Chinese Journal of Animal Nutrition >

2026 , Vol. 38 >Issue 7: 5091 - 5105

DOI: https://doi.org/10.12418/CJAN2026.408

RESEARCH PAPER

Effects of Dietary Metabolizable Energy and Crude Protein Levels on Laying Performance, Egg Quality, Serum Reproductive Hormone Contents and Follicle Development Indices of Langya Chickens During Early Laying Period

  • CHEN Xiaolong , 1 ,
  • ZHOU Chuanfeng 2 ,
  • WANG Fubao 3 ,
  • SHI Xueping 1 ,
  • ZHANG Bolin 1 ,
  • ZHU Yimin 1 ,
  • WANG Menghan 1 ,
  • GONG Xiangwei 4 ,
  • WANG Shubai , 1, *
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  • 1 College of Animal Science and Technology, Qingdao Agricultural University, Qingdao 266109, China
  • 2 College of Life Sciences, Qingdao Agricultural University, Qingdao 266109, China
  • 3 Qingdao Ke Chuang Xin Da Technology Co., Ltd., Qingdao 266109, China
  • 4 Shandong Qingdao Poultry Treasure Langya Chicken Breeding Co., Ltd., Qingdao 266500, China
* professor, E-mail:

Received date: 2025-12-26

  Online published: 2026-07-13

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Abstract

The aim of this experiment was to investigate the effects of dietary metabolizable energy (ME) and crude protein (CP) levels on the laying performance, egg quality, serum reproductive hormone contents, follicle numbers and expression of genes [estrogen receptor 1 (ESR1), follicle stimulating hormone receptor (FSHR) and prolactin receptor (PRLR)] related to follicular development of Langya hens during early laying period. A total of 540 healthy 168-day-old Langya chickens were selected and randomly divided into 9 groups according to the principle of equal body weight and laying rate, with 5 replicates per group and 12 chickens per replicate. A 3×3 two-factor and three-level experimental design was adopted, dietary ME levels were 10.8, 11.3 and 11.8 MJ/kg, respectively, and CP levels were 14.5%, 15.5% and 16.5%, respectively. The pre-trial period was 7 days, and the formal trial period was 35 days. The results showed as follows: 1) the average daily egg mass, laying rate, egg weight, albumen height, serum reproductive hormone contents and mRNA relative expression levels of ESR1, FSHR and PRLR of 11.3 MJ/kg ME group were significantly higher than those of 10.8 and 11.8 MJ/kg ME groups (P<0.05), and the feed to egg ratio was significantly lower than that of 10.8 and 11.8 MJ/kg ME groups (P<0.05); the small yellow follicle number and big white follicle number of 11.3 and 11.8 MJ/kg ME groups were significantly higher than those of 10.8 ME group (P<0.05). 2) The average daily egg mass and laying rate of 15.5% CP group were significantly higher than those of 14.5% CP group (P<0.05), and the feed to egg ratio was significantly lower than that of 14.5% CP group (P<0.05); the eggshell thickness and eggshell strength of 16.5% CP group were significantly higher than those of 15.5% CP group (P<0.05), and the egg weight, serum reproductive hormone contents, small yellow follicle number, big white follicle number and mRNA relative expression levels of ESR1, FSHR and PRLR of 15.5% CP group were significantly higher than those of 14.5% and 16.5% CP groups (P<0.05). 3) Dietary ME and CP levels had significant interaction effects on the average daily egg mass, laying rate, feed to egg ratio, eggshell strength, egg weight, albumen height, Haugh unit, serum reproductive hormone contents, small yellow follicle number and mRNA relative expression levels of ESR1, FSHR and PRLR (P<0.05). In conclusion, the dietary appropriate ME and CP levels can improve the laying performance and egg quality of Langya hens during early laying period, promote the secretion of reproductive hormone and follicle development, up-regulate the expression of genes related to follicular development. The appropriate dietary ME and CP levels for Langya hens during early laying period are 11.3 MJ/kg and 15.5%, respectively.

Key words： Langya hens; metabolizable energy; crude protein; laying performance; egg quality; genes related to follicular development

Cite this article

CHEN Xiaolong , ZHOU Chuanfeng , WANG Fubao , SHI Xueping , ZHANG Bolin , ZHU Yimin , WANG Menghan , GONG Xiangwei , WANG Shubai . Effects of Dietary Metabolizable Energy and Crude Protein Levels on Laying Performance, Egg Quality, Serum Reproductive Hormone Contents and Follicle Development Indices of Langya Chickens During Early Laying Period[J]. Chinese Journal of Animal Nutrition, 2026 , 38(7) : 5091 -5105 . DOI: 10.12418/CJAN2026.408

琅琊鸡是山东省地方品种鸡,属肉蛋兼用型品种,其外貌美观,抗病力、抗逆性强,肉质鲜美、蛋风味独特,深受人们喜爱[1]。2018年上海合作组织青岛峰会期间,琅琊鸡和鸡蛋为会议指定特色农产品[2],2008年被收录入《国家级畜禽遗传资源品种名录》,2009年被收录入《山东省畜禽遗传资源保护名录》[3]。琅琊鸡因肉、蛋品质极佳的优势,肉、蛋产品市场售价远高于白羽肉鸡和海兰褐等高产蛋鸡[4]。琅琊鸡开产期一般在21~24周龄[5],产蛋高峰期在27~36周龄[6],年产蛋数150~200个,产蛋量偏少,饲粮报酬偏低,导致饲养成本偏高[7]。品种的遗传特性是其产蛋性能不佳的原因之一,另一原因是目前针对该品种的标准化饲养营养参数体系尚未建立,饲料配制缺乏针对性的参考标准,养殖场多以现行高产蛋鸡饲养标准或养殖人员经验为参考依据,难以精准满足琅琊鸡的营养需求。
饲粮代谢能(ME)和粗蛋白质(CP)水平直接影响家禽的生产性能和产品品质,其水平过高或过低均会对家禽生产性能和产品品质等产生负面影响[8-9]。蛋白质作为生命活动的物质基础,是家禽维持生长发育与蛋、肉生产必需的[10-11]。研究表明,五华鸡从开产期至产蛋高峰期饲粮ME水平维持在11.59 MJ/kg时,可促进生殖器官发育,进而提升其繁殖性能[12]。付胜勇等[13]研究发现,饲粮ME水平为11.82 MJ/kg、CP水平为16.5%时,有利于25~34周龄海兰灰蛋鸡生产性能的发挥。有研究发现,20~35周龄爱拔益加(AA)肉种鸡饲粮ME水平为14.04 MJ/kg时可显著提高蛋黄颜色,ME水平为11.7 MJ/kg时可显著提高蛋重[14]。
目前,饲粮ME和CP水平对产蛋期琅琊鸡产蛋性能、蛋品质及繁殖性能等的影响未见报道。针对上述问题,本试验以产蛋前期琅琊鸡为试验动物,探究饲粮ME和CP水平对其产蛋性能、蛋品质、血清生殖激素含量、卵巢卵泡数量及卵巢卵泡发育相关基因表达的影响,旨在探究琅琊鸡产蛋前期适宜的饲粮ME和CP水平,为其饲粮的合理配制提供科学依据。

1 材料与方法

1.1 伦理声明

动物试验由青岛农业大学动物科技学院实验动物伦理委员会审查批准,伦理批准编号为DKY20240522-2。

1.2 试验设计和饲养管理

试验选择540只体重相近、健康的168日龄琅琊鸡母鸡,按照体重、产蛋率均等原则分为9个组,每组5个重复,每个重复12只鸡。采用3×3二因素三水平试验设计,饲粮ME设置3个水平,分别为10.8、11.3、11.8 MJ/kg;饲粮CP设置3个水平,分别为14.5%、15.5%、16.5%。试验分组见表1。
表1 试验分组

Table 1 Experimental grouping

项目
Item		 饲粮粗蛋白质水平Dietary CP levels/%
14.5 15.5 16.5


饲粮代谢能水平
Dietary ME levels/(MJ/kg)		 10.8 Ⅰ组 Ⅱ组 Ⅲ组
11.3 Ⅳ组 Ⅴ组 Ⅵ组
11.8 Ⅶ组 Ⅷ组 Ⅸ组
试验饲粮组成及营养水平见表2,除ME、CP水平外,各组其他营养素水平基本一致。饲粮CP、钙、总磷、氨基酸含量分别采用GB/T 6432—2018、GB/T 6436—2018、GB/T 6437—2018、GB/T 18246—2019的方法测定,ME和非植酸磷含量参照《中国饲料成分及营养价值表(2022年第33版)》进行计算。试验鸡均采用笼养,饲养于同栋鸡舍,自由采食和饮水,智能化环境控制设备调控舍内环境温度、相对湿度等环境条件。预试期7 d,正试期35 d。
表2 试验饲粮组成及营养水平(风干基础)

Table 2 Composition and nutrient levels of experimental diets (air-dry basis) %

项目
Items		 组别Groups
Ⅰ Ⅱ Ⅲ Ⅳ Ⅴ Ⅵ Ⅶ Ⅷ Ⅸ
原料Ingredients
玉米Corn 64.24 62.50 60.87 65.37 65.21 62.21 65.64 66.24 60.05
小麦麸Wheat bran 3.75 4.61 3.40 3.00 1.00 1.03 0.32 1.40 1.00
豆粕Soybean meal 17.84 19.91 23.02 16.57 17.28 20.93 14.36 18.72 21.36
玉米蛋白粉Corn protein meal 1.00 0.10 0.50 1.00 2.00 2.00 2.78 1.85 2.22
豆油Soybean oil 0.10 0.10 0.30 1.01 1.00 0.91 2.56 1.66 2.50
赖氨酸Lys 0.12 0.04 0.14 0.13 0.03 0.20 0.08
蛋氨酸Met 0.15 0.13 0.12 0.15 0.12 0.10 0.14 0.12 0.11
磷酸氢钙CaHPO4 1.34 1.23 1.35 1.20 1.35 1.35 1.37 1.20 1.37
石粉Limestone 7.15 7.25 7.13 7.25 7.13 7.13 7.12 6.00 7.08
食盐NaCl 0.31 0.31 0.31 0.31 0.31 0.31 0.31 0.31 0.31
苜蓿草粉Alfalfa meal 3.00 2.82 2.00 3.00 3.47 3.00 4.20 1.42 3.00
预混料Premix1) 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00
合计Total 100.00 100.00 100.00 100.00 100.00 100.00 100.00 100.00 100.00
营养水平Nutrient levels2)
代谢能ME/(MJ/kg) 10.80 10.80 10.80 11.30 11.30 11.30 11.80 11.80 11.80
粗蛋白质CP 14.50 15.49 16.50 14.52 15.54 16.50 14.48 15.49 16.52
钙Ca 2.99 2.97 2.96 3.00 2.98 2.99 3.01 2.96 2.96
总磷TP 0.58 0.61 0.60 0.59 0.61 0.60 0.59 0.59 0.60
非植酸磷NPP 0.35 0.34 0.35 0.35 0.35 0.35 0.35 0.36 0.35
蛋氨酸Met 0.37 0.38 0.35 0.37 0.35 0.36 0.36 0.35 0.37
赖氨酸Lys 0.77 0.78 0.76 0.76 0.75 0.79 0.77 0.78 0.77

1)预混料为每千克饲粮提供 The premix provided the following per kg of diets:VA 7 500 IU,VD3 3 250 IU,VE 10 mg,生物素 biotin 0.15 mg,叶酸 folic acid 0.5mg,甲萘醌 menadione 1 mg,硝酸硫胺 thiamine mononitrate 1.3 mg,核黄素 riboflavin 4.5mg,吡哆醇 pyridoxine 3 mg,氰钴胺 cyanocobalamin 0.01 mg,烟酰胺 nicotinamide 20 mg,泛酸 pantothenic acid 20 mg,Cu (as copper sulfate) 10 mg,Fe (as ferrous sulfate) 120 mg,Mn (as manganese sulfate) 100 mg,Zn (as zinc sulfate) 95 mg,Se (as sodium selenite) 0.3 mg。

2)代谢能和非植酸磷为计算值,其余营养水平为实测值。ME and NPP were calculated values, while the other nutrient levels were measured values.

1.3 检测指标与方法

1.3.1 产蛋性能

以重复为单位记录试验期间的喂料量和剩料量、产蛋数、总蛋重等,计算平均日产蛋量、产蛋率、平均日采食量和料蛋比,计算公式参照《家禽生产性能名词术语和度量计算方法》(NY/T 823—2020)。

1.3.2 蛋品质

于正试期第21天,每组每个重复随机取2枚鸡蛋进行蛋品质测定。蛋重、蛋白高度、哈氏单位、蛋黄颜色采用多功能蛋品质测定仪(ETP-01,北京天翔飞域科技有限公司)测定;蛋壳厚度采用数显千分尺测定;蛋壳强度采用蛋壳强度测定仪(KQ-1A,北京天翔飞域科技有限公司)测定;采用游标卡尺测量鸡蛋长径和短径,蛋形指数计算公式如下:
蛋形指数=长径/短径。

1.3.3 血清生殖激素含量

于正试期第35天,每组每个重复随机选取2只试验鸡,采集血液,分别装入5 mL离心管中,静置30 min后1 006.2×g离心10 min分离血清,装于EP管中,-20 ℃保存待测。采用酶标仪(SYNERGY H4,美国伯腾仪器有限公司)检测血清促黄体生成素(LH)、促卵泡生成素(FSH)、雌二醇(E2)、孕酮(P4)和促性腺激素释放激素(GnRH)含量。试剂盒均购自上海酶联生物科技有限公司。

1.3.4 卵巢卵泡数量

于正试期第35天,每组每个重复随机选取2只试验鸡,采血后立即屠宰解剖,打开腹腔,剪断卵巢系膜,取出卵巢,分离卵泡,记录等级卵泡的数量。将卵泡直径>8 mm以上划分为等级卵泡,卵泡直径在4~8 mm划分为小黄卵泡,卵泡直径在2~4 mm划分为大白卵泡[15]。

1.3.5 卵巢卵泡发育相关基因表达

于正试期第35天,每组每个重复随机选取2只试验鸡,解剖采集卵巢组织,液氮速冻后转至-80 ℃冰箱冷冻待测。采用荧光定量PCR法检测卵巢中卵泡发育相关基因[雌激素受体1(ESR1)、促卵泡激素受体(FSHR)和催乳素受体(PRLR)]的mRNA相对表达量。按照RNA试剂盒(货号:AG21102,艾科瑞生物工程有限公司)说明书提取卵巢中的总RNA,采用反转录试剂盒(货号:AG11728,艾科瑞生物工程有限公司)进行反转录,采用荧光定量试剂盒(货号:AG11701,艾科瑞生物工程有限公司)进行相关基因mRNA相对表达量的测定。引物由武汉擎科生物技术有限公司合成,基因引物序列见表3。以β-肌动蛋白(β-actin)为内参,按照2-ΔΔCt计算目的基因的mRNA相对表达量。
表3 基因引物序列

Table 3 Primer sequences of genes

基因
Genes		 引物序列
Primer sequences (5'—3')		 登录号
Accession number
雌激素受体1
ESR1		 F: TGCTGGAATGTGCCTGGTTAGA
R: CCATGCCCTCTACACATTTCCC		 NM_205183.2
促卵泡激素受体
FSHR		 F: CCTATCCTAGCCATTGCTGTGC
R: GCTGCAGATCGTCTGTGTTTCC		 NM_205079.2
催乳素受体
PRLR		 F: TCACTGCAAACCTGACCACCAT
R: GACACCCAAAACGATCCACACA		 NM_001397566.1
β-肌动蛋白
β-actin		 F: TTCTTGGGTATGGAGTCCTGTGGT
R: CCAGACAGCACTGTGTTGGCATA		 NM_205518.2

1.4 数据处理与统计分析

采用SPSS 26.0软件对数据进行统计分析,对数据进行方差齐性检验和正态性检验,满足检验条件后进行双因素方差分析、ME和CP的主效应以及两者间的互作效应分析,当互作效应显著时,对组间进行单因素方差分析,采用Duncan氏法进行组间的差异显著性检验。数据采用平均值和均值标准误(SEM)表示,P<0.05表示差异显著。
不同指标之间相关性分析采用SPSS 26.0软件进行Pearson相关分析,Pearson相关系数>0,两者之间为正相关;Pearson相关系数<0,两者之间为负相关;Pearson相关系数绝对值越接近1,相关性越强。P<0.05为显著相关,P<0.01为极显著相关。

2 结果

2.1 饲粮ME和CP水平对产蛋前期琅琊鸡产蛋性能的影响

由表4可知,主效应分析显示,饲粮ME和CP水平对产蛋前期琅琊鸡的平均日产蛋量、产蛋率、料蛋比有显著影响(P<0.05);11.3 MJ/kg ME组平均日产蛋量和产蛋率显著高于10.8、11.8 MJ/kg ME组(P<0.05),11.3 MJ/kg ME组料蛋比显著低于10.8、11.8 MJ/kg ME组(P<0.05);15.5%和16.5% CP组平均日产蛋量显著高于14.5%组(P<0.05),15.5% CP组产蛋率显著高于14.5% CP组(P<0.05),15.5%、16.5% CP组料蛋比显著低于14.5% CP组(P<0.05)。饲粮ME和CP水平对平均日产蛋量、产蛋率和料蛋比有显著互作效应(P<0.05)。Ⅴ组平均日产蛋量和产蛋率显著高于其他各组(P<0.05),Ⅵ组显著高于Ⅰ、Ⅱ、Ⅲ、Ⅶ组(P<0.05);Ⅴ组料蛋比最低,其次是Ⅵ组,Ⅴ组显著低于Ⅰ、Ⅱ、Ⅲ、Ⅳ、Ⅶ、Ⅷ、Ⅸ组(P<0.05),Ⅵ组显著低于Ⅰ、Ⅱ、Ⅲ、Ⅶ组(P<0.05)。
表4 饲粮ME和CP水平对产蛋前期琅琊鸡产蛋性能的影响

Table 4 Effects of dietary ME and CP levels on laying performance of Langya hens during early laying period

项目
Items		 平均日产蛋量
Average daily
egg mass/(g/d)		 产蛋率
Laying rate/%		 料蛋比
Feed/egg		 平均日采食量
Average daily
feed intake/g
组别Groups
Ⅰ 16.10d 39.83c 5.72a 91.80
Ⅱ 16.36cd 39.95c 5.52ab 90.34
Ⅲ 15.54d 39.51c 5.77a 90.69
Ⅳ 17.72b 44.16b 5.11bc 90.50
Ⅴ 19.68a 47.39a 4.54d 89.30
Ⅵ 18.14b 44.42b 4.88cd 88.31
Ⅶ 15.64d 39.60c 5.90a 88.80
Ⅷ 17.21bc 42.80b 5.21bc 89.23
Ⅸ 17.94b 43.76b 5.01c 89.92
SEM 0.356 0.886 0.145 2.138
主效应Main effects


代谢能
ME/(MJ/kg)		 10.8 16.00c 39.76c 5.67a 90.95
11.3 18.51a 45.32a 4.84c 89.37
11.8 16.93b 42.05b 5.37b 89.32


粗蛋白质
CP/%		 14.5 16.49b 41.19b 5.58a 90.37
15.5 17.75a 43.38a 5.09b 89.63
16.5 17.21a 42.56ab 5.22b 89.64
P值P-value
代谢能ME <0.001 <0.001 <0.001 0.575
粗蛋白质CP 0.001 0.016 0.001 0.889
代谢能×粗蛋白质ME×CP 0.001 0.041 0.032 0.952

同列数据肩标不同小写字母表示差异显著(P<0.05),相同或无字母表示差异不显著(P>0.05)。表5至表8同。

In the same column, values with different small letter superscripts mean significant difference (P<0.05), while with the same or no letter superscripts mean no significant difference (P>0.05). The same as Table 5 to Table 8.

2.2 饲粮ME和CP水平对产蛋前期琅琊鸡蛋品质的影响

由表5可知,主效应分析显示,饲粮ME水平对蛋重、蛋白高度有显著影响(P<0.05),11.3 MJ/kg ME组蛋重和蛋白高度显著高于10.8、11.8 MJ/kg ME组(P<0.05);饲粮CP水平对蛋壳厚度、蛋壳强度、蛋重、蛋白高度有显著影响(P<0.05),16.5% CP组蛋壳厚度显著高于14.5%和15.5% CP组(P<0.05),16.5% CP组蛋壳强度显著高于15.5% CP组(P<0.05),15.5% CP组蛋重显著高于14.5%、16.5% CP组(P<0.05),15.5%、16.5% CP组蛋白高度显著高于14.5% CP组(P<0.05)。饲粮ME和CP水平对蛋壳强度、蛋重、蛋白高度、哈氏单位有显著互作效应(P<0.05)。Ⅵ组蛋壳强度显著高于Ⅱ、Ⅳ、Ⅷ组(P<0.05),Ⅴ组显著高于Ⅷ组(P<0.05);Ⅴ组蛋重显著高于其他各组(P<0.05),Ⅰ、Ⅲ组显著低于其他各组(P<0.05);Ⅴ、Ⅵ组蛋白高度显著高于其他各组(P<0.05),Ⅶ组显著低于其他各组(P<0.05);Ⅵ组哈氏单位显著高于Ⅳ、Ⅶ组(P<0.05)。
表5 饲粮ME和CP水平对产蛋前期琅琊鸡蛋品质的影响

Table 5 Effects of dietary ME and CP levels on egg quality of Langya hens during early laying period

项目
Items		 蛋壳厚度
Eggshell
thickness/
mm		 蛋壳强度
Eggshell
strength/
(kg/cm2)		 蛋重
Egg
weight/g		 蛋白高度
Albumen
height/mm		 哈氏单位
Haugh unit		 蛋黄颜色
Yolk color		 蛋形指数
Egg shape
index
组别Groups
Ⅰ 0.36 3.98ab 40.56c 4.81b 76.27ab 5.20 1.29
Ⅱ 0.37 3.66bc 43.18b 4.94b 73.88ab 5.00 1.32
Ⅲ 0.40 3.99ab 40.72c 4.93b 72.14ab 6.00 1.31
Ⅳ 0.36 3.59bc 42.38b 4.85b 69.95b 5.50 1.32
Ⅴ 0.37 4.16ab 44.86a 5.24a 73.67ab 5.80 1.28
Ⅵ 0.38 4.35a 43.01b 5.15a 79.31a 6.10 1.31
Ⅶ 0.36 4.00ab 42.45b 4.11c 68.47b 5.50 1.30
Ⅷ 0.34 3.19c 42.55b 4.98b 78.32a 5.30 1.30
Ⅸ 0.37 4.00ab 42.54b 4.90b 75.59ab 5.40 1.32
SEM 0.010 0.198 0.418 0.057 2.547 0.310 0.014
主效应Main effects


代谢能
ME/(MJ/kg)		 10.8 0.37 3.88 41.49c 4.89b 74.10 5.40 1.31
11.3 0.37 4.03 43.41a 5.08a 74.31 5.80 1.31
11.8 0.36 3.73 42.51b 4.66c 74.13 5.40 1.31


粗蛋白质
CP/%		 14.5 0.36b 3.86ab 41.79b 4.59b 71.56 5.40 1.31
15.5 0.36b 3.67b 43.53a 5.05a 75.29 5.37 1.30
16.5 0.38a 4.11a 42.09b 4.99a 75.68 5.83 1.31
P值P-value
代谢能ME 0.125 0.195 <0.001 <0.001 0.994 0.203 0.936
粗蛋白质CP 0.014 0.033 <0.001 <0.001 0.106 0.135 0.531
代谢能×粗蛋白质ME×CP 0.424 0.021 0.016 <0.001 0.038 0.463 0.180

2.3 饲粮ME和CP水平对产蛋前期琅琊鸡血清生殖激素含量的影响

由表6可知,主效应分析显示,饲粮ME和CP水平对血清E2、FSH、GnRH、LH、P4含量有显著影响(P<0.05);11.3 MJ/kg ME组血清E2、FSH、GnRH、LH、P4含量显著高于10.8和11.8 MJ/kg ME组(P<0.05),15.5% CP组血清E2、FSH、GnRH、LH、P4含量显著高于14.5%、16.5% CP组(P<0.05)。饲粮ME和CP水平对血清E2、FSH、GnRH、LH、P4含量有显著互作效应(P<0.05)。Ⅴ组血清E2含量显著高于其他各组(P<0.05),Ⅳ组显著高于Ⅰ、Ⅲ、Ⅶ、Ⅸ组(P<0.05);Ⅴ组血清FSH含量显著高于其他各组(P<0.05),Ⅵ组显著高于Ⅰ、Ⅱ、Ⅲ、Ⅳ、Ⅶ、Ⅷ、Ⅸ组(P<0.05);Ⅴ组血清GnRH含量显著高于Ⅰ、Ⅱ、Ⅲ、Ⅳ、Ⅶ、Ⅷ、Ⅸ组(P<0.05),Ⅵ组显著高于Ⅲ、Ⅳ、Ⅶ、Ⅸ组(P<0.05);Ⅴ组血清LH含量显著高于其他各组(P<0.05),Ⅱ组显著高于Ⅰ、Ⅲ、Ⅶ、Ⅸ组(P<0.05);Ⅴ组血清P4含量显著高于其他各组(P<0.05),Ⅵ组显著高于Ⅰ、Ⅲ、Ⅶ、Ⅷ、Ⅸ组(P<0.05)。
表6 饲粮ME和CP水平对产蛋前期琅琊鸡血清生殖激素含量的影响

Table 6 Effects of dietary ME and CP levels on contents of reproductive hormone in serum of Langya hens during early laying period

项目
Items		 雌二醇
E2/
(pg/mL)		 促卵泡素
FSH/
(mIU/mL)		 促性腺激素
释放激素
GnRH/
(mIU/mL)		 黄体生成素
LH/
(ng/mL)		 孕酮
P4/
(ng/mL)
组别Groups
Ⅰ 518.68f 8.58e 112.82bcd 147.19c 20.56de
Ⅱ 625.33bcde 9.56c 113.48bc 158.42b 21.46bc
Ⅲ 606.70de 8.36e 108.12d 146.34c 20.08e
Ⅳ 650.30b 9.53c 111.82cd 150.58bc 21.58bc
Ⅴ 696.09a 10.40a 120.18a 178.34a 22.88a
Ⅵ 641.91bc 10.08b 117.18ab 154.78bc 21.94b
Ⅶ 610.68cde 8.46e 108.03d 147.33c 20.49de
Ⅷ 633.09bcd 9.33cd 112.78bcd 149.72bc 21.26c
Ⅸ 594.95e 9.02d 109.24cd 146.91c 20.99cd
SEM 10.954 0.110 1.586 3.113 0.190
主效应Main effects


代谢能
ME/(MJ/kg)		 10.8 583.57c 8.83b 111.48b 150.65b 20.70b
11.3 662.77a 10.00a 116.40a 161.23a 22.13a
11.8 612.91b 8.94b 110.02b 147.99b 20.91b


粗蛋白质
CP/%		 14.5 593.22c 8.86c 110.89b 148.37b 20.87b
15.5 651.50a 9.77a 115.48a 162.16a 21.87a
16.5 614.52b 9.15b 111.52b 149.34b 21.00b
P值P-value
代谢能ME <0.001 <0.001 <0.001 <0.001 <0.001
粗蛋白质CP <0.001 <0.001 0.002 <0.001 <0.001
代谢能×粗蛋白质ME×CP <0.001 0.001 0.041 0.003 0.027

2.4 饲粮ME和CP水平对产蛋前期琅琊鸡卵巢卵泡数量的影响

由表7可知,主效应分析显示,饲粮ME和CP水平对小黄卵泡数和大白卵泡数有显著影响(P<0.05),11.3和11.8 MJ/kg ME组小黄卵泡数和大白卵泡数显著高于10.8 MJ/kg ME组(P<0.05),15.5% CP组小黄卵泡数和大白卵泡数显著高于14.5%和16.5% CP组(P<0.05)。饲粮ME和CP水平对小黄卵泡数有显著互作效应(P<0.05)。Ⅴ组小黄卵泡数显著高于Ⅰ、Ⅱ、Ⅲ、Ⅵ组(P<0.05),Ⅳ、Ⅷ组显著高于Ⅰ、Ⅲ组(P<0.05)。
表7 饲粮ME和CP水平对产蛋前期琅琊鸡卵巢卵泡数量的影响

Table 7 Effects of dietary ME and CP levels on ovarian follicle numbers of Langya hens during early laying period 个

项目
Items		 等级卵泡数
Graded follicle
number		 小黄卵泡数
Small yellow follicle
number		 大白卵泡数
Big white follicle
number
组别Groups
Ⅰ 5.2 5.2d 9.4
Ⅱ 5.2 7.0bc 10.6
Ⅲ 5.4 6.6c 9.4
Ⅳ 5.6 7.6ab 11.8
Ⅴ 6.0 8.0a 12.8
Ⅵ 5.6 7.0bc 11.6
Ⅶ 5.4 7.4abc 9.8
Ⅷ 5.6 7.6ab 12.6
Ⅸ 5.4 7.2abc 11.6
SEM 0.32 0.30 0.49
主效应Main effects


代谢能
ME/(MJ/kg)		 10.8 5.27 6.27b 9.80b
11.3 5.73 7.53a 12.07a
11.8 5.47 7.40a 11.33a


粗蛋白质
CP/%		 14.5 5.40 6.73b 10.33b
15.5 5.60 7.53a 12.00a
16.5 5.47 6.93b 10.87b
P值P-value
代谢能ME 0.221 <0.001 <0.001
粗蛋白质CP 0.744 0.007 0.001
代谢能×粗蛋白质ME×CP 0.917 0.013 0.219

2.5 饲粮ME和CP水平对产蛋前期琅琊鸡卵巢卵泡发育相关基因表达的影响

由表8可知,主效应分析显示,饲粮ME和CP水平对ESR1、FSHR、PRLR的mRNA相对表达量有显著影响(P<0.05),11.3 MJ/kg ME组ESR1、FSHR、PRLR的mRNA相对表达量显著高于10.8和11.8 MJ/kg ME组(P<0.05),15.5% CP组ESR1、FSHR、PRLR的mRNA相对表达量显著高于14.5%和16.5% CP组(P<0.05)。饲粮ME和CP水平对ESR1、FSHR、PRLR的mRNA相对表达量有显著互作效应(P<0.05)。Ⅴ组ESR1、FSHR、PRLR的mRNA相对表达量显著高于其他各组(P<0.05),Ⅶ组ESR1的mRNA相对表达量显著高于Ⅰ、Ⅲ、Ⅷ、Ⅸ组(P<0.05),Ⅱ、Ⅳ组FSHR的mRNA相对表达量显著高于Ⅰ、Ⅲ、Ⅶ、Ⅸ组(P<0.05),Ⅱ、Ⅳ、Ⅵ、Ⅷ组PRLR的mRNA相对表达量显著高于Ⅰ、Ⅲ、Ⅶ组(P<0.05)。
表8 饲粮ME和CP水平对产蛋前期琅琊鸡卵巢卵泡发育相关基因表达的影响

Table 8 Effects of dietary ME and CP levels on expression of genes related to follicular development in ovary of Langya hens during early laying period

项目
Items		 雌激素受体1
ESR1		 促卵泡激素受体
FSHR		 催乳素受体
PRLR
组别Groups
Ⅰ 1.03e 1.00f 1.00d
Ⅱ 1.39bc 1.53b 1.22b
Ⅲ 1.00e 1.28e 1.05d
Ⅳ 1.41bc 1.52b 1.24b
Ⅴ 1.61a 1.93a 1.41a
Ⅵ 1.32bcd 1.45bc 1.23b
Ⅶ 1.43b 1.39cd 1.12c
Ⅷ 1.20d 1.45bc 1.20b
Ⅸ 1.26cd 1.36de 1.17bc
SEM 0.051 0.029 0.023
主效应Main effects


代谢能
ME/(MJ/kg)		 10.8 1.14c 1.27c 1.09c
11.3 1.45a 1.63a 1.29a
11.8 1.29b 1.40b 1.17b


粗蛋白质
CP/%		 14.5 1.29b 1.31c 1.12b
15.5 1.40a 1.64a 1.28a
16.5 1.19c 1.37b 1.15b
P值P-value
代谢能ME <0.001 <0.001 <0.001
粗蛋白质CP <0.001 <0.001 <0.001
代谢能×粗蛋白质ME×CP <0.001 <0.001 0.006

2.6 卵泡发育相关基因表达与产蛋性能、蛋品质、卵泡数量的相关性分析

由表9可知,ESR1的mRNA相对表达量与平均日产蛋量、产蛋率、蛋重、小黄卵泡数呈极显著正相关(P<0.01),与料蛋比呈显著负相关(P<0.05)。FSHR的mRNA相对表达量与平均日产蛋量、产蛋率、蛋重、小黄卵泡数、蛋白高度呈显著或极显著正相关(P<0.05或P<0.01),与料蛋比呈极显著负相关(P<0.01)。PRLR的mRNA相对表达量与平均日产蛋量、产蛋率、蛋重、蛋白高度、小黄卵泡数呈极显著正相关(P<0.01),与料蛋比呈极显著负相关(P<0.01)。
表9 卵泡发育相关基因表达与产蛋性能、蛋品质、卵泡数量的相关性分析

Table 9 Correlation analysis of expression of gene related to follicular development with laying performance, egg quality and follicle numbers

项目
Items		 平均日
产蛋量
Average daily
egg mass		 产蛋率
Laying
rate		 料蛋比
Feed/
egg		 蛋壳强度
Eggshell
strength		 蛋重
Egg
weight		 蛋白高度
Albumen
height		 哈氏单位
Haugh unit		 小黄卵泡数
Small yellow
follicle
number
雌激素受体1
ESR1		 0.43** 0.41** -0.32* 0.14 0.66** 0.04 -0.19 0.49**
促卵泡激素受体
FSHR		 0.62** 0.59** -0.55** 0.08 0.74** 0.34* -0.05 0.60**
催乳素受体
PRLR		 0.69** 0.65** -0.64** 0.06 0.78** 0.46** 0.06 0.50**

*代表显著相关(P<0.05),**代表极显著相关(P<0.01)。

* represented significant correlation (P<0.05), and ** represented extremely significant correlation (P<0.01).

3 讨论

3.1 饲粮ME和CP水平对产蛋前期琅琊鸡产蛋性能的影响

付胜勇等[13]研究发现,与饲粮ME水平为11.00 MJ/kg时相比,饲粮ME水平为11.82 MJ/kg时可显著提高25~28周龄海兰灰蛋鸡的产蛋率和料蛋比,降低平均日采食量。张利敏[16]研究报道,与饲粮ME水平为10.04 MJ/kg时相比,饲粮ME水平为11.30、11.92 MJ/kg时可显著提高26周龄罗曼褐壳蛋鸡的产蛋率,降低料蛋比和平均日采食量。据报道,饲粮CP水平由15%提升至16%,24周龄淮南麻黄鸡种鸡的产蛋率显著上升,料蛋比显著下降[17]。张妮娅等[18]研究发现,饲粮ME和CP水平对28周龄海兰褐蛋鸡平均日采食量和料蛋比有显著互作效应。本试验结果显示,饲粮ME水平为11.3 MJ/kg、CP水平为15.5%时可显著提高产蛋期前期琅琊鸡的平均日产蛋量和产蛋率,降低料蛋比;饲粮ME和CP水平对平均日产蛋量、产蛋率和料蛋比有显著互作效应。由此可见,饲粮不同ME和CP水平对蛋鸡产蛋性能有不同程度的影响,适宜的饲粮ME和CP水平可改善蛋鸡产蛋性能。
据报道,饲粮ME水平过高可促进蛋鸡脂肪沉积,腹部脂肪沉积过多会挤压卵巢,影响卵泡发育和排卵,造成产蛋率下降[19]。饲粮CP水平升高可使蛋鸡机体蛋白质沉积向鸡蛋的分配比例增加,进而提高日产蛋量[20]。本试验相关性分析显示,产蛋前期琅琊鸡卵巢ESR1、FSHR、PRLR的mRNA相对表达量与平均日产蛋量、产蛋率呈极显著正相关,与料蛋比呈显著或极显著负相关。由此可见,饲粮不同ME和CP水平通过影响卵泡发育相关基因表达,对琅琊鸡产蛋性能发挥了不同调控作用。

3.2 饲粮ME和CP水平对产蛋前期琅琊鸡蛋品质的影响

据报道,与饲粮ME水平为11.82 MJ/kg时相比,饲粮ME水平为11.00 MJ/kg时可显著提升21~24周龄海兰灰蛋鸡的蛋重和蛋白高度,而对蛋壳厚度、蛋壳强度和蛋黄颜色无显著影响[13]。陈万虹等[21]研究发现,22~25周龄雪山鸡饲喂不同CP水平饲粮,16.5% CP组较14.5%、15.5% CP组蛋壳厚度显著增加,14.5% CP组蛋形指数最高,饲粮ME和CP水平对蛋品质无显著互作效应。耿爱莲等[22]研究发现,与饲粮CP水平为15.5%和15.0%时相比。饲粮CP水平为16.0%时可显著增加30周龄北京油鸡蛋重,对其他蛋品质指标无显著影响。饲粮CP水平过高或过低均会降低鸡蛋的哈氏单位和蛋白高度[23]。本试验发现,饲粮ME水平为11.3 MJ/kg、CP水平为15.5%时可提高产蛋前期琅琊鸡的蛋重和蛋白高度,饲粮CP水平为16.5%时可提高蛋壳厚度和蛋壳强度;饲粮ME和CP水平对蛋壳强度、蛋重、蛋白高度、哈氏单位有显著互作效应;相关性分析显示,ESR1的mRNA相对表达量与蛋重呈极显著正相关,FSHR、PRLR的mRNA相对表达量与蛋重、蛋白高度呈显著或极显著正相关。由此可见,饲粮不同ME和CP水平通过影响卵泡发育相关基因表达,进而影响蛋品质。

3.3 饲粮ME和CP水平对产蛋前期琅琊鸡血清生殖激素含量的影响

据报道,饲粮ME和CP水平的变化会导致蛋白质和脂质代谢平衡发生变化,影响动物机体内分泌活动和生殖激素的分泌[24]。低CP水平饲粮可增加参与脂肪酸合成的代谢物数量,改变脂质代谢,CP水平过高会导致肝脏损伤;ME摄入不足会导致蛋鸡腹脂沉积下降,体况降低,ME摄入过多会导致蛋鸡脂肪细胞代谢异常,并增加肝脏和脂肪组织中甘油三酯的积累,进而抑制生殖激素的分泌[25-26]。杨玉等[27]研究发现,与饲粮ME水平为10.87 MJ/kg时相比,饲粮ME水平为13.34 MJ/kg时可显著提高19~28周龄海兰褐壳蛋鸡血清FSH、LH、P4含量。赵晓钰等[28]研究报道,与饲粮CP水平为11.20%时相比,饲粮CP水平为13.86%和16.48%时可显著提高伊犁鹅血清E2、LH和GnRH含量,显著降低血清催乳素(PRL)含量。本试验结果显示,饲粮ME水平为11.3 MJ/kg、CP水平为15.5%时可显著提高产蛋前期琅琊鸡血清E2、FSH、GnRH、LH、P4含量;饲粮ME和CP水平对血清E2、FSH、GnRH、LH、P4含量有显著互作效应。由此可见,饲粮不同ME和CP水平对蛋鸡血清生殖激素含量有不同程度的影响,饲粮适宜的ME和CP水平可提高血清生殖激素含量。

3.4 饲粮ME和CP水平对产蛋前期琅琊鸡卵巢卵泡数量的影响

据报道,饲粮ME水平过高可促进蛋鸡脂肪沉积,卵巢被脂肪浸润,导致小生长卵泡数量下降,闭锁卵泡数量增加[29]。Zhang等[30]研究发现,饲粮CP水平(13.5%~17.5%)对荆江鸭的小黄卵泡数、大白卵泡数无显著影响。Shi等[31]研究发现,不同CP水平(14%~16%)对皖西白鹅卵泡数量有显著影响,16% CP组大白卵泡数显著增加。孟润泽[32]研究发现,11.30和12.60 MJ/kg ME组的24周龄北京鸭成熟卵泡数显著高于10.00、10.65、11.95 MJ/kg ME组。本试验发现,11.3和11.8 MJ/kg ME组小黄卵泡数和大白卵泡数显著高于10.8 MJ/kg ME组,15.5% CP组小黄卵泡数和大白卵泡数显著高于14.5%和16.5% CP组。由此可见,饲粮不同ME和CP水平对禽类卵泡发育有不同程度影响。据报道,适宜的ME水平有利于蛋鸡卵泡发育,长期高水平ME摄入会诱发高胰岛素血症和高脂血症,导致蛋鸡卵泡发育受损[33]。

3.5 饲粮ME和CP水平对产蛋前期琅琊鸡卵泡发育相关基因表达的影响

Xin等[34]研究发现,与饲粮ME水平为11.7 MJ/kg、CP水平为16.5%组相比,饲粮ME水平为11.3 MJ/kg、CP水平为15.0%时可上调15周龄海兰褐蛋鸡卵巢ESR1的mRNA相对表达量。赵鑫等[35]研究发现,与饲粮ME水平为11.28 MJ/kg、CP水平为14.48%组相比,饲粮ME水平为10.26 MJ/kg、CP水平为11.42%组浙东白鹅母鹅卵巢FSH的mRNA相对表达量显著上调,卵巢FSHR和ESR1的mRNA相对表达量无显著差异。本试验发现,饲粮ME水平为11.3 MJ/kg、CP水平为15.5%时可显著上调产蛋前期琅琊鸡卵巢ESR1、FSHR、PRLR的mRNA相对表达量;饲粮ME和CP水平对卵巢ESR1、FSHR、PRLR的mRNA相对表达量有显著互作效应。由此可见,饲粮ME和CP水平过高或过低不利于卵巢ESR1、FSHR、PRLR的表达,饲粮适宜的ME和CP水平可上调琅琊鸡卵泡发育相关基因表达,从而提高其产蛋性能。
据报道,改变卵巢促性腺激素受体的表达,进而调节卵巢对促性腺激素刺激的敏感性,是饲粮ME水平调控卵泡发育与排卵过程的重要途径之一[36]。周虚等[37]研究发现,高ME水平会促进卵巢FSHR的表达,而低ME水平则会抑制其表达。

3.6 卵泡发育相关基因表达与产蛋性能、蛋品质、卵泡数量的相关性分析

孙鹃等[38]研究报道,绿壳蛋鸡血液ESR1的mRNA相对表达量与产蛋量呈正相关。研究发现,血液雌激素受体(ESR)的表达对文昌鸡产蛋数有显著影响[39]。研究发现,茶花鸡卵巢FSHR的mRNA相对表达量与产蛋数呈显著正相关[40-41]。邹坤等[42]研究发现,雷州黑鸭卵巢ESR1和FSHR的mRNA相对表达量与产蛋量呈显著正相关,与蛋品质无显著相关关系。本试验发现,产蛋前期琅琊鸡卵巢ESR1的mRNA相对表达量与平均日产蛋量、产蛋率、蛋重、小黄卵泡数呈极显著正相关;FSHR、PRLR的mRNA相对表达量与平均日产蛋量、产蛋率、蛋重、蛋白高度、小黄卵泡数呈显著或极显著正相关;ESR1、FSHR和PRLR的mRNA相对表达量与料蛋比呈显著或极显著负相关,此结果与上述报道基本一致。由此可见,饲粮ME和CP水平通过影响蛋鸡卵泡发育相关基因表达途径调控其卵泡发育、产蛋性能和蛋品质。

4 结论

① 饲粮适宜的ME和CP水平可上调卵巢卵泡发育相关基因表达,从而促进卵泡发育和生殖激素分泌,最终提高产蛋前期琅琊鸡产蛋性能和蛋品质。
② 饲粮ME水平为11.3 MJ/kg、CP水平为15.5%时,产蛋前期琅琊鸡产蛋率最高,料蛋比最低。
③ 饲粮CP水平为16.5%时,产蛋前期琅琊鸡可获得较好的蛋壳厚度和蛋壳强度。
④ 产蛋前期琅琊鸡饲粮ME和CP水平分别以11.3 MJ/kg和15.5%为宜。

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