微量元素和饲料添加剂调控蛋壳品质的研究进展

张亚男　王　晶　武书庚　张海军　齐广海

(中国农业科学院饲料研究所，农业部饲料生物技术重点开放实验室，生物饲料开发国家工程研究中心，北京100081)



微量元素和饲料添加剂调控蛋壳品质的研究进展

张亚男王晶武书庚*张海军齐广海*

(中国农业科学院饲料研究所，农业部饲料生物技术重点开放实验室，生物饲料开发国家工程研究中心，北京100081)

蛋壳破损是家禽养殖业的重要问题，受日龄、基因、环境、营养和蛋鸡的健康状况等影响。近年来，营养调控蛋壳品质的研究主要集中于微量元素和饲料添加剂。本文简述了微量元素和饲料添加剂影响蛋壳品质的研究进展，以期为生产实践中蛋壳品质的调控提供新措施。饲粮中添加一定水平和形式的锰、微生态制剂、有机酸和中药提取物等均可改善蛋壳品质。

蛋壳品质；微量元素；微生态制剂；有机酸；精油和植物提取物

蛋壳品质降低是蛋鸡养殖过程中的重要问题，影响产蛋量和经济效益，降低孵化率和胚胎的成活率[1]。蛋壳可保护胚胎免受外界环境有害因素的影响，调节空气和水分交换，为胚胎发育提供钙等[2]。改善蛋壳品质，减少鸡蛋破损，可避免致病菌侵入鸡蛋，因而蛋壳品质广受关注。

蛋壳破损严重影响蛋鸡产业的健康发展。据统计，破损蛋占总产量的6%～10%，造成家禽产业和蛋品加工业的经济损失，以及消费者对鸡蛋品质的担忧[3]。劣质鸡蛋中破损蛋占80%～90%[4]；随日龄的增加，蛋禽体内维生素D3代谢紊乱[5]，产蛋后期破损率甚至超过总产量的20%[6]。因此，蛋鸡行业亟需改善蛋壳品质的有效措施。

影响蛋壳矿化和品质的因素较多，包括日龄、基因、环境、营养及蛋鸡健康状况等。早期营养调控蛋壳品质的研究主要集中于钙、磷等矿物质和维生素D3，近来研究表明，适当来源和水平的锰等微量元素，或功能性饲料添加剂，可促进酶的活性或影响胃肠道的代谢，促进蛋壳矿化，改善蛋壳品质。张亚男等[7-9]曾对营养素和糖胺聚糖对蛋壳品质的影响进行了综述，简述了锰、锌对蛋壳品质的影响。本文基于近年来的研究，综述了微量元素和饲料添加剂对蛋壳品质的调控进展。

1　微量元素

在蛋壳和骨骼矿化过程中，与微量元素相关的酶(如碳酸酐酶、葡萄糖醛酸基转移酶)发挥重要作用。研究表明，微量元素(如锌、锰、铜)可作为某些酶的辅助因子，通过影响碳酸钙形成和蛋壳的晶体结构从而调控蛋壳的机械特性[4]。

1.1锌

关于单独锌对蛋壳品质的研究较少。在基础饲粮(锌含量32.6 mg/kg)中添加60 mg/kg的锌，可显著改善老龄鸡(69～82周龄)的蛋壳强度；但因饲粮中同时加入了60 mg/kg的锰和10 mg/kg的铜，蛋壳品质改善作用是否因为锌尚未可知[5]。饲粮添加锌/锰(30/0、65/30和100/60 mg/kg)可提高开产蛋鸡的蛋壳厚度[10]，200/160 mg/kg可显著提高蛋壳强度，但蛋壳厚度无显著差异[11]。本课题组前期研究表明，锌可显著提高蛋壳厚度、壳重比例和蛋壳指数，与硫酸锌相比，氨基酸锌效果更佳，但未见显著影响蛋壳强度[12-13]。锌是碳酸酐酶的活性因子，饲粮锌通过提高血浆和蛋壳腺内碳酸酐酶的活性，促进碳酸钙的沉积，从而提高蛋壳重量和厚度。锌还可影响蛋壳弹性和韧性，可能与锌对基质蛋白或蛋壳结晶质地的影响有关[14]。

机体内锌的代谢需要保持一定的平衡状态。锌不足时，蛋鸡生长受到抑制、骨骼矿化不全[15]，碳酸酐酶活性降低，蛋壳品质下降[13]。饲粮锌的一般推荐剂量为70～135 mg/kg[16]，再高(170或200 mg/kg)对蛋壳品质无显著改善作用[14,17]，过高(137～655 mg/kg)则影响蛋鸡生产性能，降低蛋壳重量[18]。

1.2锰

相对于锌，锰对蛋壳品质的影响研究较多，且结果显著。饲粮添加80 mg/kg锰(氧化锰)可增加蛋壳厚度[19]。换羽后蛋鸡饲粮，随锰剂量(40～200 mg/kg)增加，蛋壳厚度和蛋壳指数线性提高[20]。100 mg/kg锰通过提高蛋壳腺内糖胺聚糖和糖醛酸的合成，改善蛋壳的超微结构和蛋壳品质，可显著提高蛋壳强度、厚度和韧性[21]。但也有研究表明，饲粮锰(25、50和70 mg/kg)不影响蛋壳品质[22]。多数研究表明，饲粮锰可改善蛋壳的机械特性，但对蛋壳重量和厚度的影响结果并不一致。锰作为半乳糖-β-1,3-葡萄糖醛酸基转移酶Ⅰ(galactose-β-1,3-glucuronosyl transferase-Ⅰ，GlcAT-Ⅰ)的活性因子，可将1个葡萄糖醛酸残基从二磷酸尿苷葡糖醛酸转移到糖胺聚糖链上，进一步催化蛋白聚糖合成[23]。饲粮116 mg/kg锰可提高蛋壳腺内GlcAT-Ⅰ mRNA表达水平和蛋白表达，提高蛋壳膜中糖胺聚糖和糖醛酸的含量，改善蛋壳超微结构，提高蛋壳强度；进一步利用比较蛋白质组学分析饲粮锰对蛋壳中蛋白种类及含量的差异，发现7种表达上调的蛋白可能与蛋壳形成有关，其中2种为半乳糖基转移酶和葡萄糖醛酸基转移酶[24]。可见，锰可能是通过调控糖胺聚糖的合成代谢，调控糖胺聚糖的形成，影响蛋壳超微结构，进而调控蛋壳品质。

饲粮缺锰可致蛋壳变薄，蛋壳超微结构(尤其乳突层)发生变化，有机基质中己糖醛酸和己糖胺的含量降低[25]。饲粮锰不足时，蛋壳腺内GlcAT-Ⅰ mRNA表达水平和蛋白表达均降低，蛋壳膜中糖胺聚糖和糖醛酸的含量降低，影响蛋壳超微结构，导致蛋壳强度降低[21]。饲粮锰适宜剂量为80～120 mg/kg，而锰在饲粮中的添加一般不超过200 mg/kg，超过适宜剂量添加对蛋壳品质再无改善作用，对生产性能亦无显著影响[26]。

1.3铜

蛋壳内的基质薄膜由大量的胶原纤维黏连而成，赖氨酰氧化酶对维持蛋壳膜的完整具有显著作用，而铜参与赖氨酰氧化酶的形成[27]。研究表明，铜缺乏可改变依赖赖氨酸的纤维交联，基质薄膜变形，导致鸡蛋大小不一，蛋壳破损[28-29]。由于铜在蛋鸡体内的需要量较低，饲粮一般不会产生缺乏，且高铜对蛋鸡生产健康状态产生负面影响。本课题组前期研究表明，125 mg/kg铜(硫酸铜)致蛋壳厚度最薄，蛋壳强度最差，6～30 mg/kg铜对蛋壳强度和蛋壳厚度较为适宜[29]。在添加植酸酶的低磷饲粮中添加有机铜不影响蛋壳品质[30]，赖氨酸铜代替硫酸铜对种鸡的蛋壳品质(如厚度、壳重比例、蛋壳指数)均无改善作用[31]。

1.4有机与无机源微量元素的对比研究

近期研究表明，微量元素的添加水平和形式(无机或有机)可影响蛋壳品质(表1)，但研究结果并不一致。以有机锌和锰(氨基酸盐)代替无机锌和锰(氧化物)，可缓解日龄对蛋壳品质的影响，提高62～70周龄蛋鸡的蛋壳强度，但对壳重比例和蛋壳厚度无显著影响[32]。饲粮中添加锌、锰和铜的氨基酸盐或硫酸盐(分别为40、40和7 mg/kg)，氨基酸盐组蛋壳厚度提高了3.8%[33]。饲粮中同时添加有机和无机锌、锰和铜，显著提高了蛋壳重量和厚度[34]。饲粮锌、锰和铜的添加可提高蛋壳强度和厚度，但微量元素来源之间并无差异[35]。基础饲粮(46.4 mg/kg锰)添加锰(120 mg/kg)可显著改善蛋壳品质，但来源之间无显著差异[36]。本课题组的研究表明，饲粮添加不同水平的有机和无机锰(25、50、100和200 mg/kg)，相对于无机锰，有机锰的相对生物学效价分别是159.1%(蛋壳强度)、107.8%(蛋壳弹性)和189.2%(蛋壳厚度)[26]。此外，关于其他元素对蛋壳品质的影响研究也逐渐增多。产蛋后期饲粮中添加丙酸铬(600 μg/kg铬)可提高蛋壳厚度[37]。亚硒酸钠和酵母硒可提高蛋壳重量和蛋壳指数，且酵母硒可提高蛋壳强度[38]，有机硒(硒代蛋氨酸)代替亚硒酸钠对蛋壳品质(如厚度、壳重比例和蛋壳指数)无显著改善作用[39]。硼可显著提高后期蛋鸡的蛋壳强度和厚度，且来源之间无显著差异[40]。镁(2.3、3.0、3.5和4.2 g/kg)可提高后期蛋鸡蛋壳强度和蛋壳厚度[41]。迄今，围绕无机和有机微量元素调控蛋壳品质进行了大量研究，但结果不一，关键在于缺乏机理的探究。

综上，微量元素锌、锰、铜在蛋壳形成过程发挥重要作用，而关于其是否存在互作或拮抗关系而影响蛋壳品质调控的研究甚少。对比研究发现微量元素间协同或拮抗共同作用对蛋壳品质的影响，可为更好地利用微量元素解决蛋壳品质问题提供理论支持，同时减少浪费。

表1　饲粮微量元素对蛋鸡蛋壳品质的影响

续表1微量元素Microelement添加形式Supplementationform添加量Supplementationlevel效果Effects参考文献References锌、锰、铜Zn，Mn，Cu无机和有机30～120mg/kg锌，35～125mg/kg锰，5～20mg/kg铜饲粮添加水平逐渐增加的锌、锰和铜，提高了蛋壳厚度和强度，来源之间无显著差异Stefanello等[35]锰Mn硫酸锰、蛋白锰和甘氨酸锰基础饲粮(46．4mg/kg锰)添加120mg/kg锰显著改善蛋壳品质，但来源之间无显著差异Venglovská等[36]锰Mn无机和有机25、50、100和200mg/kg相对于无机锰，有机锰在蛋壳强度、蛋壳弹性和蛋壳厚度上的相对生物学效价分别是159．1%、107．8%和189．2%Xiao等[26]

2　微生态制剂

研究表明，微生态制剂、有机酸等添加剂可提高钙等矿物元素的利用率[42]，改善蛋壳品质(表2)。饲粮中添加1%果聚糖可显著提高蛋壳强度和壳重比例，提高骨骼粗灰分、钙和磷水平[43]。产蛋后期蛋鸡饲粮中添加菊粉或果寡糖可提高壳重比例、蛋壳密度和强度[44]。在含有较高水平玉米干酒糟及其可溶物(DDGS)的蛋鸡(50周龄)饲粮中添加菊粉，可显著提高壳重比例、蛋壳厚度和蛋壳密度[45]。含有乳酸菌的饲粮中添加益生元(3%或4%)可显著改善蛋壳品质，主要系因益生菌和乳糖促进了肠道内短链脂肪酸的生成[46]。但也有研究表明，饲粮中添加菊粉对蛋壳重量、厚度和强度均无显著影响[47]。饲粮中添加菊粉对蛋壳粗灰分、钙含量和壳重比例无显著影响[48]。含有酵母培养物的饲粮中添加益生元对蛋壳品质亦无改善[49]。

饲粮添加益生菌可显著改善蛋壳品质[50-52]。老龄鸡(64周龄)饲粮中添加枯草芽孢杆菌(2.3×108CFU/g)可显著提高产蛋率、蛋重和厚度，降低破蛋率[53]。枯草芽孢杆菌(0.10%)和菊粉(0.10%)分别或共同添加于饲粮中，可通过增加微生物区系在绒毛吸收部位的定植，改善生产性能、蛋壳品质和钙的储存[54]。益生元和益生菌对蛋壳品质的改善作用与体内矿物元素的吸收利用有关，可提高矿物元素的溶解度。这主要是通过增加底物的合成促进短链脂肪酸的形成；肠黏膜的改变和肠细胞的定植可提高小肠吸收面积；益生素降解植酸酶，提高钙结合蛋白的表达；从而促进肠道的健康生长[55]。但也有研究表明，饲粮中添加益生菌对蛋壳品质无显著改善作用[45,56]。

3　有机酸

产蛋后期蛋鸡饲粮中添加短链脂肪酸(SCFA，0.05%)可提高蛋壳强度，降低软、破蛋率[57]，SCFA(0.078%)可提高后期蛋鸡(70周龄)的蛋壳厚度，降低破蛋率，但对蛋壳重量无显著影响，系因有机酸促进了钙的吸收，增加了血钙，从而提高了蛋壳品质[58]。蛋鸡(46～70周龄)饲粮中添加中链脂肪酸(MCFA)可提高蛋壳的壳重比例、密度和强度，是由于MCFA提高了小肠的绒毛高度，降低了肠道上部的pH，从而提高了钙、磷的利用率，改善蛋壳品质[43]。而甲酸和丙酸对罗曼蛋鸡的蛋壳厚度和强度无改善作用[59]。

4　精油和植物提取物

饲粮中添加精油复合物(牛至、月桂叶、鼠尾草、香桃木叶、茴香籽和柑橘果皮)使产蛋后期蛋鸡(54～74周龄)的软破蛋率降低15.5%[60]，精油复合物(鼠尾草、百里香和薄荷的提取物，0.015%或0.030%)提高了蛋壳硬度和重量[61]。植物提取物(红三叶草和大蒜素，0.10%)提高了蛋壳强度[62]，黑茴香籽的提取物(1%、2%和3%)可显著提高蛋壳厚度和强度[63]，饲粮中添加中药提取物使得蛋壳厚度增加10.0%、强度增加15.2%、破蛋率降低2.5%[64]。中草药复合物可增加产蛋后期蛋鸡的蛋壳强度(19.3%)，胫骨强度也增加，表明中草药改善蛋壳品质的机理可能与降低骨骼流失和促进矿物质吸收有关，但关于中草药的活性成分并不清楚[65]，尚需进一步研究。也有研究表明，饲粮添加精油或中草药提取物对蛋壳品质并无改善[45,66]。

多数研究表明，饲粮添加微生态制剂、有机酸和植物提取物可改善产蛋后期蛋鸡的蛋壳品质，其改善蛋壳品质的机理主要是通过调控肠道的健康代谢，促进矿物元素的吸收利用，从而促进蛋壳的矿化。尤其在产蛋后期，蛋鸡体内的代谢较青年期紊乱，容易产生吸收代谢问题。然而，上述饲料添加剂如何调控矿物元素吸收利用的机理尚不清楚，通过分子生物学技术如代谢组学等，筛选出肠道内与调控蛋壳品质相关菌群，并进行验证，可更好地解释改善蛋壳品质的机理。此外，将微量元素和微生态制剂、有机酸或提取物结合，共同调控蛋壳品质，可为更好地利用添加剂改善蛋壳品质提供依据，这将是营养调控蛋壳品质的新方向。

表2　饲料添加剂对蛋鸡蛋壳品质的影响

续表2饲料添加剂Feedadditive添加量Supplementationlevel效果Effects参考文献References枯草芽孢杆菌、菊粉及共用Bacillussubtilis，inulinandboth2．3×108CFU/g，0．10%和0．10%提高蛋壳重量(21．7%、24．6%和27．5%)、厚度(9．1%、18．2%和21．2%)和密度(4．6%、2．2%和4．0%)Abdelqader等[53]枯草芽孢杆菌Bacillussubtilis2．3×108CFU/g，0．05%或0．10%增加了后期蛋鸡的蛋壳厚度(8．4%和7．5%)，降低了软、破、脏蛋率Abdelqader等[54]黑茴香籽提取物Blackcuminextract1%、2%和3%可显著提高蛋壳厚度和强度Boka等[63]植物提取物(红三叶草和大蒜素)Plantextract(redcloverandallicin)0．10%蛋壳强度显著提高29．8%，但不影响蛋壳重量、厚度和蛋壳中钙、磷、镁含量Lokaewmanee等[62]脱脂奶粉(添加于含嗜酸乳酸杆菌的饲粮)SkimmedmilkpowderaddedtoadietcontainingLactobacillusacidophilus3．0%和4．0%提高了蛋壳厚度(4．2%和6．0%)和密度(0．1%和0．3%)Cesari等[46]

5　小　结

饲粮添加一定形式和水平的锰、微生态制剂、有机酸和中药提取物均可改善蛋壳品质，且在老龄蛋鸡的效果更佳，可应用于生产。饲粮添加微量元素等添加剂调控蛋壳品质结果的不一致，与蛋鸡日龄、生理状态、添加物的形式和组成等有关，根本原因在于其调控蛋壳品质的机理尚不清楚。

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(责任编辑田艳明)

Research Progress of Microelement and Feed Additives on Eggshell Quality Regulation

ZHANG YananWANG JingWU Shugeng*ZHANG HaijunQI Guanghai*

(Key Laboratory of Feed Biotechnology of Ministry of Agriculture, Feed Research Institute, Chinese Academy of >Agricultural Sciences, National Engineering Research Center of Biological Feed, Beijing 100081, China)

Poor eggshell quality is an important problem in poultry industry, which is affected by many factors such as age, genetic, environmental and nutritional factors, as well as the health status of hens. Many recent studies regarding the effect of nutrition on eggshell quality regulation have focused on dietary microelement and feed additives. This study was summarized the research progress of microelement and feed additives on eggshell quality regulation, in order to provide new measures for regulating eggshell quality in production practice. And the eggshell quality may be positively affected in certain conditions by optimal dietary level and form of manganese, as well as by the addition of probiotics, organic acids and herb extracts.[ChineseJournalofAnimalNutrition, 2016, 28(10)：3015-3024]

eggshell quality; microelement; probiotics; organic acids; essential oils and plant extracts

s: WU Shugeng, professor, E-mail: wushugeng@caas.cn; QI Guanghai, professor, E-mail: qiguanghai163@163.com

10.3969/j.issn.1006-267x.2016.10.001

2016-03-28

现代农业产业技术体系(CARS-41-K13)；家禽产业技术体系北京市创新团队(CARS-PSTP)；国家自然科学基金(31572426)

张亚男(1988—)，女，山东德州人，博士研究生，从事蛋品质营养调控研究。E-mail： zyn3299@126.com

武书庚，研究员，硕士生导师，E-mail: wushugeng@ caas.cn；齐广海，研究员，博士生导师，E-mail: qiguanghai163@163.com

S816.7

A

1006-267X(2016)10-3015-10