Effects of Rhodopseudomonas palustris on Growth Performance, Antioxidant Capacity, Muscle Quality, and Liver and Intestine Health of Largemouth Bass (Micropterus salmoides)

ZHENG Jieqiu; FU Bing; XING Dongxu; LI Qingrong; ZOU Yuxiao; ZHOU Donglai

doi:10.12418/CJAN2026.231

2026 , Vol. 38 >Issue 4: 2866 - 2885

DOI: https://doi.org/10.12418/CJAN2026.231

RESEARCH PAPER

Effects of Rhodopseudomonas palustris on Growth Performance, Antioxidant Capacity, Muscle Quality, and Liver and Intestine Health of Largemouth Bass (Micropterus salmoides)

ZHENG Jieqiu ^,¹^,² ,
FU Bing ² ,
XING Dongxu ² ,
LI Qingrong ² ,
ZOU Yuxiao ^,¹^,²^,^* ,
ZHOU Donglai ^,²^,^*

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¹ College of Light Industry and Food Sciences, Zhongkai University of Agriculture and Engineering, Guangzhou 510225, China
² Guangdong Key Laboratory of Agricultural Products Processing, Sericultural & Agri-Food Research Institute, Guangdong Academy of Agricultural Sciences, Guangzhou 510640, China

*ZOU Yuxiao, professor, E-mail: yuxiaozou@163.com;

ZHOU Donglai, associate professor, E-mail: zhoudonglai.2007@163.com

Received date: 2025-09-18

Online published: 2026-04-14

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Abstract

This experiment aimed to investigate the effects of different dietary supplementation methods of Rhodopseudomonas palustris on the growth performance, antioxidant capacity, muscle quality, and liver and intestine health of largemouth bass (Micropterus salmoides). A total of 270 largemouth bass with an initial body weight of (106.73±0.33) g were randomly divided into 3 groups, with 3 replicates per group and 30 fish per replicate. The control group (CK group) was fed a basal diet, and the experimental groups were fed the basal diets supplemented with Rhodopseudomonas palustris by spraying live bacteria (MG group, live bacterial count of 2.25×10⁷ CFU/mL) and by spraying bacteria and then drying (DG group, dead bacteria), respectively. The experiment lasted for 61 days. The results showed as follows: 1) compared with CK group, the condition factor in MG and DG groups was significantly decreased (P<0.05). Meanwhile, the weight gain rate in DG group was significantly lower than that in MG group (P<0.05), and the feed coefficient was significantly higher than that in MG group (P<0.05). 2) Compared with CK group, the muscle crude fat content in DG group was significantly decreased (P<0.05); the contents of polyunsaturated fatty acids in muscle in MG and DG groups were significantly increased (P<0.05), the contents of eicosapentaenoic acid and docosahexaenoic acid in muscle were significantly increased (P<0.05), and the muscle hardness, elasticity, cohesion and resilience were significantly increased (P<0.05); the muscle chewiness and adhesiveness in DG group were significantly increased (P<0.05); the myofiber density in MG and DG groups was significantly increased (P<0.05), and the myofiber diameter in DG group was significantly decreased (P<0.05). 3) Compared with CK group, the serum reduced glutathione (GSH) content and activities of catalase (CAT) and superoxide dismutase in MG and DG groups were significantly increased (P<0.05); the activities of serum alkaline phosphatase (AKP) and acid phosphatase (ACP) in MG group were significantly increased (P<0.05), and the serum malondialdehyde (MDA) content and activities of aspartate aminotransferase and alanine aminotransferase were significantly decreased (P<0.05); the serum total antioxidant capacity (T-AOC) and lysozyme activity in DG group were significantly increased (P<0.05), and the activities of serum AKP and ACP were significantly decreased (P<0.05). 4) Compared with CK group, the liver T-AOC in MG and DG groups was significantly increased (P<0.05), the liver GSH content in MG group was significantly increased (P<0.05), and the liver CAT activity and MDA content in MG and DG groups were significantly decreased (P<0.05). 5) Compared with CK group, the activities of intestinal trypsin and lipase in MG group were significantly increased (P<0.05); the intestinal villus height, villus width and muscular thickness in MG and DG groups were significantly increased (P<0.05); the intestinal Pseudomonadota relative abundance in MG group was increased, the Mesomycoplasma relative abundance was decreased, and the Acinetobacter relative abundance was increased; the intestinal Bacillota relative abundance in DG group was increased, and the relative abundances of Sphingomonas and Pseudomonas were also increased. In conclusion, dietary supplementation of Rhodopseudomonas palustris can enhance the muscle quality and antioxidant capacity of largemouth bass, and improve the liver and intestine health. Under the conditions of this experiment, the effect of live Rhodopseudomonas palustris was better.

Key words： Rhodopseudomonas palustris; largemouth bass (Micropterus salmoides); antioxidant capacity; muscle quality; liver health; intestine health

Cite this article

ZHENG Jieqiu , FU Bing , XING Dongxu , LI Qingrong , ZOU Yuxiao , ZHOU Donglai . Effects of Rhodopseudomonas palustris on Growth Performance, Antioxidant Capacity, Muscle Quality, and Liver and Intestine Health of Largemouth Bass (Micropterus salmoides)[J]. Chinese Journal of Animal Nutrition, 2026 , 38(4) : 2866 -2885 . DOI: 10.12418/CJAN2026.231

大口黑鲈(Micropterus salmoides)又称加州鲈,是我国重要的淡水经济养殖鱼类之一,其因具有生长周期短、经济价值高、肉质鲜美和无肌间刺等优势,备受养殖者和消费者青睐^[1-2]。为了追求更高的经济效益,目前养殖户多采用集约化高密度养殖模式,这导致水质恶化、病害频发和肉品质下降等问题,严重制约了产业的可持续发展^[3-4]。目前,病害防治高度依赖抗生素,但长期使用抗生素不仅导致病原菌耐药性增强,破坏鱼类肠道微生态平衡,而且这些药物通过食物链富集后会影响食品安全,危害人体健康^[5-6]。研究表明,在饲料中合理补充益生菌^[7]、益生元^[8]或植物活性成分^[9]等添加剂有助于改善水产动物肠道健康,并且对水产动物的生长性能、抗氧化能力和营养品质等具有明显的促进作用^[10-12]。因此,益生菌已经成为水产养殖业中用来促进动物生长、增强动物免疫力的重要选择^[13-14]。

沼泽红假单胞菌(Rhodopseudomonas palustris)属于红螺菌科(Rhodospirillaceae)、红假单胞菌属(Rhodopseudomonas),为革兰阴性、紫色非硫细菌。沼泽红假单胞菌不仅代谢功能强大,而且营养丰富,富含多种氨基酸、维生素、类胡萝卜素和辅酶Q等多种营养成分^[15-16],具有多种生物学功能,作为益生菌中的代表菌种被广泛应用于水产养殖业^[14,17]。研究表明,沼泽红假单胞菌不仅能够提高鱼体生长性能^[17],还可以增强鱼体的抗氧化能力^[18]、免疫力和抗病性^[19]。研究发现,饲料添加沼泽红假单胞菌可以改善罗非鱼(Oreochromis niloticus)生长性能和抗氧化能力^[20],促进皱纹盘鲍(Haliotis discus hannai)生长并提高其存活率^[19],增强凡纳滨对虾(Litopenaeus vannamei)的抗病性^[21]。

近年来,沼泽红假单胞菌作为水质净化剂或饲料添加剂广泛应用在水产养殖中,但目前的研究多为直接拌料活菌或者泼洒到养殖水体中,并且多数研究更多关注水质和鱼类的生长性能等方面^[14,17],沼泽红假单胞菌的不同添加方式对鱼的健康和肉品质的影响研究甚少,而鱼肉品质是消费者最为关心的。因此,本研究旨在探究饲料中沼泽红假单胞菌的不同添加方式对大口黑鲈生长性能、抗氧化能力、肌肉品质以及肝脏和肠道健康的影响,以期为沼泽红假单胞菌在水产养殖中的合理应用提供参考。

1 材料与方法

1.1 试验设计和饲料制备

选取270尾初始体重为(106.73±0.33) g的大口黑鲈,随机分为3组,分别命名为对照组(CK组)、拌料组(MG组)和烘干组(DG组),每组3个重复,每个重复30尾鱼;试验鱼随机分配到9个直径为75 cm的养殖桶中(养殖水体为250 L)。CK组饲喂基础饲料(商品配合饲料,广东杰大饲料有限公司提供),试验组分别饲喂在基础饲料上喷洒沼泽红假单胞菌(MG组,活菌)和喷洒沼泽红假单胞菌并烘干(DG组,死菌)的试验饲料,基础饲料营养水平见表1。沼泽红假单胞菌为本实验室提供,活菌数为2.25×10⁷ CFU/mL。试验期61 d。

表1 基础饲粮营养水平(风干基础)

Table 1 Nutrient levels of the basal diet (air-dry basis) %

项目 Item	粗蛋白质 Crude protein	粗脂肪 Crude lipid	粗纤维 Crude fiber	粗灰分 Crude ash	总磷 Total phosphorus	赖氨酸 Lysine	水分 Moisture
含量Content	49.23	10.76	1.48	17.23	1.21	3.64	8.06

	营养水平为实测值。
	Nutrient levels were measured values.

各组饲料制作如下:MG组,每1 000 g基础饲料添加30 mL沼泽红假单胞菌菌液,将菌液均匀喷洒在基础饲料表面,充分搅拌混合均匀,常温风干后待用,现用现配;DG组,每1 000 g基础饲料添加30 mL沼泽红假单胞菌菌液,将菌液均匀喷洒在基础饲料表面,充分搅拌混合均匀后,平铺于烘箱,60 ℃烘干(2 h)后4 ℃储存待用;CK组,每1 000 g基础饲料添加30 mL无菌水,将无菌水均匀喷洒在基础饲料表面,充分搅拌混合均匀,常温风干后待用,现用现配。

1.2 饲养管理

试验所用大口黑鲈为市售,试验鱼先在广东省农业科学院蚕业与农产品加工研究所室内循环水养殖系统中暂养2周。正式试验前,停料24 h,按照试验设计准备试验鱼。试验期间,每天投喂2次(09:00和16:30各1次),饱食投喂。将投喂0.5 h后未摄食完的饲料捞出,烘干水分后称重,准确记录每天饲料采食量和试验鱼死亡情况。养殖期间水温为25~28 ℃,溶氧量为≥5 mg/L,氨氮含量≤0.2 mg/L,pH为7.0~8.2,亚硝酸盐含量≤0.05 mg/L,自然光照周期。动物试验经广东省农业科学院动物福利伦理委员会批准(批准号:GDAAS2022015)。

1.3 样品采集

试验结束后,试验鱼禁食24 h,记录每桶中存活鱼数,计算成活率。每个重复随机选取8尾鱼,测量体长、体重后尾静脉采血,4 ℃静置2 h,然后4 ℃、1 324×g离心10 min,取上清液置于-80 ℃保存,用于血清生化和抗氧化指标的测定。随后将试验鱼在冰上解剖,分离内脏团、肝脏并称重,计算脏体比(VSI)和肝体比(HSI)。每组随机选取3尾鱼的中肠(1 cm)和肝脏,一部分置于4%多聚甲醛,室温保存,用于肠道和肝脏组织形态学检测;另取部分中肠和肝脏样品,用液氮速冻后于-80 ℃保存,用于肠道消化酶活性和肝脏抗氧化指标测定;同时收集后肠液氮速冻后于-80 ℃保存,用于肠道菌群测序分析。背肌去皮后,取肌肉块(0.5 cm×0.5 cm×0.5 cm),完全浸没于装有4%多聚甲醛的离心管中,室温保存,用于肌肉纤维形态学测定;剩余背肌一部分用于质构特性分析,另一部分液氮速冻后于-80 ℃保存,用于肌肉营养成分含量测定。

1.4 测定指标及方法

1.4.1 饲料营养成分

饲料中粗蛋白质、粗脂肪、粗纤维、粗灰分、总磷、赖氨酸和水分含量分别参照GB/T 6432—2018、GB/T 6433—2006、GB/T 6434—2022、GB/T 6438—2007、GB/T 6437—2018、GB/T 18246—2019和GB/T 6435—2014中方法进行测定。

1.4.2 生长性能

生长性能相关指标计算公式如下:

增重率(WGR,%)=100×(终末体重-初始体重)/初始体重;

特定生长率(SGR,%/d)=100×(ln终末体重-ln初始体重)/饲养天数;

摄食率(FR,%/d)=100×摄食量(FI)/[(初始体重+终末体重)/2]/饲养天数;

FI(g/尾)=摄食饲料总量/[(初始尾数+终末尾数)/2];

饲料系数(FCR)=摄食饲料总量/(终末体重-初始体重);

肥满度(CF,g/cm³)=100×体重(g)/体长(cm)³;

VSI(%)=100×内脏团重量(g)/体重(g);

HSI(%)=100×肝脏重量(g)/体重(g)。

1.4.3 肌肉营养成分含量

称取每组样品适量,平行测定6次。肌肉水分含量按照《食品中水分的测定》(GB 5009.3—2016)中的直接干燥法测定;粗灰分含量按照《食品中灰分的测定》(GB 5009.4—2016)中的马弗炉高温灼烧法测定;粗蛋白质含量按照《食品中蛋白质的测定》(GB 5009.5—2016)中的凯氏定氮法测定;粗脂肪含量按照《食品中脂肪的测定》(GB 5009.6—2016)中的索氏抽提法测定;脂肪酸组成及含量按照《食品中脂肪酸的测定》(GB 5009.168—2016)中方法测定,氨基酸组成及含量按照《食品中氨基酸的测定》(GB 5009.124—2016)中方法测定。

1.4.4 肌肉质构特性分析

肌肉质构特性分析参照本实验室前期建立的方法^[22]并稍作修改。将大口黑鲈背肌切成2.0 cm×2.0 cm×1.5 cm的小块后,采用TA.XT Plus物性分析仪(Stable Micro System,英国)进行质构特性分析,探头为P50探头,参数设定为:测前速度、测试速度和测后速度均为1.0 mm/s,应变比为65%,测定间隔时间为5 s,数据获得速率为200 pps,测试温度为常温,压缩次数为2次。每个平行取6尾鱼,每尾鱼分别取3块肌肉进行测定。

1.4.5 肌肉、肝脏和肠道组织形态

肌肉、肝脏和肠道组织经4%的多聚甲醛固定24 h,通过苏木精-伊红(HE)染色后,采用全景切片扫描仪(PANNORAMIC-1000,3DHISTECH)采集切片照片,通过CaseViewer 2.4软件(3DHISTECH)截取照片,利用Image-Pro Plus 6.0(Media Cybernetics)软件测量肠道绒毛高度、绒毛宽度和肌层厚度,每个切片分别测量6组数值。

1.4.6 血清生化和抗氧化指标

将-80 ℃冻存的血清样品在冰上解冻,按照南京建成生物工程研究所试剂盒说明书的操作步骤对血清样品进行生化和抗氧化指标的测定,指标包括总抗氧化能力(T-AOC)以及还原型谷胱甘肽(GSH)、丙二醛(MDA)、过氧化氢酶(CAT)、超氧化物歧化酶(SOD)、碱性磷酸酶(AKP)、酸性磷酸酶(ACP)、溶菌酶(LZM)、谷草转氨酶(AST)和谷丙转氨酶(ALT)含量或活性。

1.4.7 肝脏抗氧化指标

将储存于-80 ℃的肝脏组织在冰上解冻,准确称取组织按重量(g)∶体积(mL)=1∶9的比例加入4 ℃生理盐水后机械匀浆,并在4 ℃、573×g下离心10 min,取上清液进行抗氧化指标测定,指标包括T-AOC以及GSH、MDA、CAT和SOD含量或活性,所用试剂盒均购自南京建成生物工程研究所,操作步骤按照说明书进行。

1.4.8 肠道消化酶活性

将肠道样品在冰上解冻,准确称取组织重量,按重量(g)∶体积(mL)=1∶9的比例加入4 ℃生理盐水后机械匀浆,并在4 ℃、573×g下离心10 min,取上清液,进行肠道α-淀粉酶(AMS)、脂肪酶(LPS)和胰蛋白酶(TRY)活性的测定,所用试剂盒均购自南京建成生物工程研究所,操作步骤按照说明书进行。

1.4.9 肠道菌群测序分析

肠道菌群测序由上海美吉生物医药科技有限公司进行。从大口黑鲈肠道中提取基因组DNA后,采用携带barcode的特异引物扩增16S rRNA基因的目标区域,使用DNA凝胶回收纯化试剂盒(PCR Clean-Up Kit,中高逾华生物技术有限公司)对回收产物进行纯化,并用Qubit 4.0(Thermo Fisher Scientific,美国)对回收产物进行检测定量,利用Illumina NextSeq 2000平台进行测序。测序结束后采用FASTP和FLASH软件对reads进行过滤和拼接,使用QIIME 2流程中的Deblur插件进行降噪处理,得到扩增子序列变异(ASVs),基于Silva 16S rRNA基因数据库(v 138.2),使用QIIME 2中的vsearch分类器进行物种分类学分析,采用mothur软件进行alpha多样性和群落组成分析。

1.5 数据统计分析

试验数据采用SPSS 27.0软件进行单因素方差分析(one-way ANOVA),并采用Duncan氏法进行组间差异显著性的多重比较检验,结果数据采用“平均值±标准差(mean±SD)”形式表示,P<0.05为差异显著。

2 结果与分析

2.1 沼泽红假单胞菌对大口黑鲈生长性能的影响

由表2可知,与CK组相比,MG组和DG组大口黑鲈CF均显著降低(P<0.05)。与MG组相比,DG组WGR显著降低(P<0.05),FCR显著提高(P<0.05)。各组间终末体重、SGR、FR、FI、SR、HSI和VSI均无显著差异(P>0.05)。

表2 沼泽红假单胞菌对大口黑鲈生长性能的影响

Table 2 Effects of Rhodopseudomonas palustris on growth performance of largemouth bass (Micropterus salmoides)

项目 Items	组别Groups
项目 Items	CK	MG	DG
初始体重IBW/g	106.37±1.34	106.79±1.42	107.02±1.58
终末体重FBW/g	231.92±6.80	232.80±4.94	221.05±5.69
增重率WGR/%	112.41±2.08^ab	113.61±3.96^a	105.15±5.28^b
特定生长率SGR/(%/d)	1.24±0.02	1.24±0.03	1.18±0.04
摄食率FR/(%/d)	1.22±0.02	1.19±0.02	1.21±0.01
摄食量FI/(g/尾)	124.42±1.10	123.83±1.00	125.68±1.31
饲料系数FCR	1.03±0.02^ab	1.01±0.04^b	1.07±0.03^a
成活率SR/%	98.89±1.92	98.89±1.92	100.00±0.00
肥满度CF/(g/cm³)	2.98±0.37^a	2.47±0.11^b	2.47±0.10^b
肝体比HSI/%	1.72±0.14	1.87±0.18	1.80±0.19
脏体比VSI/%	6.19±0.34	6.20±0.31	6.34±0.25

	同行数据肩标不同字母表示差异显著(P<0.05),相同字母或无字母表示差异不显著(P>0.05)。下表同。
	In the same row, values with different letter superscripts mean significant difference (P<0.05), while with no letter or the same letter superscripts mean no significant difference (P>0.05). The same as below.

2.2 沼泽红假单胞菌对大口黑鲈肌肉品质的影响

2.2.1 沼泽红假单胞菌对大口黑鲈肌肉营养成分含量的影响

由表3可知,与CK组相比,DG组大口黑鲈肌肉粗脂肪含量显著降低(P<0.05),MG组和DG组肌肉水分含量显著降低(P<0.05)。与MG组相比,DG组肌肉粗脂肪含量显著降低(P<0.05),肌肉粗蛋白质含量显著提高(P<0.05)。各组间肌肉粗灰分含量无显著差异(P>0.05)。

表3 沼泽红假单胞菌对大口黑鲈肌肉常规营养成分含量的影响(鲜样基础)

Table 3 Effects of Rhodopseudomonas palustris on muscle conventional nutrient contents of largemouth bass (Micropterus salmoides) (fresh sample basis) %

项目 Items	组别Groups
项目 Items	CK	MG	DG
粗脂肪Crude lipid	5.84±0.48^a	5.38±0.17^a	4.11±0.75^b
粗蛋白质Crude protein	26.67±0.96^ab	25.60±0.47^b	27.25±0.46^a
粗灰分Crude ash	1.32±0.14	1.18±0.05	1.19±0.19
水分Moisture	72.09±0.57^a	70.69±0.55^b	70.77±0.14^b

由表4可知,在大口黑鲈肌肉中共检测出22种脂肪酸,其中9种饱和脂肪酸,13种不饱和脂肪酸,包括6种单不饱和脂肪酸和7种多不饱和脂肪酸。与CK组相比,MG组和DG组肌肉单不饱和脂肪酸含量显著降低(P<0.05),肌肉多不饱和脂肪酸含量显著提高(P<0.05);DG组肌肉总不饱和脂肪酸含量显著降低(P<0.05),肌肉总饱和脂肪酸含量显著提高(P<0.05)。此外,与CK组相比,MG组和DG组肌肉二十碳五烯酸(EPA)和二十二碳六烯酸(DHA)含量显著提高(P<0.05)。

表4 沼泽红假单胞菌对大口黑鲈肌肉脂肪酸组成及含量的影响

Table 4 Effects of Rhodopseudomonas palustris on muscle fatty acid composition and contents of largemouth bass (Micropterus salmoides) %

项目 Items	组别Groups
项目 Items	CK	MG	DG
月桂酸C12∶0^*	0.04±0.00^a	0.04±0.00^ab	0.03±0.00^b
豆蔻酸C14∶0^*	1.71±0.06^a	1.53±0.04^b	1.39±0.09^c
豆蔻一烯酸C14∶1^#	0.03±0.00^a	0.03±0.00^a	0.02±0.00^b
十五烷酸C15∶0^*	0.17±0.00^a	0.16±0.00^b	0.15±0.00^b
棕榈酸C16∶0^*	21.60±0.10^b	21.63±0.06^b	22.27±0.15^a
棕榈油酸C16∶1^#	3.82±0.14^a	3.72±0.10^a	2.97±0.24^b
十七烷酸C17∶0^*	0.16±0.01^b	0.17±0.01^b	0.20±0.02^a
硬脂酸C18∶0^*	4.49±0.17^b	4.89±0.14^b	5.56±0.31^a
油酸C18∶1^#	33.80±0.79^a	30.86±0.65^b	28.33±1.12^c
亚油酸C18∶2^▲	18.51±0.19^a	17.79±0.09^c	18.10±0.10^b
亚麻酸C18∶3^▲	1.96±0.05^a	1.71±0.05^b	1.66±0.09^b
花生酸C20∶0^*	0.17±0.00	0.18±0.00	0.18±0.02
二十碳一烯酸C20∶1^#	1.12±0.01^a	1.00±0.03^c	1.04±0.02^b
二十碳二烯酸C20∶2^▲	0.53±0.02^b	0.56±0.00^b	0.67±0.04^a
二十碳三烯酸C20∶3^▲	0.32±0.01^b	0.37±0.01^a	0.39±0.01^a
二十碳四烯酸C20∶4^▲	1.56±0.13^c	2.02±0.09^b	2.31±0.18^a
山嵛酸C22∶0^*	0.11±0.00^b	0.10±0.00^b	0.12±0.00^a
芥酸C22∶1^#	0.16±0.01^c	0.21±0.00^b	0.23±0.02^a
二十碳五烯酸C20∶5^▲	1.34±0.08^c	1.58±0.06^b	1.71±0.04^a
木焦油酸C24∶0^*	0.08±0.01^b	0.10±0.00^a	0.10±0.00^a
二十四碳一烯酸C24∶1^#	0.31±0.03^b	0.44±0.03^a	0.48±0.04^a
二十二碳六烯酸C22∶6^▲	7.97±0.69^b	10.89±0.58^a	12.07±0.85^a
单不饱和脂肪酸MUFA	39.23±0.91^a	36.26±0.75^b	33.09±1.30^c
多不饱和脂肪酸PUFA	32.19±0.71^c	34.93±0.62^b	36.90±1.02^a
总不饱和脂肪酸TUFA	71.43±0.23^a	71.19±0.13^a	69.99±0.30^b
总饱和脂肪酸TSFA	28.54±0.22^b	28.80±0.13^b	30.00±0.33^a

	*:饱和脂肪酸;#:单不饱和脂肪酸;▲:多不饱和脂肪酸。
	*: saturated fatty acids; #: monounsaturated fatty acids; ▲: polyunsaturated fatty acids.

由表5可知,在大口黑鲈肌肉中共检测出16种氨基酸,其中8种必需氨基酸,2种半必需氨基酸,6种非必需氨基酸(3种属于鲜味氨基酸)。与CK组相比,DG组肌肉总氨基酸、必需氨基酸、半必需氨基酸、非必需氨基酸和鲜味氨基酸含量均显著降低(P<0.05),MG组肌肉上述氨基酸含量无显著差异(P>0.05)。

表5 沼泽红假单胞菌对大口黑鲈肌肉氨基酸组成及含量的影响

Table 5 Effects of Rhodopseudomonas palustris on muscle amino acid composition and contents of largemouth bass (Micropterus salmoides) %

项目 Items	组别Groups
项目 Items	CK	MG	DG
丝氨酸Ser^#	0.88±0.01^a	0.87±0.02^a	0.74±0.02^b
谷氨酸Glu^#Δ	3.35±0.05^a	3.33±0.13^a	2.84±0.05^b
天冬氨酸Asp^#Δ	2.23±0.01^a	2.23±0.07^a	1.89±0.04^b
甘氨酸Gly^#Δ	1.18±0.01^a	1.06±0.04^b	0.89±0.02^c
丙氨酸Ala^#	1.36±0.01^a	1.31±0.02^b	1.06±0.02^c
脯氨酸Pro^#	0.79±0.00^a	0.72±0.00^c	0.75±0.02^b
苏氨酸Thr^*	0.97±0.00^a	0.97±0.03^a	0.81±0.02^b
缬氨酸Val^*	1.07±0.01^a	1.06±0.04^a	0.85±0.02^b
蛋氨酸Met^*	0.44±0.02^a	0.48±0.05^a	0.33±0.01^b
异亮氨酸Ile^*	0.99±0.01^a	1.01±0.03^a	0.79±0.01^b
亮氨酸Leu^*	1.76±0.01^a	1.75±0.06^a	1.39±0.02^b
苯丙氨酸Phe^*	0.89±0.01^a	0.89±0.03^a	0.72±0.01^b
赖氨酸Lys^*	2.03±0.02^a	2.04±0.08^a	1.72±0.01^b
组氨酸His^*	0.54±0.01^a	0.54±0.02^a	0.42±0.00^b
酪氨酸Tyr^▲	0.68±0.00^a	0.69±0.03^a	0.51±0.01^b
精氨酸Arg^▲	1.30±0.02^a	1.27±0.02^a	1.00±0.03^b
总氨基酸TAA	20.46±0.09^a	20.24±0.62^a	16.70±0.28^b
必需氨基酸EAA	8.69±0.01^a	8.75±0.34^a	7.03±0.08^b
半必需氨基酸SEAA	1.98±0.02^a	1.97±0.05^a	1.51±0.04^b
非必需氨基酸NEAA	9.79±0.07^a	9.51±0.24^a	8.17±0.17^b
鲜味氨基酸FAA	6.76±0.05^a	6.62±0.20^a	5.61±0.11^b

	*:必需氨基酸;▲:半必需氨基酸;#:非必需氨基酸;Δ:鲜味氨基酸。
	*: essential amino acids; ▲: semi-essential amino acids; #: non-essential amino acids; Δ: flavor amino acids.

2.2.2 沼泽红假单胞菌对大口黑鲈肌肉质构特性的影响

由表6可知,与CK组相比,MG组和DG组大口黑鲈肌肉硬度、弹性、内聚力和回弹性显著提高(P<0.05),DG组肌肉咀嚼性和胶黏性显著提高(P<0.05),而MG组肌肉咀嚼性和胶黏性无显著差异(P>0.05)。

表6 沼泽红假单胞菌对大口黑鲈肌肉质构特性的影响

Table 6 Effects of Rhodopseudomonas palustris on muscle texture characteristics of largemouth bass (Micropterus salmoides)

项目 Items	组别Groups
项目 Items	CK	MG	DG
硬度Hardness/g	10 424.95±314.91^b	11 833.23±272.29^a	12 189.74±173.16^a
咀嚼性Chewiness/g	2 334.58±83.54^b	2 334.50±47.57^b	2 504.90±23.72^a
弹性Elasticity/mm	0.42±0.02^c	0.52±0.02^b	0.58±0.03^a
胶黏性Adhesiveness/g	4 398.81±154.62^b	4 693.78±199.58^b	5 827.24±126.54^a
内聚力Cohesion	0.36±0.03^b	0.43±0.02^a	0.45±0.02^a
回弹性Resilience	0.21±0.02^b	0.24±0.01^a	0.24±0.01^a

2.2.3 沼泽红假单胞菌对大口黑鲈肌肉纤维形态的影响

如图1所示,CK组大口黑鲈肌肉纤维间隙较大,排列松散;饲料添加沼泽红假单胞菌后纤维间隙变小,排列紧密,肌纤维密度变高。进一步统计分析发现(表7),与CK组相比,MG组和DG组肌纤维密度均显著提高(P<0.05),DG组肌纤维直径显著降低(P<0.05)。

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图1 沼泽红假单胞菌对大口黑鲈肌肉纤维形态的影响

A~C:横切面肌肉纤维结构(20×)。A:CK组;B:MG组;C:DG组。

Fig.1 Effects of Rhodopseudomonas palustris on myofiber morphology of largemouth bass (Micropterus salmoides)

A to C: myofiber structure of cross sections (20×). A: CK group; B: MG group; C: DG group.

表7 沼泽红假单胞菌对大口黑鲈肌肉纤维形态的影响

Table 7 Effects of Rhodopseudomonas palustris on myofiber morphology of largemouth bass (Micropterus salmoides)

项目 Items	组别Groups
项目 Items	CK	MG	DG
肌纤维直径Myofiber diameter/μm	148.02±5.81^a	136.09±3.06^ab	127.19±6.92^b
肌纤维密度Myofiber density/(个/mm²)	106.23±1.66^b	131.37±3.45^a	127.47±8.13^a

2.3 沼泽红假单胞菌对大口黑鲈血清生化和抗氧化指标的影响

由表8可知,与CK组相比,MG组和DG组大口黑鲈血清GSH含量以及CAT和SOD活性均显著提高(P<0.05);MG组血清AKP和ACP活性显著提高(P<0.05),血清MDA含量以及AST和ALT活性显著降低(P<0.05);DG组血清T-AOC和LZM活性显著提高(P<0.05),血清AKP和ACP活性显著降低(P<0.05)。

表8 沼泽红假单胞菌对大口黑鲈血清生化和抗氧化指标的影响

Table 8 Effects of Rhodopseudomonas palustris on serum biochemical and antioxidant indices of largemouth bass (Micropterus salmoides)

项目 Items	组别Groups
项目 Items	CK	MG	DG
抗氧化指标Antioxidant indices
总抗氧化能力T-AOC/(mmol/L)	0.44±0.04^b	0.47±0.05^b	0.60±0.09^a
还原型谷胱甘肽GSH/(μmol/L)	84.62±2.07^c	382.80±2.46^a	218.06±7.99^b
丙二醛MDA/(nmol/mL)	15.81±2.11^a	12.08±0.47^b	13.40±2.20^ab
过氧化氢酶CAT/(U/mL)	21.03±0.93^b	36.36±1.85^a	37.53±1.25^a
超氧化物歧化酶SOD/(U/mL)	109.45±0.48^b	119.67±1.82^a	120.70±1.11^a
生化指标Biochemical indices
碱性磷酸酶AKP/(金氏单位/dL)	10.05±0.65^b	11.44±0.56^a	6.08±0.60^c
酸性磷酸酶ACP/(金氏单位/dL)	14.11±0.60^b	17.00±1.66^a	11.10±0.45^c
溶菌酶LZM/(μg/mL)	13.88±0.52^b	15.37±0.47^b	16.67±0.27^a
谷草转氨酶AST/(U/L)	13.50±2.08^a	9.35±1.08^b	12.10±1.07^ab
谷丙转氨酶ALT/(U/L)	3.11±0.43^a	2.31±0.14^b	2.55±0.26^ab

2.4 沼泽红假单胞菌对大口黑鲈肝脏健康的影响

2.4.1 沼泽红假单胞菌对大口黑鲈肝脏抗氧化指标的影响

由表9可知,与CK组相比,MG组和DG组大口黑鲈肝脏T-AOC显著提高(P<0.05),MG组肝脏GSH含量显著提高(P<0.05),MG组和DG组肝脏CAT活性和MDA含量显著降低(P<0.05)。

表9 沼泽红假单胞菌对大口黑鲈肝脏抗氧化指标的影响

Table 9 Effects of Rhodopseudomonas palustris on liver antioxidant indices of largemouth bass (Micropterus salmoides)

项目 Items	组别Groups
项目 Items	CK	MG	DG
总抗氧化能力T-AOC/(nmol/g prot)	0.12±0.03^c	0.20±0.01^b	0.53±0.03^a
还原型谷胱甘肽GSH/(μmol/g prot)	28.41±1.58^b	34.15±0.88^a	29.96±0.42^b
丙二醛MDA/(nmol/mg prot)	1.28±0.16^a	0.80±0.06^b	0.98±0.07^b
过氧化氢酶CAT/(U/mg prot)	1.70±0.14^a	1.02±0.06^b	1.34±0.25^b
超氧化物歧化酶SOD/(U/mg prot)	4.47±0.11	4.54±0.15	4.60±0.05

2.4.2 沼泽红假单胞菌对大口黑鲈肝脏组织形态的影响

如图2所示,CK组大口黑鲈肝脏细胞有明显的空泡化,细胞膜明显破裂,细胞核出现移位;而MG组和DG组肝脏细胞空泡化和细胞核移位情况均有不同程度改善,肝脏细胞结构完整,轮廓清晰。

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图2 沼泽红假单胞菌对大口黑鲈肝脏组织形态的影响

A~C:横切面肝脏组织形态(40×)。A:CK组;B:MG组;C:DG组。

Fig.2 Effects of Rhodopseudomonas palustris on liver tissue morphology of largemouth bass (Micropterus salmoides)

A to C: liver tissue morphology of cross sections (40×). A: CK group; B: MG group; C: DG group.

2.5 沼泽红假单胞菌对大口黑鲈肠道健康的影响

2.5.1 沼泽红假单胞菌对大口黑鲈肠道消化酶活性的影响

由表10可知,与CK组相比,MG组大口黑鲈肠道TRY和LPS活性显著提高(P<0.05),MG组和DG组AMS活性显著降低(P<0.05)。

表10 沼泽红假单胞菌对大口黑鲈肠道消化酶活性的影响

Table 10 Effects of Rhodopseudomonas palustris on intestinal digestive enzyme activities of largemouth bass (Micropterus salmoides)

项目 Items	组别Groups
项目 Items	CK	MG	DG
α-淀粉酶AMS/(U/mg prot)	2.36±0.09^a	1.61±0.36^b	1.54±0.15^b
胰蛋白酶TRY/(U/mg prot)	2 069.80±38.85^b	2 468.78±116.14^a	2 082.05±77.72^b
脂肪酶LPS/(U/g prot)	100.67±2.21^b	141.30±19.84^a	113.69±16.17^ab

2.5.2 沼泽红假单胞菌对大口黑鲈肠道组织形态的影响

如图3所示,各组大口黑鲈肠道组织结构完整,肠绒毛排列有序、无脱落现象。进一步统计分析发现(表11),与CK组相比,MG组和DG组肠道绒毛更加纤长紧密,绒毛高度、绒毛宽度和肌层厚度均显著提高(P<0.05)。

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图3 沼泽红假单胞菌对大口黑鲈肠道组织形态的影响

A~C:横切面肠道组织形态(5×)。A:CK组;B:MG组;C:DG组。

Fig.3 Effects of Rhodopseudomonas palustris on intestinal tissue morphology of largemouth bass (Micropterus salmoides)

A to C: intestinal tissue morphology of cross sections (5×). A: CK group; B: MG group; C: DG group.

表11 沼泽红假单胞菌对大口黑鲈肠道组织形态的影响

Table 11 Effects of Rhodopseudomonas palustris on intestinal tissue morphology of largemouth bass (Micropterus salmoides) μm

项目 Items	组别Groups
项目 Items	CK	MG	DG
绒毛高度Villus height	838.19±9.66^c	944.59±8.90^a	866.92±3.81^b
绒毛宽度Villus width	154.16±3.73^c	205.48±4.65^a	162.15±4.29^b
肌层厚度Muscular thickness	143.22±3.49^c	175.16±6.83^b	213.13±4.71^a

2.5.3 沼泽红假单胞菌对大口黑鲈肠道菌群的影响

如图4-A所示,通过韦恩(Venn)图统计分析发现,大口黑鲈肠道菌群共得到709个ASVs,其中3组共有ASVs为296个。由表12可知,大口黑鲈肠道菌群α多样性分析结果表明,与CK组相比,MG组和DG组Sobs指数和ACE指数显著降低(P<0.05),DG组Simpson指数显著提高(P<0.05);各组间Chao1指数、Shannon指数和Goods覆盖度无显著差异(P>0.05)。

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图4 沼泽红假单胞菌对大口黑鲈肠道菌群的影响

A:肠道菌群扩增子序列变异韦恩图;B:肠道菌群在门水平上组成;C:肠道菌群在属水平上组成。

Fig.4 Effects of Rhodopseudomonas palustris on intestinal microbiota of largemouth bass (Micropterus salmoides)

A: Venn diagram of ASVs in intestinal microbiota; B: intestinal microbiota composition at phylum level; C: intestinal microbiota composition at genus level.

表12 大口黑鲈肠道菌群alpha多样性分析

Table 12 Alpha diversity analysis of intestinal microbiota of largemouth bass (Micropterus salmoides)

项目 Items	组别Groups
项目 Items	CK	MG	DG
Sobs指数Sobs index	602.33±26.10^a	462.00±10.82^b	479.00±42.23^b
ACE指数ACE index	615.86±40.92^a	487.35±14.47^c	543.53±19.29^b
Chao1指数Chao1 index	583.66±55.38	487.35±14.47	500.33±56.83
Shannon指数Shannon index	3.85±0.25	3.65±0.20	3.51±0.02
Simpson指数Simpson index	0.06±0.01^b	0.06±0.01^b	0.68±0.19^a
Goods覆盖度Goods coverage	0.999±0.001	0.999±0.001	0.999±0.000

如图4-B所示,在门水平上,大口黑鲈肠道菌群中的优势菌门(相对丰度由高到低)为假单胞菌门(Pseudomonadota,54.22%~61.49%)、芽孢杆菌门(Bacillota,28.07%~39.99%)、放线菌门(Actinomycetota,1.24%~3.08%)、绿弯菌门(Chloroflexota,1.08%~1.95%)和拟杆菌门(Bacteroidota,0.79%~1.45%)。与CK组相比,MG组假单胞菌门相对丰度提高了6.87个百分点,放线菌门和拟杆菌门相对丰度分别提高了0.43和0.23个百分点;DG组芽孢杆菌门相对丰度提高了8.69个百分点,假单胞菌门、放线菌门和拟杆菌门相对丰度分别降低了0.40、1.41和0.42个百分点。

如图4-C所示,在属水平上,大口黑鲈肠道菌群中的优势菌属(相对丰度由高到低)为中间支原体属(Mesomycoplasma,23.45%~37.66%)、玫瑰杆菌属(Roseateles,8.05%~9.43%)、缓食菌属(Tardiphaga,6.49%~8.20%)、鞘氨醇单胞菌属(Sphingomonas,6.27%~8.51%)、不动杆菌属(Acinetobacter,4.82%~6.53%)和假单胞菌属(Pseudomonas,5.44%~6.00%)。与CK组相比,MG组中间支原体属相对丰度降低了3.55个百分点,玫瑰杆菌属、假单胞菌属和不动杆菌属相对丰度分别提高了1.38、0.51和0.43个百分点;DG组中间支原体属、鞘氨醇单胞菌属和假单胞菌属相对丰度分别提高了10.66、1.63和0.56个百分点,不动杆菌属相对丰度降低了1.27个百分点。

3 讨论

3.1 沼泽红假单胞菌对大口黑鲈生长性能的影响

鱼类的生长性能是衡量其生长和营养状况的基础依据,也是评定其商品经济价值的关键依据^[23]。研究表明,饲料添加沼泽红假单胞菌可以提高草鱼(Ctenopharyngodon idella)和罗非鱼终末体重和WGR等,促进其生长^[17,20]。也有研究表明,饲料添加光合细菌(荚膜红假单胞菌)对尖吻鲈(Lates calcarifer)生长性能无显著影响^[24]。最近有研究表明,用酶解沼泽红假单胞菌粉替代5%鱼粉对大口黑鲈终末体重和WGR等生长性能指标无显著影响^[25]。本试验结果发现,与CK组相比,MG组和DG组大口黑鲈终末体重、SGR、FR、FI、SR、HSI和VSI等生长性能指标均无显著差异;与CK组相比,DG组WGR和SGR有所降低,FCR有所升高。由于肠道消化酶活性结果显示,DG组AMS活性显著低于CK组。因此推测DG组大口黑鲈消化和吸收碳水化合物的能力降低,能量物质摄入减少,可能会利用一些脂肪来提供能量,导致大口黑鲈在摄入同等饲料量和营养物质时,表现出WGR、SGR降低和FCR升高。CF常用于反映鱼体营养状况及肥瘦程度,其数值下降通常认为是能量储备的消耗,如储存的肝糖原或体脂肪被消耗^[26]。本试验结果发现,与CK组相比,MG组和DG组CF均显著降低,这说明饲粮添加沼泽红假单胞菌能够促进大口黑鲈肝糖原和体脂肪的代谢,但具体的作用机制有待进一步研究。

3.2 沼泽红假单胞菌对大口黑鲈肌肉品质的影响

肌肉的营养成分、脂肪酸、氨基酸组成及含量以及质构特性等是评价鱼肉品质的重要指标。脂肪酸分为饱和脂肪酸和不饱和脂肪酸,一般认为,饱和脂肪酸具有诱发多种疾病的潜在危害^[27],而不饱和脂肪酸对人体的身体健康及生理功能有着极为重要的作用^[28]。多不饱和脂肪酸具有抑制癌症细胞生长、调节血压和血脂、预防心脑血管疾病以及促进免疫调节和生长发育等多种功能^[29],其中EPA和DHA是2种重要的不饱和脂肪酸,具有促进人体脑细胞发育、改善血液循环等作用^[30]。本研究中,与CK组相比,MG组和DG组大口黑鲈肌肉多不饱和脂肪酸含量显著提高,特别是EPA和DHA含量均显著提高。研究表明,饲料添加益生菌可以提高头孔无须魮(Poropuntius kontumensis)肌肉中不饱和脂肪酸含量^[31]。研究发现,一些鱼类肠道微生物已被认为可能是多不饱和脂肪酸的生产者^[32]。因此,推测沼泽红假单胞菌可能通过影响脂质代谢相关酶活性或者生成多不饱和脂肪酸,从而影响大口黑鲈肌肉中脂肪酸的组成,但其具体机制还有待进一步研究。氨基酸组成及含量反映水产品中蛋白质营养价值的高低。研究显示,饲料添加益生菌能显著提高淇河鲫鱼(Carassius auratus)和云斑尖塘鳢(Oxyeleotris marmoratus)肌肉中风味氨基酸和必需氨基酸含量^[33-34]。但在本研究中,与CK组相比,MG组大口黑鲈肌肉总氨基酸、必需氨基酸、半必需氨基酸、非必需氨基酸和鲜味氨基酸含量均无显著差异,DG组肌肉总氨基酸、必需氨基酸、半必需氨基酸、非必需氨基酸和鲜味氨基酸含量均显著降低。同时,本试验中2个试验组肌肉粗蛋白质含量与CK组相比均无显著差异,而肌肉氨基酸含量主要检测的是水解氨基酸含量,蛋白质在酸解过程中会导致部分氨基酸(如色氨酸和半胱氨酸等)被破坏,其含量不能被检出。推测DG组中这些氨基酸的含量可能较高,因此导致总氨基酸和其他氨基酸相对含量都显著降低。

鱼肉的口感品质常用质构特性来衡量。质构特性与肌肉中的水分和脂肪含量以及肌纤维直径和肌纤维密度有关。一般认为,鱼肉硬度大、弹性强时则口感更好^[35-36],而硬度与肌肉脂肪含量有关,脂肪含量越高时肉质越细嫩^[37-38]。肌肉中脂肪含量的降低会增加肌束之间的相互摩擦,降低肌肉嫩度,提高肌肉硬度^[39-40]。肌纤维直径与鱼肉的硬度、弹性和咀嚼性呈负相关。一般肌纤维直径减小、密度升高时,肌肉组织的硬度、弹性和咀嚼性会增大^[41]。本试验结果也验证了这一观点。与CK组相比,MG组和DG组肌纤维直径显著降低,肌纤维密度显著提高,同时肌肉硬度、弹性、内聚力和回弹性均显著提高。此外,与CK组相比,DG组肌肉粗脂肪含量显著降低,这可能是由于DG组试验鱼消化吸收的碳水化合物减少,机体利用粗脂肪来提供一部分能量,用于维持机体能量运转和提高机体的抗氧化能力和免疫功能等^[42]。这与国内外其他学者报道的结果保持一致。黄灵等^[43]发现,饲料添加沼泽红假单胞菌显著提高棕点石斑鱼(Epinephelus fuscoguttatus)肌肉粗蛋白质含量,降低肌肉粗脂肪含量。周秀珍等^[44]发现,饲料添加混合益生菌能够提高大口黑鲈肌肉硬度、胶黏性和咀嚼性。类似的研究也表明,饲料添加干酪乳杆菌K17显著提高大口黑鲈肌肉硬度、咀嚼性和肌纤维密度^[45]。总体来看,饲料添加沼泽红假单胞菌可以提升大口黑鲈肌肉品质和口感。

3.3 沼泽红假单胞菌对大口黑鲈血清生化和抗氧化指标的影响

动物机体内的抗氧化系统包括酶促系统(T-AOC、SOD、CAT等)和非酶促系统(GSH等)^[46]。T-AOC是反映血清中总体抗氧化能力的重要指标^[47]。SOD是生物体内至关重要的抗氧化酶之一,可以清除机体内超氧自由基。CAT可以将过氧化氢分解为完全无害的水和氧气,从而保护机体细胞免受氧自由基的损伤^[48]。MDA含量主要反映机体的脂质过氧化程度^[49]。GSH作为最重要的非酶促抗氧化物,具有清除自由基和过氧化氢等功能。研究发现,饲料添加沼泽红假单胞菌可以显著提高罗非鱼血清SOD和CAT活性^[18]。沼泽红假单胞菌也能够显著降低鲫鱼血浆MDA含量^[50]。此外,饲料添加适量球形红假单胞菌可以显著提高草鱼血清SOD活性^[51]。同样,在本试验中,与CK组相比,MG组和DG组大口黑鲈血清SOD、CAT活性及GSH含量均显著提高,MG组血清MDA含量显著降低,DG组血清T-AOC显著提高,表明饲料添加沼泽红假单胞菌活菌和死菌均可以显著增强大口黑鲈抗氧化能力。

非特异性免疫是鱼类主要的免疫形式,而AKP、ACP和LZM是反映鱼类非特异免疫力的重要指标^[52]。ACP和AKP能够水解细菌表面的磷酸单酯,提高吞噬细胞降解异物的速度,是动物溶酶体的重要组成部分,能够保护机体防御病原体入侵^[53]。而AST和ALT是一类广泛存在于肝脏等器官内的酶,当肝脏出现氧化损伤时,会导致血清AST和ALT活性提高^[54-55]。研究发现,饲料添加沼泽红假单胞菌能够提高鲫鱼血浆AKP和LZM活性^[50]。饲料添加适量球形红假单胞菌显著提高草鱼血清AKP和ACP活性^[51]。此外,饲料添加荚膜红假单胞菌、沼泽红假单胞菌和球形红假单胞菌的混合菌能够降低卵形鲳鲹(Trachinotus ovatus)血清AST和ALT活性^[56]。同样,本试验发现,饲料添加沼泽红假单胞菌活菌(MG组)能够显著提高大口黑鲈血清AKP和ACP活性,显著降低血清AST和ALT活性,提高大口黑鲈的免疫功能,同时缓解大口黑鲈的肝脏损伤。分析原因可能是由于沼泽红假单胞菌中含有大量的氨基酸、B族维生素、辅酶Q和金属离子等,是多种酶和辅酶的基本成分,因此能够激活水产动物体内AKP、ACP和LZM等免疫相关酶活性,提高抗病性^[57-59]。

3.4 沼泽红假单胞菌对大口黑鲈肝脏健康的影响

肝脏是鱼类机体中最大的消化腺、解毒及代谢器官,肝脏功能可从不同指标上反映试验动物受到外源物质的损伤程度,亦可佐证肝脏组织病理学上的潜在改变^[60-61]。研究发现,饲料添加高硒光合细菌显著提高中华绒螯蟹(Eriocheir sinensis)肝胰腺T-AOC,显著降低肝胰腺MDA含量^[62]。本研究发现,与CK组相比,2个试验组大口黑鲈肝脏T-AOC、GSH含量和SOD活性均有所提高,肝脏MDA含量显著降低,表明饲料添加沼泽红假单胞菌能够增强大口黑鲈肝脏抗氧化能力,有效调控肝脏功能相关指标,对肝脏起到保护作用。肝脏的病变被认为是动物健康受损的警示信号^[63]。研究表明,饲料添加适量沼泽红假单胞菌可以改善棕点石斑鱼肝脏组织的病变和脂肪空泡化现象^[43]。本试验发现,饲喂添加沼泽红假单胞菌的饲料后,大口黑鲈肝脏的病理变化得到了明显改善,特别是肝脏细胞空泡化和细胞核移位现象得到了明显缓解。

3.5 沼泽红假单胞菌对大口黑鲈肠道健康的影响

肠道是鱼类消化吸收的主要场所,一般通过测定肠道消化酶活性来评估水产动物的消化水平^[64]。研究发现,饲料添加沼泽红假单胞菌可以增强草鱼肠道淀粉酶和纤维素酶活性^[65]。饲料添加沼泽红假单胞菌提高了青蛤(Cyclina sinensis)幼体阶段肠道AMS和TRY活性,增强了青蛤幼体的消化能力^[66]。此外,研究发现,饲料添加红假单胞菌显著提高了红螯螯虾(Cherax quadricarinatus)肝胰腺TRY活性,而对LPS和AMS活性无显著影响^[67]。另有研究发现,沼泽红假单胞菌可以提高棕点石斑鱼肠道TRY和AMS活性,而对LPS活性无显著影响^[43]。类似的,本试验发现,与CK组相比,MG组大口黑鲈肠道TRY和LPS活性显著提高,而MG组和DG组AMS活性显著降低。研究报道,饲料添加沼泽红假单胞菌会影响肠道内生酶的活性^[43]。沼泽红假单胞菌作为一种益生菌,可以改变肠道内碳水化合物、脂类和糖类的含量,刺激鱼体肠道,从而提高消化酶活性^[68]。

除通过消化酶提高机体消化功能外,肠道绒毛也可以增加消化物与肠壁的接触,促进食物消化和营养吸收^[69]。肠道绒毛高度和肌层厚度与肠道营养物质的吸收成正比^[70]。研究发现,饲料添加适量沼泽红假单胞菌可以提高棕点石斑鱼肠道绒毛高度^[43]。本试验发现,饲料添加沼泽红假单胞菌可以提高大口黑鲈肠道绒毛高度、绒毛宽度和肌层厚度,增加绒毛面积,促进对饲料中营养物质的吸收。分析其原因,肠道中的微生物代谢会生成酸类物质如丁酸、乳酸等,可以促进肠道细胞的增殖和肠道的蠕动^[68]。由此说明,沼泽红假单胞菌对大口黑鲈肠道健康有一定的改善作用。

肠道菌群结构在宿主营养吸收和健康方面发挥着重要作用,一旦菌群失调,对鱼类的消化和免疫系统可能都会产生影响,导致疾病发生,而益生菌在调节鱼类肠道菌群中具有重要作用^[71]。研究发现,饲料添加复合益生菌降低了黄鳝(Monopterus albus)肠道菌群alpha多样性^[72],在虹鳟(Oncorhynchus mykiss)中同样得出类似结论^[73]。本试验中,与CK组相比,试验组大口黑鲈肠道菌群Sobs指数和ACE指数显著降低,Chao1指数和Shannon指数略有下降;同时,MG组假单胞菌门相对丰度提高,DG组芽孢杆菌门相对丰度提高。这说明沼泽红假单胞菌进入鱼体后降低了大口黑鲈肠道菌群的丰富度。假单胞菌门可以分泌抗菌物质,抑制病原菌生长,优化肠道微生态平衡;而芽孢杆菌门对宿主生长、免疫力也有积极的作用,能促进肠道菌群平衡^[74]。进一步分析发现,与CK组相比,2个试验组大口黑鲈肠道玫瑰杆菌属相对丰度提高;MG组不动杆菌属相对丰度提高,中间支原体属相对丰度降低。玫瑰杆菌属在调节肠道微生物群落方面起积极作用;不动杆菌属对宿主的生长发育、抗病能力也具有一定的促进作用^[75];而支原体属是水生动物中的致病菌,能够破坏宿主免疫系统,诱导其他致病菌的产生和发展^[76]。这些结果表明,饲料添加沼泽红假单胞菌能够调节大口黑鲈肠道菌群结构,改善肠道健康状况,提高鱼体免疫力。

4 结论

饲料添加沼泽红假单胞菌对大口黑鲈生长性能无显著影响,但能够提高肌肉品质和机体抗氧化能力,改善肝脏健康,同时提高肠道消化酶活性,调节肠道菌群,并改善肠道健康。在本试验条件下,在大口黑鲈饲料中拌食沼泽红假单胞菌活菌效果更佳。

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Abstract

Cite this article

1 材料与方法

1.1 试验设计和饲料制备

表1 基础饲粮营养水平(风干基础)

1.2 饲养管理

1.3 样品采集

1.4 测定指标及方法

1.4.1 饲料营养成分

1.4.2 生长性能

1.4.3 肌肉营养成分含量

1.4.4 肌肉质构特性分析

1.4.5 肌肉、肝脏和肠道组织形态

1.4.6 血清生化和抗氧化指标

1.4.7 肝脏抗氧化指标

1.4.8 肠道消化酶活性

1.4.9 肠道菌群测序分析

1.5 数据统计分析

2 结果与分析

2.1 沼泽红假单胞菌对大口黑鲈生长性能的影响

表2 沼泽红假单胞菌对大口黑鲈生长性能的影响

2.2 沼泽红假单胞菌对大口黑鲈肌肉品质的影响

2.2.1 沼泽红假单胞菌对大口黑鲈肌肉营养成分含量的影响

表3 沼泽红假单胞菌对大口黑鲈肌肉常规营养成分含量的影响(鲜样基础)

表4 沼泽红假单胞菌对大口黑鲈肌肉脂肪酸组成及含量的影响

表5 沼泽红假单胞菌对大口黑鲈肌肉氨基酸组成及含量的影响

2.2.2 沼泽红假单胞菌对大口黑鲈肌肉质构特性的影响

表6 沼泽红假单胞菌对大口黑鲈肌肉质构特性的影响

2.2.3 沼泽红假单胞菌对大口黑鲈肌肉纤维形态的影响

图1 沼泽红假单胞菌对大口黑鲈肌肉纤维形态的影响

表7 沼泽红假单胞菌对大口黑鲈肌肉纤维形态的影响

2.3 沼泽红假单胞菌对大口黑鲈血清生化和抗氧化指标的影响

表8 沼泽红假单胞菌对大口黑鲈血清生化和抗氧化指标的影响

2.4 沼泽红假单胞菌对大口黑鲈肝脏健康的影响

2.4.1 沼泽红假单胞菌对大口黑鲈肝脏抗氧化指标的影响

表9 沼泽红假单胞菌对大口黑鲈肝脏抗氧化指标的影响

2.4.2 沼泽红假单胞菌对大口黑鲈肝脏组织形态的影响

图2 沼泽红假单胞菌对大口黑鲈肝脏组织形态的影响

2.5 沼泽红假单胞菌对大口黑鲈肠道健康的影响

2.5.1 沼泽红假单胞菌对大口黑鲈肠道消化酶活性的影响

表10 沼泽红假单胞菌对大口黑鲈肠道消化酶活性的影响

2.5.2 沼泽红假单胞菌对大口黑鲈肠道组织形态的影响

图3 沼泽红假单胞菌对大口黑鲈肠道组织形态的影响

表11 沼泽红假单胞菌对大口黑鲈肠道组织形态的影响

2.5.3 沼泽红假单胞菌对大口黑鲈肠道菌群的影响

图4 沼泽红假单胞菌对大口黑鲈肠道菌群的影响

表12 大口黑鲈肠道菌群alpha多样性分析

3 讨论

3.1 沼泽红假单胞菌对大口黑鲈生长性能的影响

3.2 沼泽红假单胞菌对大口黑鲈肌肉品质的影响

3.3 沼泽红假单胞菌对大口黑鲈血清生化和抗氧化指标的影响

3.4 沼泽红假单胞菌对大口黑鲈肝脏健康的影响

3.5 沼泽红假单胞菌对大口黑鲈肠道健康的影响

4 结论

References