（PPARγ或C/EBPα）相关基因的表达（P < 0.05），猪肉的摄入下调了脂肪分解相关基因的表达，但上调了脂肪细胞分化相关基因的表达（P < 0.05）。尿嘧啶和二十碳五烯酸等特定血清差异代谢物与脂质代谢相关基因的表达显著相关（P < 0.05）。这些代谢物主要参与脂质代谢、蛋白质消化吸收、胆汁分泌和氨基酸代谢等通路。以上结果说明，与植物基仿制肉相比，肉通过特定的血清代谢物直接或间接的调节脂质代谢稳态，进而减少脂质积累，且不同种类的肉间存在一定的差异。

5. 猪肉、牛肉及其植物基仿制肉对小鼠结肠稳态的影响及相关机制研究

膳食中未消化吸收的部分进入大肠后可能影响肠道微生物组成和肠道屏障功能。因此进一步探究了4种膳食对结肠稳态的影响。结果显示：肉和植物基仿制肉的短期摄入（20天）不会引发小鼠结肠组织的病理现象。然而，膳食干预68天后，植物基仿制肉组小鼠结肠组织出现炎症浸润，肉组中没有观察到相似的现象（P < 0.05）。此外，与植物基仿制肉相比，肉的长期摄入通过降低组织细胞的凋亡增加了杯状细胞的数量和粘蛋白的分泌（P < 0.01）。同时，与植物基仿制肉相比，肉的长期摄入通过上调紧密连接蛋白（JAM、ZO-1、Claudin-1或Occludin）和粘附连接蛋白（E-cadherin）的表达以及下调肌球蛋白轻链激酶（MLCK）的表达增强了上皮屏障的完整性（P < 0.01）。进一步探究发现，与植物基仿制肉相比，肉的摄入显著改变了微生物的结构和组成。在门水平上，肉的摄入显著增加了厚壁菌门的相对丰度，降低了拟杆菌门和变形杆菌门等的相对丰度（P < 0.05）。在属水平上，肉的摄入降低了促炎菌（如Bacteroides、Desulfovibrio或Colidextribacter）或致病菌（如Mucispirillum和Helicobacter）的相对丰度，增加了有益菌（如Lactobacillus、Bifidobacterium或Roseburia）的相对丰度（P < 0.05）。而且，与植物基仿制肉相比，肉的摄入显著降低了尸胺、腐胺或精胺等微生物代谢物的水平（P < 0.05）。深入探究发现，与植物基仿制肉相比，肉的长期摄入均显著降低了血清中脂多糖的含量以及结肠组织中促炎通路NF-κB上关键分子（TLR4、MyD88、IKKα/β、IκBα和p65）的基因表达或蛋白磷酸化水平（P < 0.05）。这些结果说明，与植物基仿制肉相比，肉的长期摄入通过降低促炎菌和致病菌以及增加有益菌和减少蛋白质的发酵增强了肠道屏障功能，进而降低了促炎通路的活化，更利于肠道稳态。

Meat is an important part of the diet, and plant-based meat analogues has attracted a lot of interest as a meat substitute. However, the nutritional differences between the two are currently unclear. The nutritional value of food is mostly determined by how well it is digested and how well nutrients are absorbed. In view of this, this study systematically explored the digestive and absorption properties of pork, beef and corresponding plant-based meat analogues in in vitro and mice. Their effects on gastrointestinal digestive function, eating behavior, lipid metabolism and colonic homeostasis in mice were also investigated. The purpose was to reveal the effects of meat and plant-based meat analogues on body health and related nutrient metabolism mechanisms in aspects of digestion, absorption, and metabolism. This will provide a theoretical reference and scientific basis for the reasonable intake of meat and plant-based meat analogues. The main findings in this study are highlighted as follows:

1. Study on in vitro digestion characteristics of pork, beef and their plant-based meat proteins

This study first assessed the variations in protein digestion of pork, beef, plant-based pork analogue, and plant-based beef analogue through in vitro model. To further determine the causes of the variations in protein digestion, the physicochemical characteristics of the digestive system and protein structure were also investigated. The results showed that there was no significant difference between the protein digestibility of pork and plant-based pork analogue after gastric digestion. The digestibility of beef protein was significantly lower than that of plant-based beef analogue after gastric digestion (P < 0.01). After intestinal digestion, the digestibility of meat protein was significantly higher than that of the corresponding plant-based meat analogues (P < 0.01). Moreover, meat digestion releases more total peptides, small molecular weight peptides, and potentially bioactive peptides than plant-based meat analogues. Further exploration found that the charge and ion distributions of meat and plant-based meat analogues digestion systems were significantly different, especially during the gastric digestion stage, which showed opposite potential states (P < 0.01). The apparent viscosity of the digestive system of the meat group was significantly lower than that of the plant-based meat analogue groups (P < 0.01). Total sulfhydryl content and surface hydrophobicity of meat protein were significantly higher than those of plant-based meat protein (P < 0.01). Compared with plant-based meat protein, the α-helix content of meat protein was higher, but the content of β-sheet, β-turn angle and random coil was lower. These results show that the lower apparent viscosity in meat digestion systems, as well as the higher surface hydrophobicity and less β-sheet structure of meat proteins, allow digestive enzymes and food proteins to interaction more fully, making them easier to digest than plant-based meat analogues.

2. Effects of pork, beef and their plant-based meat analogues on gastrointestinal digestive function and digestion and absorption in mice

Furthermore, mice were employed as a model to examine how the four diets affected gastrointestinal digestive function. Also, its digestion and absorption in vivo after prolonged ingestion were investigated. The results showed that the long-term intake (68d) of meat significantly increased the number of gastric parietal cells. The levels of extracellular gastrin, acetylcholine, corresponding receptors, intracellular Ca2+, calcium/calmodulin- dependent protein kinase II, protein kinase C, and protein kinase were also increased as compared to plant-based meat analogues (P < 0.05). Long-term intake of meat also increased the activity of H+, K+-ATPase, and pepsin, duodenal villi height, and the ratio of villi height to crypt depth compared with plant-based meat analogues (P < 0.05). Further investigation found that meat released more peptides during digestion in vivo, upregulating the expression of gastrointestinal nitrogen sensors (GPR92, GPRC6A, T1R1, T1R2, or T1R3) when compared to plant-based meat analogues (P < 0.05). At the same time, meat intake changed the composition of host-derived and microbial-derived peptides by reducing the secretion of host gastrointestinal proteins and cell shedding compared with plant-based meat analogues. In addition, the content of amino acids available for absorption in the jejunum of mice in the pork and the plant-based beef analogue groups was significantly higher than that in the beef and the plant-based pork analogue groups, but the serum amino acid content in the beef group was higher (P < 0.05). These results indicate that meat protein has better digestion and absorption performance in mice than plant-based meat protein. After long-term ingestion, more peptides and amino acids released from meat enhanced the gastrointestinal digestive function of mice by upregulating the expression of nitrogen nutrient sensors, and changed the composition of host-derived and microbial-derived peptides in the gastrointestinal tract.

3. Effects of pork, beef and their plant-based meat analogues on the eating behavior of mice and related mechanisms

Based on the aforementioned research findings, gastrointestinal motility and appetite regulation were further examined to determine the effects of these four diets with variable digestion and absorption properties on the feeding behavior of mice. The reasons for the differences were evaluated through the sensory and digestive characteristics of the diet. The results showed that, in terms of gastrointestinal motility, the long-term intake of meat significantly reduced the muscle layer thickness of the duodenum and jejunum of mice. The activity of gastrointestinal cells of Cajal were also inhibited by downregulating the expression of insulin-like growth factor 1 and its receptors (IGF-1 and IGF-1R), stem cell factor (SCF) and muscle proto-oncogene receptor tyrosine kinase (c-kit) as compared to plant-based meat analogues (P < 0.01). At the same time, meat intake significantly decreased the content of 5-HT in serum by downregulating the expression of 5-HT synthase (TPH1) but upregulating its transporter (SERT) and catabolic enzyme (Maoa), and changed the expression of 5-HT receptors 2A, 2B, 3A, 3B, 4 or 7 in the intestine (P < 0.05). In terms of appetite regulation, meat intake significantly reduced levels of appetite-stimulating factors (ghrelin, NPY, and Y5R) in the peripheral or hypothalamus but increased levels of appetite-suppressing factors (PYY, Y2R, GLP-1, GLP-1R, POMC, and MC4R) compared to plant-based meat analogues (P < 0.05). These two changes caused by meat intake enhanced satiety, which further reduced the food intake of mice (P < 0.01). Further exploration found that some volatile compounds in the diet were significantly associated with appetite regulators (P < 0.05). Among them, the content of volatile compounds such as linalool in the meat diets was significantly lower than that of plant-based meat analogues (P < 0.05). KEGG enrichment analysis showed that peptides in gastric and duodenal contents mainly exert hormone-like effects or affect endocrine cell secretion through neuroactive ligand-receptor interaction pathways, thereby regulating feeding behavior. These results suggest that meat-specific organoleptic odor properties and digestive properties enhance satiety by slowing gastrointestinal emptying and altering the balance between appetite-regulating hormones compared to plant-based meat analogues, thereby reducing food intake in mice.

4. Effects of pork, beef and their plant-based meat analogues on lipid metabolism and related mechanisms in mice

In order to further explore the effects of four diets on lipid accumulation, the composition of serum metabolites and the expression of lipid metabolism-related genes were measured. The findings showed that meat intake significantly reduced the size of fat cells and lipid buildup in liver tissue compared with plant-based meat analogues. Simultaneously, meat intake significantly reduced the levels of pro-inflammatory factors (leptin, resistin, F4/80, SAA3, MCP-1, IL6, IL-1β, and TNFα) and increased the level of the anti-inflammatory factor (adiponectin) in serum or adipose tissue (P < 0.05). Further exploration found that meat intake significantly changed the composition of serum metabolites and downregulated the expression of genes related to lipid uptake (CD36, FABP4, FATP2 or FATP5) and synthesis (SCD-1, SREBP-1, ACCα or FAS) but upregulated the expression of genes related to lipid oxidation (ACSL1, CPT1α, PPARα or PGC-1α) in the liver compared with plant-based meat analogues in mice (P < 0.05). At the same time, beef intake downregulated the expression of genes related to lipid synthesis, lipolysis (ATGL or HSL) and adipocyte differentiation (PPARγ or C/EBPα) in adipose tissue compared with plant-based beef analogue (P < 0.05). Pork intake downregulated the expression of genes related to lipolysis and upregulated the expression of genes related to adipocyte differentiation compared with plant-based pork analogue (P < 0.05). Specific serum differential metabolites, such as uracil and eicosapentaenoic acid, were significantly associated with the expression of lipid metabolism-related genes (P < 0.05). These metabolites were mainly involved in lipid metabolism, protein digestion and absorption, bile secretion, and amino acid metabolism. The above results show that meat directly or indirectly regulates lipid metabolism homeostasis through specific serum metabolites compared with plant-based meat analogues, which further reduces lipid accumulation. There are certain differences between different meat types.

5. Effects of pork, beef and their plant-based meat analogues on colonic homeostasis and related mechanisms in mice

The undigested and unabsorbed parts of the diet entering the large intestine may affect gut microbial composition and intestinal barrier function. Therefore, the effects of four diets on colonic homeostasis were further explored. The results showed that short-term intake (20d) of meat and plant-based meat analogues did not cause pathological phenomena. However, long-term consumption (68d) of plant-based meat analogues showed inflammatory infiltration in the colonic tissue of mice, which was not observed in the meat groups (P < 0.05). In addition, long-term intake of meat increased the number of goblet cells and mucin secretion by reducing apoptosis of histiocytes compared with plant-based meat analogues (P < 0.01). At the same time, long-term intake of meat enhanced the integrity of the epithelial barrier. As it upregulated the expression of tight junction proteins (JAM, ZO-1, Claudin-1 or Occludin) and adherens junction protein (E-cadherin) and downregulated the expression of myosin light chain kinase (MLCK) compared with plant-based meat analogues (P < 0.01). Further investigation found that meat intake significantly changed the structure and composition of gut microbiota as compared to plant-based meat analogues. At the phylum level, meat intake significantly increased the relative abundance of Firmicutes but decreased the relative abundance of Bacteroides and Proteobacteria (P < 0.05). At the genus level, meat intake reduced the relative abundance of pro-inflammatory bacteria (such as Bacteroides, Desulfovibrio or Colidextribacter) or pathogenic bacteria (such as Mucispirillum and Helicobacter) (P < 0.05). It also increased the relative abundance of beneficial bacteria (such as Lactobacillus, Bifidobacterium or Roseburia) (P < 0.05). Moreover, meat intake decreased the levels of microbial metabolites such as cadaverine, putrescine or spermine as compared to plant-based meat analogues (P < 0.05). In-depth exploration showed that long-term intake of meat significantly reduced the content of lipopolysaccharides in serum as well as the gene expression or phosphorylation levels of key molecules (TLR4, MyD88, IKKα/β, IκBα and p65) on the pro-inflammatory pathway NF-κB in colonic tissue compared with plant-based meat analogues (P < 0.05). These results indicate that long-term intake of meat enhances the intestinal barrier function by reducing pro-inflammatory and pathogenic bacteria, increasing beneficial bacteria, and reducing the fermentation of protein, thereby reducing the activation of pro-inflammatory pathways and facilitating intestinal homeostasis.

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