High Sugar/fat Ratio Feeds Favor Growth Performance and Muscle Fatty Acid Quality of Yellow River Carp （Cyprinus carpio）

doi:10.13304/j.nykjdb.2025.0272

Abstract

Abstract:

To determine the optimal sugar/fat ratio in the diet of Yellow River carp， on the premise of meeting its basic requirements for n-3 essential fatty acids （n-3 EFA）， a commercial feed with a conventional formula （sugar/fat ratio of 25/7） was used as the control group （T0）. Mixed soybean oil and linseed oil served as the fat source， and the sugar/fat ratios of the experimental diets were adjusted to 40/4 （T1）， 32/7 （T2） and 25/9 （T3） by modifying the addition amounts of carbohydrate and fat. The effects of different sugar/fat ratios on Yellow River carp were analyzed. The results showed that there were no significant differences in the morphological indices （hepatosomatic index and viscerosomatic index）， crude protein content in whole fish and muscle， serum cholesterol content and malondialdehyde content among different dietary groups. However， the specific growth rate and weight gain rate significantly increased with the elevation of dietary sugar/fat ratio. The serum glucose level in T2 treatment and the serum insulin content in T1 treatment were significantly higher than those in T0 treatment. The activities of hepatic glucokinase and 6-phosphofructokinase-1 in T1 treatment were significantly higher than those in T3 treatment. Additionally， the contents of serum triglyceride， high-density lipoprotein cholesterol， as well as hepatic triglyceride and non-esterified fatty acids significantly decreased as the sugar/fat ratio increased. Regarding hepatic antioxidant capacity， the total antioxidant capacity in T1~T3 treatments was significantly higher than that in T0 treatment. In terms of muscle fatty acid composition， the levels of eicosatrienoic acid， docosahexaenoic acid， n-6 polyunsaturated fatty acids and total polyunsaturated fatty acids in the muscle and liver of Yellow River carp in T1~T3 treatments were significantly higher than those in T0 treatment. The expression levels of Δ6 fad-β and elovl5-α in the liver of Yellow River carp in T0 and T1 treatments were significantly higher than those in T2 treatment， and the hepatic Δ6fad-α expression level in T1 treatment was significantly higher than that in T0 group treatment. These findings indicated that a dietary sugar/fat ratio of 40/4 （T1） was more conducive to the synthesis of highly unsaturated fatty acids in the fish liver. In conclusion， when the essential fatty acid content in the diet was sufficient， the utilization rate of high sugar/fat ratio diets could be improved， which in turn enhanced hepatic glucose and lipid metabolism， promoted the growth of Yellow River carp， and improved the quality of muscle fatty acids.

Key words: Yellow River carp, sugar/fat ratio, growth performance, glycolipid metabolism, highly unsaturated fatty acid （HUFA） deposition

摘要：

为明确黄河鲤饲料中适宜糖/脂添加比例，在满足其对n-3必需脂肪酸（n-3 essential fatty acids， n-3 EFA）基础需求的前提下，采用商品饲料常规配方糖脂比25/7为对照（T0），以豆油与亚麻籽油为混合脂肪源，通过调整碳水化合物与脂肪的添加量，使饲料的糖脂比分别达到40/4（T1）、32/7（T2）和 25/9（T3），分析不同糖脂比对黄河鲤的影响。结果表明，各饲料组黄河鲤的形态学指标（肝体比、脏体比）、全鱼及肌肉粗蛋白、血清胆固醇和丙二醛含量无显著差异，特定生长率、增重率随饲料糖脂比升高而显著升高，T2处理的血清葡萄糖水平和T1处理的胰岛素含量显著高于T0处理，T1处理的肝脏葡萄糖激酶、6-磷酸果糖激酶-1活性显著高于T3处理。黄河鲤血清甘油三酯、高密度脂蛋白胆固醇及肝脏甘油三酯、游离脂肪酸含量随糖脂比的升高而显著降低，在肝脏抗氧化能力方面，T1~T3处理的总抗氧化能力显著高于T0处理。在肌肉脂肪酸组成方面，T1~T3处理黄河鲤肌肉和肝脏二十碳三烯酸、二十二碳六烯酸、n-6 多不饱和脂肪酸、多不饱和脂肪酸水平显著高于T0处理，T0和T1处理鱼体肝脏Δ6 fad-β与 elovl5-α表达水平均显著高于T2处理，且T1处理鱼体肝脏Δ6 fad-α显著高于T0处理。以上表明，饲料糖脂比为 40/4（T1）时更适合鱼体肝脏合成高不饱和脂肪酸。综上可知，在饲料必需脂肪酸充足条件下，提高饲料糖脂比可提升高糖脂比饲料的利用率，改善黄河鲤肝脏糖和脂代谢，促进鱼体生长和肌肉脂肪酸品质。

关键词: 黄河鲤, 糖脂比, 生长性能, 糖脂代谢, 高不饱和脂肪酸沉积

CLC Number:

S963

Xinlu LI, Yuhang SANG, Xinyi LI, Zijie HE, Chaobin QIN, Guoxing NIE, Yuanyou LI, Dizhi XIE. High Sugar/fat Ratio Feeds Favor Growth Performance and Muscle Fatty Acid Quality of Yellow River Carp （Cyprinus carpio）[J]. Journal of Agricultural Science and Technology, 2025, 27(11): 8-22.

李新禄, 桑宇航, 黎心怡, 和子杰, 秦超彬, 聂国兴, 李远友, 谢帝芝. 高糖/脂比饲料对黄河鲤生长性能和肌肉脂肪酸品质的影响[J]. 中国农业科技导报, 2025, 27(11): 8-22.

Figures/Tables 9

References 60

[1]	郭立, 井润珍, 程镇燕, 等. 降低鲤鱼饲料蛋白质的初步研究[J]. 饲料工业, 2013, 34(8): 41-45.
[2]	翟钰姗, 姜晓娜, 瞿飞虎, 等. 饲料中蛋白质对鱼类的影响及其研究进展[J]. 河北渔业, 2023(9): 40-46.
	ZHAI Y S, JIANG X N, QU F H, et al.. The effect of protein in feed on fish and its research progress [J]. Hebei Fish., 2023(9):40-46.
[3]	杨明容, 李达, 曹岩. 泥鳅幼鱼配合饲料中蛋白质、脂肪、糖类及纤维素的适宜含量研究[J]. 饲料工业, 2017(10): 15-20.
	YANG M R, LI D, CAO Y. Studies on optimum contents of protein,fat,carbohydrate and cellulose in formulated diet of Misgurnus anguillcaudatus Juvenile [J].Feed. Ind., 2017(10): 15-20.
[4]	CAO X F, DAI Y J, LIU M Y, et al.. High-fat diet induces aberrant hepatic lipid secretion in blunt snout bream by activating endoplasmic reticulum stress-associated IRE1/XBP1 pathway [J].Biochim. Biophys. Acta Mol. Cell Biol. Lipds., 2019, 1864(3): 213-223.
[5]	PAN Y X, LUO Z, ZHUO M Q, et al.. Liver X receptor (LXR) in yellow catfish Pelteobagrus fulvidraco:molecular characterization,mRNA tissue expression and transcriptional regulation by insulin in vivo and in vitro [J]. Comp. Biochem. Physiol. Part B Biochem. Mol. Biol., 2016, 191: 13-19.
[6]	王菲, 李向飞, 李贵锋, 等. 不同糖脂比对建鲤幼鱼生长、体组成、消化及糖酵解能力的影响[J]. 水产学报, 2015, 39(9): 1386-1394.
	WANG F, LI X F, LI G F, et al.. Effects of dietary carbohydrate-to-lipid ratios on growth,body composition,digestion and glycolysis of juvenile Jian carp (Cyprinus carpio Var. Jian) [J]. J. Fish. China, 2015, 39(9): 1386-1394.
[7]	孙金辉, 范泽, 程镇燕, 等. 饲料中玉米淀粉添加水平对鲤生长性能、消化酶活性及血清生化指标的影响[J]. 动物营养学报, 2016, 28(4): 1152-1159.
	SUN J H, FAN Z, CHENG Z Y, et al.. Effects of dietary corn starch supplemental level on growth performance,digestive enzyme activities and serum biochemical indices of common carp [J]. Chin. J. Anim. Nutr., 2016, 28(4): 1152-1159.
[8]	TAN Q, XIE S, ZHU X, et al.. Effect of dietary carbohydrate-to-lipid ratios on growth and feed utilization in Chinese longsnout catfish (Leiocassis longirostris Günther) [J].J. Appl. Ichthyol., 2007, 23(5): 605-610.
[9]	华颖. 黑鲷幼鱼对饲料中碳水化合物利用及其机理研究[D]. 杭州: 浙江大学, 2017.
	HUA Y. Utilization and mechanism of carbohydrates in diet of juvenile black sea bream, Acanthopagrus schlegelii [D]. Hangzhou: Zhejiang University, 2017.
[10]	WANG J T, HAN T, LI X Y, et al.. Effects of dietary protein and lipid levels with different protein-to-energy ratios on growth performance,feed utilization and body composition of juvenile red-spotted grouper,Epinephelus akaara [J]. Aquac. Nutr., 2017, 23(5): 994-1002.
[11]	HAN D, SHAN X J, ZHANG W B, et al.. A revisit to fishmeal usage and associated consequences in Chinese aquaculture [J]. Rev. Aquac., 2018, 10(2): 493-507.
[12]	赵燕静, 王良炎, 刘慧芬, 等. 河南省野生和养殖黄河鲤肌肉营养品质比较研究[J]. 河南水产, 2020(2): 26-31.
	ZHAO Y J, WANG L Y, LIU H F, et al.. The comparative study of muscle nutrient quality in muscle between wild and cultured common carp (Cyprinus carpio) in Henan province [J]. Henan Fish., 2020(2): 26-31.
[13]	MA Y C, LI M M, XIE D Z, et al.. Fishmeal can be replaced with a high proportion of terrestrial protein in the diet of the carnivorous marine teleost (Trachinotus ovatus) [J/OL]. Aquaculture, 2020, 519: 734910 [2025-03-16]. .
[14]	谢帝芝, 于若梦, 陈芳, 等. 饲料LNA/LA比对鲤幼鱼生长性能,肝脏脂肪酸组成及Δ6fad,elovl5 mRNA表达的影响[J]. 水产学报, 2017, 41(5): 757-765.
	XIE D Z, YU R M, CHEN F, et al.. Effects of dietary LNA/LA ratio on growth,hepatic fatty acid profiles and Δ6fad, elovl5 mRNA expression in common carp(Cyprinus carpio) [J]. J. Fish. China, 2017, 41(5): 757-765.
[15]	麻永财. 卵形鲳鲹配合饲料中陆生复合蛋白替代鱼粉的研究[D]. 广州: 华南农业大学, 2020.
	MA Y C. Study on replacement of fishmeal with terrestrial compound protein in the formulated diets for Trachinotus ovatus [D]. Guangzhou:South China Agricultural University, 2020.
[16]	李贵锋. 不同蛋能比和糖脂比对建鲤幼鱼生长性能、体组成、消化及代谢酶活性的影响[D]. 南京: 南京农业大学, 2012.
	LI G F. Effects of dietary protein to energy rations and carbohydrate to lipid rations on growth performance, body composition, digestive and metabolism enzymes activities of juvenile jian carp (Cyprinus carpio var. Jian) [D]. Nanjing: Nanjing Agricultural University, 2012.
[17]	李远友, 李孟孟, 汪萌, 等. 卵形鲳鲹营养需求与饲料研究进展[J]. 渔业科学进展, 2019, 40(1): 167-177.
	LI Y Y, LI M M, WANG M, et al.. Research advances in nutritional requirement and feed of Trachinotus ovatus [J]. Prog. Fish. Sci., 2019, 40(1): 167-177.
[18]	CASTRO C, CORRAZE G, FIRMINO-DIÓGENES A, et al.. Regulation of glucose and lipid metabolism by dietary carbohydrate levels and lipid sources in gilthead sea bream juveniles [J]. Br. J. Nutr., 2016, 116(1): 19-34.
[19]	曲木, 李长娥, 刘宏超, 等. 饲料不同糖脂比对鲤鱼生长、体成分及消化酶活性的影响[J]. 动物营养学报, 2016, 28(7): 2069-2078.
	QU M, LI C E, LIU H C, et al.. Effects of different dietary carbohydrate-to-lipid ratios on growth,body composition and digestive enzyme activities of common carp (Cyprinus carpoio) [J]. Chin. J. Anim. Nutr., 2016, 28(7): 2069-2078.
[20]	何吉祥, 崔凯, 徐晓英, 等. 异育银鲫幼鱼对蛋白质、脂肪及碳水化合物需求量的研究[J]. 安徽农业大学学报, 2014, 41(1): 30-37.
	HE J X, CUI K, XU X Y, et al.. Optimial dietary requirements of protein,lipid and carbohydrate for juvenile gibel carp(Carassius auratus gibelio) [J]. J. Anhui Agric. Univ., 2014, 41(1): 30-37.
[21]	张世亮, 艾庆辉, 徐玮, 等. 饲料中糖/脂肪比例对瓦氏黄颡鱼生长、饲料利用、血糖水平和肝脏糖酵解酶活力的影响[J]. 水生生物学报, 2012, 36(3): 466-473.
	ZHANG S L, AI Q H, XU W, et al.. Effects of dietary carbohydrate to lipid ratio on growth,feed utilization,plasma glucose and glycolytic enzyme activities of juvenile darkbarbel catfish,Pelteobagrus vachelli [J]. Acta Hydrobiol. Sin., 2012, 36(3): 466-473.
[22]	李婷婷. 杂交鲟幼鱼饲料中适宜脂肪源和糖脂比研究[D]. 上海: 上海海洋大学, 2021.
	LI T T. The dietary appropriate lipid sourcesand carbohydrate to lipid level of juvenile hybrid sturgeon, Acipenser baerii♀×A . schrenckii♂ [D]. Shanghai: Shanghai Ocean University, 2021.
[23]	杨丽萍, 郑文佳, 秦超彬, 等. 饲料糖脂比对吉富罗非鱼生长、血液指标和肝脏糖代谢关键酶活性及基因表达的影响[J]. 水产学报, 2016, 40(9): 1376-1386.
	YANG L P, ZHENG W J, QIN C B, et al.. Effects of dietary carbohydrate-to-lipid ratios on growth,blood biochemical indices,key enzymes activity and gene expression of hepatic glucose metabolism of juvenile GIFT Oreochromis niloticus [J]. J. Fish. China, 2016, 40(9): 1376-1386.
[24]	鲍清华, 夏辉, 杨煜洁, 等. 三种酶解蛋白肽分别替代部分鱼粉对凡纳滨对虾幼虾生长、体组成和抗氧化性能的影响[J]. 河北渔业, 2024(1): 1-8.
[25]	孙瑞健, 徐玮, 米海峰, 等. 饲料脂肪水平和投喂频率对大黄鱼生长、体组成及脂肪沉积的影响[J]. 水产学报, 2015, 39(3): 401-409.
	SUN R J, XU W, MI H F, et al.. Effects of dietary lipid level and feeding frequency on growth,body composition and lipid deposition in juvenile large yellowcroaker (Larimichthys crocea) [J]. J. Fish. China, 2015, 39(3): 401-409.
[26]	朱婷婷, 李琦, 朱浩拥, 等. 饲料脂肪水平对俄罗斯鲟幼鱼生长、血液生化指标及抗氧化性能的影响[J]. 海洋渔业, 2017, 39(1): 58-67.
	ZHU T T, LI Q, ZHU H Y, et al.. Effects of dietary lipid level on growth performance,blood biochemical index and antioxidant status of juvenile Acipenser gueldenstaedtii [J]. Mar. Fish., 2017, 39(1): 58-67.
[27]	NANTON D A, LALL S P, MCNIVEN M A. Effects of dietary lipid level on liver and muscle lipid deposition in juvenile haddock, Melanogrammus aeglefinus L. [J].Aquac. Res., 2001, 32(s1): 225-234.
[28]	周小秋, 李洪琴, 刘汉元, 等. ω3UFA和ω6UFA对幼建鲤生产性能和免疫功能的影响[J]. 中国畜牧杂志, 2005, 41(8): 27-29.
	ZHOU X Q, LI H Q, LIU H Y, et al.. Effect of ω3UFA and ω6UFA on growth performance and immune function of young carp [J]. Chin. J. Anim. Sci., 2005, 41(8): 27-29.
[29]	WANG B K, LIU W B, XU C, et al.. Dietary carbohydrate levels and lipid sources modulate the growth performance,fatty acid profiles and intermediary metabolism of blunt snout bream Megalobrama amblycephala in an interactive pattern [J]. Aquaculture, 2017, 481: 140-153.
[30]	JIN M, LU Y, YUAN Y, et al.. Regulation of growth,antioxidant capacity,fatty acid profiles,hematological characteristics and expression of lipid related genes by different dietary n-3 highly unsaturated fatty acids in juvenile black seabream (Acanthopagrus schlegelii) [J]. Aquaculture, 2017, 471: 55-65.
[31]	AN W Q, DONG X H, TAN B P, et al.. Dietary n-3 long chain polyunsaturated fatty acids affect the serum biochemical parameters,lipid-metabolism-related of gene expression and intestinal health of juvenile hybrid grouper (♀ Epinephelus fuscoguttatus × ♂ Epinephelus lanceolatu) [J]. Aquac. Nutr., 2021, 27(5): 1383-1395.
[32]	JUMP D B. N-3 polyunsaturated fatty acid regulation of hepatic gene transcription [J]. Curr. Opin. Lipidol., 2008, 19(3): 242-247.
[33]	JESTER J V, POTMA E, BROWN D J. PPARγ regulates mouse meibocyte differentiation and lipid synthesis [J]. Ocul. Surf., 2016, 14(4): 484-494.
[34]	ZHENG J L, LUO Z, ZHU Q L, et al.. Molecular cloning and expression pattern of 11 genes involved in lipid metabolism in yellow catfish Pelteobagrus fulvidraco [J]. Gene, 2013, 531(1): 53-63.
[35]	王念民, 杨合霖, 丰超杰, 等. 碳酸盐碱度对3月龄杂交鲟(Huso dauricus♀ × Acipenser schrenckii♂)生长与血清生化指标的影响[J]. 上海海洋大学学报, 2023, 32(1): 98-107.
	WANG N M, YANG H L, FENG C J, et al.. Effects of different carbonate alkalinity on survival,growth and biochemical indexes in serum of three-month-old hybrid sturgeon (Huso dauricus♀ × Acipenser schrenckii♂) [J].J. Shanghai Ocean. Univ., 2023, 32(1): 98-107.
[36]	陆游, 周飘萍, 袁野, 等. 不同小麦淀粉和脂肪水平对大黄鱼的生长性能、饲料利用及糖代谢关键酶活力的影响[J]. 水产学报, 2017, 41(2): 297-310.
	LU Y, ZHOU P P, YUAN Y, et al.. Effects of different wheat starch and lipid levels on growth performance,feed utilization and hepatic carbohydrate metabolism key enzymes activities in large yellow croaker (Larimichthys crocea) [J]. J. Fish. China, 2017, 41(2): 297-310.
[37]	李玉, 徐怡钟, 梁婷, 等. 绿茶粉和绿茶多酚对犬的降脂作用[J]. 动物营养学报, 2018, 30(12): 5107-5117.
	LI Y, XU Y Z, LIANG T, et al.. Lipid-lowering effect of green tea powder and green tea polyphenols in canines [J]. Chin. J. Anim. Nutr., 2018, 30(12): 5107-5117.
[38]	方揽月, 邓志刚, 王洋, 等. 稀土、胆汁酸及其复配物对大口黑鲈生长性能和肠肝结构功能的影响[J]. 中国饲料, 2023(3): 89-94.
	FANG L Y, DENG Z G, WANG Y, et al.. Effects of rare earth elements,bile acids and their complexes on growth performance and enterohepatic structure and function of largemouth bass [J]. China Feed., 2023(3): 89-94.
[39]	谢瑞涛. 饲料脂肪与蛋白质对杂交石斑鱼(褐点石斑鱼♀×清水石斑鱼♂)生长及代谢的影响[D]. 湛江:广东海洋大学, 2021.
	XIE R T. Effects of dietary lipid levels and protein levels ongrowth and physiological metabolism of the hybrid (Epinephelus fuscoguttatus♀×E . polyphekadion♂) [D]. Zhanjiang: Guangdong Ocean University, 2021.
[40]	刘襄河, 叶超霞, 沈碧端, 等. 饲料中糖/脂肪比对暗纹东方幼鱼生长、血液指标、肝代谢酶活性及PEPCK基因表达的影响[J]. 水产学报, 2014, 38(8): 1149-1158.
	LIU X H, YE C X, SHEN B D, et al.. Effect of dietary carbohydrate-to-lipid ratio on growth,blood biochemical indices,hepatic metabolic enzymes and PEPCK gene expression of juvenile obscure puffer (Takifugu obscurus) [J]. J. Fish. China, 2014, 38(8): 1149-1158.
[41]	ZHOU P P, WANG M Q, XIE F J, et al.. Effects of dietary carbohydrate to lipid ratios on growth performance,digestive enzyme and hepatic carbohydrate metabolic enzyme activities of large yellow croaker (Larmichthys crocea) [J]. Aquaculture, 2016, 452: 45-51.
[42]	WANG L N, LIU W B, LU K L, et al.. Effects of dietary carbohydrate/lipid ratios on non-specific immune responses,oxidative status and liver histology of juvenile yellow catfish Pelteobagrus fulvidraco [J]. Aquaculture, 2014, 426: 41-48.
[43]	李向, 华雪铭, 魏翔, 等. 饲料中维生素D₃含量对大口黑鲈生长和抗氧化能力的影响[J]. 上海海洋大学学报, 2021, 30(1): 94-102.
	LI X, HUA X M, WEI X, et al.. Effects of dietary vitamin D₃ on the growth and antioxidant capacity of largemouth bass(Micropterus salmoides) [J].J. Shanghai Ocean. Univ., 2021, 30(1): 94-102.
[44]	唐首杰, 刘辛宇, 吴太淳, 等. 慢性氨氮胁迫对“新吉富” 罗非鱼幼鱼生长及血清生化指标的影响[J]. 水产科学, 2019, 38(6): 741-748.
	TANG S J, LIU X Y, WU T C, et al.. Effects of chronic exposure to ammonia on growth and serum biochemical indices in juvenile new GIFT strain of Nile Tilapia oreochromis niloticus [J]. Fish. Sci., 2019, 38(6): 741-748.
[45]	龚福来, 林雪, 王红权. 不同维生素C源对吉富罗非鱼生长性能、抗氧化能力和免疫力的影响[J]. 动物营养学报, 2021, 33(4): 2378-2389.
	GONG F L, LIN X, WANG H Q. Effects of different vitamin C sources on growth performance,antioxidant capacity and immunity of genetically improved farmed Tilapia [J].Chin. J. Anim. Nutr., 2021, 33(4): 2378-2389.
[46]	曲桂娟, 孟鹤媛, 魏佳淇, 等. 复方中草药对黄金鲫幼鱼生长及抗氧化能力影响的研究[J]. 中国饲料, 2023(23): 103-106.
	QU G J, MENG H Y, WEI J Q, et al.. Effects of compound Chinese herbal medicine on growth and antioxidant capacity of juvenile golden crucian carp [J]. China Feed., 2023(23): 103-106.
[47]	RAŠKOVIĆ A, MILANOVIĆ I, PAVLOVIĆ N, et al.. Antioxidant activity of rosemary (Rosmarinus officinalis L.) essential oil and its hepatoprotective potential [J/OL]. BMC Complement. Altern. Med., 2014, 14: 225 [2025-03-16]. .
[48]	GUIMARÃES N S S, RAMOS V S, PRADO-SOUZA L F L, et al.. Rosemary (Rosmarinus officinalis L.) glycolic extract protects liver mitochondria from oxidative damage and prevents acetaminophen-induced hepatotoxicity [J/OL]. Antioxidants, 2023,12(3): 628 [2025-03-16]. .
[49]	ABU-ELALA N M, YOUNIS N A, ABUBAKR H O, et al.. Influence of dietary fermented Saccharomyces cerevisiae on growth performance,oxidative stress parameters,and immune response of cultured Oreochromis niloticus [J].Fish Physiol. Biochem., 2020, 46(2): 533-545.
[50]	翟晓虎, 杨海锋, 陈慧英, 等. 丙二醛的毒性作用及检测技术研究进展[J]. 上海农业学报, 2018, 34(1): 144-148.
	ZHAI X H, YANG H F, CHEN H Y, et al.. Research progress in the toxicity and determination technology of malondialdehyde [J]. Acta Agric. Shanghai, 2018, 34(1): 144-148.
[51]	刘丽杰. 胰岛素及其信号通路在黄斑蓝子鱼Lc-Pufa合成调控中的作用及机制研究[D]. 汕头: 汕头大学, 2022.
	LIU L J. Studies on the rple and mechanism of insulin and its signaling pathway in the regulation of LC-PUFA biosynthesis in rabbitfish Siganus canaliculatus [D]. Shantou: Shantou University, 2022.
[52]	KUDA O, JELENIK T, JILKOVA Z, et al.. N-3 fatty acids and rosiglitazone improve insulin sensitivity through additive stimulatory effects on muscle glycogen synthesis in mice fed a high-fat diet [J]. Diabetologia, 2009, 52(5): 941-951.
[53]	MATSUURA B, KANNO S, MINAMI H, et al.. Effects of antihyperlipidemic agents on hepatic insulin sensitivity in perfused Goto-Kakizaki rat liver [J]. J. Gastroenterol., 2004, 39(4): 339-345.
[54]	MOUSTAÏD N, BEYER R S, SUL H S. Identification of an insulin response element in the fatty acid synthase promoter [J]. J. Biol. Chem., 1994, 269(8): 5629-5634.
[55]	LATASA M J, MOON Y S, KIM K H, et al.. Nutritional regulation of the fatty acid synthase promoter in vivo:Sterol regulatory element binding protein functions through an upstream region containing a sterol regulatory element [J]. Proc. Natl. Acad. Sci. USA, 2000, 97(19): 10619-10624.
[56]	FORETZ M, PACOT C, DUGAIL I, et al.. ADD1/SREBP-1c is required in the activation of hepatic lipogenic gene expression by glucose [J]. Mol. Cell. Biol., 1999, 19(5): 3760-3768.
[57]	PALMER D G, RUTTER G A, TAVARÉ J M. Insulin-stimulated fatty acid synthase gene expression does not require increased sterol response element binding protein 1 transcription in primary adipocytes [J]. Biochem. Biophys. Res. Commun., 2002, 291(3): 439-443.
[58]	ZHUO M Q, LUO Z, WU K, et al.. Regulation of insulin on lipid metabolism in freshly isolated hepatocytes from yellow catfish (Pelteobagrus fulvidraco) [J]. Comp. Biochem. Physiol. Part B Biochem. Mol. Biol., 2014, 177: 21-28.
[59]	CHENG Z Y, LI J H, ZHANG B L, et al.. Verification of protein sparing by feeding carbohydrate to common carp Cyprinus carpio [J]. Chin. J. Oceanol. Limnol.,2017, 35(2): 251-257.
[60]	黄望旺. 高糖日粮中添加甜菜碱或/和Tmao对团头鲂肝脂积累、糖代谢及氨基酸代谢的影响[D]. 武汉: 华中农业大学, 2023.
	HUANG W W. Effects of adding betaine and TMAO in high-carbohydrate diets on hepatic lipic accumulation,glycometabolism,and amino acid metabolism in Megalobrama amblycephala [D]. Wuhan: Huazhong Agricultural University, 2023.

成分Ingredient	处理Treatment
成分Ingredient	T0	T1	T2	T3
豆粕 Soybean meal	27.00	20.00	20.00	20.00
国产鱼粉 Domestic fish meal	5.50	2.00	2.00	3.00
猪肉粉 Pork powder	7.00	1.00	1.00	1.00
菜籽粕 Rapeseed meal	15.00	10.00	10.00	10.00
棉籽粕 Cottonseed meal	13.00	10.00	9.00	10.00
玉米淀粉 Corn starch	6.00	20.00	11.00	9.00
麦麸 Wheat bran	5.00	15.00	17.00	7.00
混合维生素 Vitamin mixture^＃	0.07	0.07	0.07	0.07
混合矿物质 Mineral mixture^※	0.11	0.11	0.11	0.11
氯化胆碱 Choline chiorlde	0.10	0.10	0.10	0.10
磷酸二氢钙 Dicalcium phosphate	2.00	2.00	2.00	2.00
豆油 Soybean oil	2.00	1.00	2.00	3.00
亚麻油 Flaxseed oil	3.00	1.50	3.00	4.50
纤维素 Cellulose	13.54	15.87	21.27	28.77
赖氨酸 L-lysine	0.50	1.00	1.10	1.10
蛋氨酸 DL-methionine	0.18	0.35	0.35	0.35
粗蛋白 Crude protein	32.00	22.16	22.03	21.94
粗脂肪 Crude lipid	7.00	4.02	6.57	8.77
亚麻酸/亚油酸 Linoleic acid/linseed acid	1.06	1.06	1.06	1.06
能量 Energy	11.31	13.59	13.21	12.8

成分Ingredient	处理Treatment
成分Ingredient	T0	T1	T2	T3
豆粕 Soybean meal	27.00	20.00	20.00	20.00
国产鱼粉 Domestic fish meal	5.50	2.00	2.00	3.00
猪肉粉 Pork powder	7.00	1.00	1.00	1.00
菜籽粕 Rapeseed meal	15.00	10.00	10.00	10.00
棉籽粕 Cottonseed meal	13.00	10.00	9.00	10.00
玉米淀粉 Corn starch	6.00	20.00	11.00	9.00
麦麸 Wheat bran	5.00	15.00	17.00	7.00
混合维生素 Vitamin mixture^＃	0.07	0.07	0.07	0.07
混合矿物质 Mineral mixture^※	0.11	0.11	0.11	0.11
氯化胆碱 Choline chiorlde	0.10	0.10	0.10	0.10
磷酸二氢钙 Dicalcium phosphate	2.00	2.00	2.00	2.00
豆油 Soybean oil	2.00	1.00	2.00	3.00
亚麻油 Flaxseed oil	3.00	1.50	3.00	4.50
纤维素 Cellulose	13.54	15.87	21.27	28.77
赖氨酸 L-lysine	0.50	1.00	1.10	1.10
蛋氨酸 DL-methionine	0.18	0.35	0.35	0.35
粗蛋白 Crude protein	32.00	22.16	22.03	21.94
粗脂肪 Crude lipid	7.00	4.02	6.57	8.77
亚麻酸/亚油酸 Linoleic acid/linseed acid	1.06	1.06	1.06	1.06
能量 Energy	11.31	13.59	13.21	12.8

基因 Gene	引物 Primer	序列 Sequence（5’-3’）
Fads6-α	Fa S	ATCGGACACCTGAAGGGAGCG
Fads6-α	Fa A	CATGTTGAGCATGTTGACATCCG
Fads6-β	Fb S	GTACCAATGGGAGGTTCGGCAC
Fads6-β	Fb A	GAGTTGAAGGTTTGGATGAAATGCATG
Elovl5-α	Eb S	GTCCTGACCATGTTCCAGACATCTTG
Elovl5-α	Eb A	CTGTAAGCGGACGAGGTGTCGTC
Elovl5-β	Eb S	GTCCTGACCATGTTCCAGACATCTTG
Elovl5-β	Eb A	CATGAAGCTCCTCTACTGCGCTG
β-actin	BS	CGCCCCAGACATCAGGGTG
β-actin	BA	CACAGATCATGTTTGAGACCTTCAACAC

基因 Gene	引物 Primer	序列 Sequence（5’-3’）
Fads6-α	Fa S	ATCGGACACCTGAAGGGAGCG
Fads6-α	Fa A	CATGTTGAGCATGTTGACATCCG
Fads6-β	Fb S	GTACCAATGGGAGGTTCGGCAC
Fads6-β	Fb A	GAGTTGAAGGTTTGGATGAAATGCATG
Elovl5-α	Eb S	GTCCTGACCATGTTCCAGACATCTTG
Elovl5-α	Eb A	CTGTAAGCGGACGAGGTGTCGTC
Elovl5-β	Eb S	GTCCTGACCATGTTCCAGACATCTTG
Elovl5-β	Eb A	CATGAAGCTCCTCTACTGCGCTG
β-actin	BS	CGCCCCAGACATCAGGGTG
β-actin	BA	CACAGATCATGTTTGAGACCTTCAACAC

指标 Index	处理Treatment
指标 Index	T0	T1	T2	T3
初始体质量 Initial body weight/g	98.60±0.02 a	98.62±0.04 a	98.68±0.07 a	98.68±0.03 a
终末体质量 Final body weight/g	267.36±4.49 a	259.52±8.39 ab	233.80±2.41 b	233.61±6.52 b
增重率 WGR/%	171.16±4.50 a	163.15±8.41 ab	136.92±2.28 b	136.74±6.54 b
特定生长率 SGR/（%·d^-1）	1.78±0.03 a	1.73±0.06 ab	1.54±0.02 bc	1.54±0.05 c
饲料系数 FCR	1.41±0.03 c	1.66±0.08 bc	1.97±0.04 ab	1.98±0.10 a
肥满度 CF	2.18±0.05 a	2.20±0.05 a	2.31±0.04 a	2.28±0.05 a
脏体比 VSI/%	7.62±0.73 a	7.85±0.47 a	7.89±0.47 a	7.27±0.53 a
肝体比 HSI/%	1.07±0.04 a	1.22±0.07 a	1.07±0.07 a	1.08±0.06 a
胴体比 Careass ratio/%	60.61±0.66 a	61.16±0.62 a	59.42±0.86 a	60.67±0.81 a