动物营养学报 >

2023 , Vol. 35 >Issue 5: 2786 - 2796

DOI: https://doi.org/10.12418/CJAN2023.261

综述

乳糖合成与分泌的机制及其与奶畜泌乳性能的相关性分析

  • 李戎诺 ,
  • 夏超 ,
  • 邓露 , *
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  • 西北农林科技大学动物科技学院,杨凌 712100
*邓 露,副教授,博士生导师,E-mail:

李戎诺(2000—),女,山东济宁人,硕士,从事分子营养学与奶牛高效肝脏糖异生相关研究。E-mail:

Copy editor: 陈 鑫

收稿日期: 2022-11-04

  网络出版日期: 2023-05-11

基金资助

国家自然科学基金面上项目(32070782)

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Mechanism of Lactose Synthesis and Secretion and Its Correlation to Lactation Performance of Dairy Animals

  • LI Rongnuo ,
  • XIA Chao ,
  • DENG Lu , *
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  • College of Animal Science and Technology, Northwest A&F University, Yangling 712100, China
*associate professor, E-mail:

Received date: 2022-11-04

  Online published: 2023-05-11

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摘要

乳糖是乳汁中主要营养成分之一,在调控奶牛产奶效率过程中发挥重要作用。本文从奶牛的乳腺结构、乳糖合成底物、乳糖合成调控、乳糖分泌等方面进行了综述;系统阐明了乳糖通过维持囊泡渗透压调控奶牛产奶效率的生物学基础;并通过分析不同泌乳动物乳中乳糖含量与产奶量、乳蛋白含量、乳脂含量的相关关系,揭示了乳糖调控产奶效率的生物学功能,为提高奶牛产奶性能提供理论基础。

关键词： 乳糖合成; 乳成分; 产奶效率; 相关性分析

本文引用格式

李戎诺 , 夏超 , 邓露 . 乳糖合成与分泌的机制及其与奶畜泌乳性能的相关性分析[J]. 动物营养学报, 2023 , 35(5) : 2786 -2796 . DOI: 10.12418/CJAN2023.261

Abstract

Lactose is one of the main nutrients in milk and plays an important role in regulating milk production in dairy cows. This paper reviews lactose synthesis from the mammary gland structure, lactose synthesis substrate, lactose synthesis regulation and lactose secretion in dairy cows; systematically elucidates the biological theory of lactose in regulating milk production by maintaining vesicular osmotic pressure. We also analyzed the relationship between lactose content and milk yield, milk protein and milk fat contents in different lactating animals to reveal the biological function of lactose in regulating milk production, and to provide a theoretical basis and potential targets for improving milk production performance.

Key words： lactose synthesis; milk composition; milk production efficiency; analysis of correlation

奶牛健康、高效养殖是奶业可持续发展的基础。产奶效率低对奶业的制约作用日益突出,在优质饲料资源紧缺的时代大背景下,提升产奶效率无疑是现代牧场节本增效的重要突破口。研究表明,乳糖能以维持乳腺上皮细胞内渗透压的方式调控乳汁分泌量,是决定产奶效率的核心因子[1-2]。因此,如何在确保奶牛健康的条件下,最大限度地提高奶牛乳糖合成效率,是提升奶产量的重要手段。
乳糖作为人体所需的重要营养物质,可以显著促进人体中钙、铁、锌等矿物质的吸收[3-4],维持机体菌群(微生物)稳态[5-6],提高肠道有益菌的浓度[7]。尽管如此,乳糖在食品中的作用仍存在较大争议。例如,未消化的乳糖在人体肠道中通过影响渗透压使液体进入肠腔,诱发渗透性腹泻[8]。此外,肠道微生物群发酵乳糖,产生挥发性脂肪酸和气体(氢气、甲烷和二氧化碳)引起人体腹痛、腹胀,即乳糖不耐症[9]。随着乳糖不耐症研究不断深入,大多奶制品生产时会去除乳糖或加入能消化乳糖的益生菌[10-11]。
目前,在全球范围内,乳糖率作为牛奶记录程序框架中的常规记录指标[12],在大多数牧场群体检测中都会被记录。临床上通用的判断奶牛乳房炎的指标是乳汁中的体细胞数和病原微生物等。近期研究发现,乳糖率与乳中体细胞数呈负相关,可作为判断奶牛乳房炎症的重要生物指标,对预防牛乳房炎症有重要意义[13]。尽管如此,关于乳糖的潜在特质只有少量的研究,我们可获得的关于这一特性的表型和遗传方面的信息还很少。且少有报道明确指出乳糖含量与产奶量、乳蛋白、乳脂含量之间的关系。因此,本综述通过分析不同泌乳动物的乳成分数据,对乳中乳糖含量与产奶量、乳脂、乳蛋白含量之间的相关性及原因进行系统阐述,为进一步提高牧场产奶量,改善牛乳品质提供参考。

1 奶牛泌乳过程

奶牛泌乳是一个极其复杂的过程,在此过程中多种营养物质被摄取利用以合成乳蛋白、乳脂及乳糖等乳中干物质,该过程不仅需要多种营养物质的驱动,还需要乳腺在不同层面进行协调。乳腺由泌乳组织和结缔组织组成。乳腺腺泡是乳腺合成并分泌乳汁的功能单位,是由乳腺上皮细胞、基底膜、肌上皮细胞围成的空腔结构[14]。多个腺泡组成乳腺小叶[15]。乳汁在乳腺上皮细胞内合成,储存于腺泡腔内,肌上皮细胞通过收缩作用将乳汁从腺泡腔挤入乳导管,最终汇集于位于乳头正上方的乳腺池中[16]。泌乳时,乳中各种营养物质通过不同的运输方式进入乳腺腺泡中,在该区域融合为牛乳并保存在乳腺腺泡、泌乳管和乳腺池中[17],如图1所示。
图1 腺泡示意图

Fig.1 Schematic diagram of acini

2 乳糖的合成机制及生物学功能

2.1 乳糖合成的底物——葡萄糖的来源和吸收

乳糖是由葡萄糖和半乳糖合成的二糖,其中D-葡萄糖作为乳糖合成的专性前体物质,在哺乳期间需求较大。奶牛泌乳所需的葡萄糖主要来源于小肠吸收或肝脏糖异生。由于瘤胃这一独特的消化生理结构,奶牛通过小肠吸收获取的葡萄糖只占体内葡萄糖的30%左右,而肝脏糖异生需要合成体内高达70%的葡萄糖[18-19]。糖异生是机体将非糖物质转变成葡萄糖的过程,是保障乳腺葡萄糖供应、奶牛产奶量及奶品质的主要途径,该过程的紊乱往往导致奶牛哺乳期酮症、食欲不振及丙酸血症等代谢性疾病的发生[20]。因此,研究糖异生的调控机制对于奶牛的高效健康养殖至关重要。有研究表明,使用一定量甘油(生糖物质)代替饲粮中的玉米,或者补饲生物素和烟酰胺均可显著促进奶牛肝脏糖异生作用[21-22]。我们的研究发现,哺乳动物雷帕霉素靶蛋白(mammalian target of rapamycin,mTOR)信号通路在奶牛肝脏中处于高活性状态,且能够通过过氧化物酶体增殖物激活受体γ (peroxisome proliferator-activated receptor γ,PPARγ) -协同激活因子-1α (PGC1-α)上调糖异生基因的表达[23]。
葡萄糖主要由腺泡上皮的基底部细胞从血液中摄取,分为主动运输和被动运输2个转运过程[24-25],受葡萄糖转运蛋白(glucose transporter,GLUT)家族调节[26]。其中,GLUT1作为主要的葡萄糖转运蛋白,在乳腺和其他细胞、组织中大量表达[27-28]。有研究发现,GLUT1和GLUT4在乳腺中的表达体现出了时空表达差异特性[29-30]。同时表达GLUT4和GLUT1的乳腺上皮细胞在怀孕前占主导地位,而只表达GLUT1的乳腺上皮细胞在怀孕和哺乳期间增殖并成为主要细胞类型。

2.2 乳糖的合成与分泌

乳腺上皮细胞是乳糖合成的场所,胞质中充满了广泛的粗面内质网网络、膨大的高尔基体、大量的线粒体和含有酪蛋白胶束的囊泡[31]。泌乳期间,哺乳动物血液中60%~85%的葡萄糖被乳腺上皮细胞摄取,用于合成乳糖[32]。研究报道,乳腺中葡萄糖摄取与乳糖合成量和产奶量呈线性正相关[33⇓⇓⇓-37]。被摄取的葡萄糖一部分在GLUT的作用下进入高尔基体[38];另外一部分经过磷酸化形成葡萄糖-6-磷酸盐,并通过磷酸葡糖激酶、UDP-葡萄糖焦磷酸化酶2、半乳糖-1-磷酸尿苷酰基转移酶和UDP-半乳糖-4-表异构酶转化为UDP-半乳糖[39-40]。UDP-半乳糖在溶质载体家族35成员A2(solute carrier family 35 member A2,SLC35A2)或溶质载体家族35成员B1(solute carrier family 35 member B1,SLC35B1)的作用下进入高尔基体[38-39,41 -42]。高尔基体内含有由α-乳清蛋白(α-lactalbumin,LALBA)和β-1,4-半乳糖基转移酶(β-1,4-galactosyltransferase Ⅰ,β-4-GalT-Ⅰ)组成的乳糖合酶[43-44],能够将葡萄糖和UDP-半乳糖合成乳糖[45-46]。在人、奶牛、山羊以及猪的哺乳期乳腺研究中均发现LALBA的表达上调[47⇓⇓⇓-51],其能显著增加β-4-GalT-Ⅰ与葡萄糖的亲和力[52-53],且乳腺上皮细胞中的LALBA浓度与乳中蛋白质、脂肪和乳糖的含量呈正相关[54]。乳糖合成与分泌过程如图2所示。随后,乳糖会迅速与相关分泌蛋白如Asi-酪蛋白、t3-酪蛋白、LALBA和ft-乳球蛋白以及离子等一同包被形成分泌囊泡,由高尔基体通过微管和微丝逐步将囊泡迁移至细胞顶端膜,通过胞吐作用分泌到腺泡腔中[55⇓-57]。
图2 乳糖合成与分泌过程

Fig.2 Process of lactose synthesis and secretion

2.3 乳糖维持囊泡渗透压的生物学功能

在乳的形成过程中,水分子通过水通道蛋白(AQP)以跨膜运输的方式进入乳腺上皮细胞[58],该过程主要由乳糖所产生的渗透压梯度驱动。研究发现,影响囊泡内渗透压的物质主要有乳糖和无机离子如钠离子(Na+)、氯离子(Cl-)等[59]。乳糖分子结构式为C12H22O11,其中暴露在外的有8个亲水羟基,极易与水分子形成分子间氢键[60],具有强亲水性,结合水的能力远大于Na+等无机离子,如图3所示,因此囊泡渗透压主要由乳糖维持。当乳糖进入到分泌囊泡中,会使分泌囊泡中渗透压远大于细胞腔,此时大量的水分子进入囊泡以维持渗透压稳定。所以乳汁稀释不影响乳糖含量,但乳汁中的其他固形物含量会随乳汁量升高而下降。乳糖作为牛乳中的固形物对维持牛乳的渗透压起着决定性作用,所以乳糖合成效率对奶牛产奶量有显著影响[61⇓-63]。
图3 乳糖与钠离子水合能力比较

Fig.3 Hydratability comparison of lactose and Na+

3 乳糖含量及其他相关指标的相关性分析

3.1 乳糖含量与产奶量的相关性分析

为验证乳糖合成效率显著影响奶牛产奶量这一结论,本综述对Medhammar等[64]的测定数据进行了整理分析,得到了关于水牛、牦牛、驯鹿、驼鹿、双峰骆驼、单峰骆驼、麝牛、独龙牛、马、驴以及奶牛乳中乳糖和水分含量相关数据。详见表1。
表1 不同物种乳中乳糖和水分含量 g/100 g 奶

Table 1 Lactose and moisture contents in milk components of different species[64]

项目
Items		 奶牛
Cow		 水牛
Buffalo		 牦牛
Yak		 独龙牛
Mithun		 麝牛
Musk OX		 马
Mare		 驴
Donkey		 单峰骆驼
Dromedary
camel		 双峰骆驼
Bactrian
camel		 大羊驼
Llama		 羊驼
Alpaca		 驯鹿
Reindeer		 驼鹿
Moose
乳糖 Lactose 5.1 4.4 4.8 4.4 4.1 6.6 6.4 4.3 4.2 6.3 5.1 2.9 2.6
水 Moisture 88.1 83.2 82.6 78.6 83.6 89.8 90.8 89 84.8 84.8 83.7 67.9 76.8
我们对上述数据进行相关性分析,得出乳糖含量与水分含量拟合曲线的P值为0.002 6,呈极显著正相关关系。结果如图4所示。
图4 乳糖与水分含量拟合曲线

Fig.4 Fitted curve of lactose and moisture contents

3.2 乳糖和其他乳成分相关性分析

此外,我们还对上述物种乳成分数据进行了汇总,具体如表2所示,并对乳糖和其他乳成分含量进行了相关性分析,得出的相关系数,如表3所示。结果显示乳中的乳糖含量与乳蛋白含量之间存在极显著负相关关系,拟合曲线如图5;乳糖与乳脂含量之间存在极显著负相关关系,拟合曲线如图6所示;乳糖与饱和脂肪酸(saturated fatty acids,SFA)含量之间存在显著负相关关系,与单不饱和脂肪酸(monounsaturated fatty acids,MUFA)含量之间存在极显著负相关关系,与多不饱和脂肪酸(polyunsaturated fatty acids,PUFA)含量之间不存在显著相关关系,拟合曲线分别如图7、图8、图9所示。
表2 各物种乳成分数据

Table 2 Milk composition data for each species[64]

项目
Items		 奶牛
Cow		 水牛
Buffalo		 牦牛
Yak		 独龙牛
Mithun		 麝牛
Musk OX		 马
Mare		 驴
Donkey		 单峰骆驼
Dromedary
camel		 双峰骆驼
Bactrian
camel		 大羊驼
Llama		 羊驼
Alpaca		 驯鹿
Reindeer		 驼鹿
Moose
乳脂
Milk fat/(g/100 g奶)		 3.3 7.4 6.8 8.9 5.4 1.6 0.7 3.2 5.0 4.2 3.2 16.1 8.6
乳糖
Lactose/(g/100 g奶)		 5.1 4.4 4.8 4.4 4.1 6.6 6.4 4.3 4.2 6.3 5.1 2.9 2.6
乳蛋白
Milk protein/
(g/100 g奶)		 3.2 4.0 5.2 6.5 5.3 2.0 1.6 3.1 3.9 4.1 5.8 10.4 10.5
饱和脂肪酸
SFA/
(mg/100 g 奶)		 1 870 5 810 3 540 — — 550 310 2 080 2 237 2 940 — 11 240 4 540
单不饱和脂肪酸
MUFA/
(mg/100 g 奶)		 812 2 170 1 180 — — 432 77 2 200 1 307 1 260 — 2 730 2 540
多不饱和脂肪酸
PUFA/
(mg/100 g 奶)		 195 454 146 — — 560 77 277 153 150 — 566 1 460

“—”表示暂无数据。

“—” indicates that there is no data yet.

表3 乳糖与各乳成分含量之间的相关系数

Table 3 Correlation coefficients between lactose and each milk component contents

项目
Item		 乳脂
Milk fat		 乳蛋白
Milk protein		 饱和脂肪酸
SFA		 单不饱和脂肪酸
MUFA		 多不饱和脂肪酸
PUFA
乳糖 Lactose 0.002 2** 0.000 6** 0.022 4* 0.000 9* 0.063 1

“**”表示极显著相关(P<0.01);“*”表示显著相关(P<0.05)。表5同。

“**” shows extremely significant correlation (P<0.01); “*” shows significant correlation (P<0.05). The same as Table 5.

图5 乳糖与乳脂含量拟合曲线

Fig.5 Fitted curve of lactose and milk fat contents

图6 乳糖与乳蛋白含量拟合曲线

Fig.6 Fitted curve of lactose and milk protein contents

图7 乳糖与饱和脂肪酸含量拟合曲线

Fig.7 Fitted curve of lactose and SFA contents

图8 乳糖与单不饱和脂肪酸含量拟合曲线

Fig.8 Fitted curve of lactose and MUFA contents

图9 乳糖与多不饱和脂肪酸含量拟合曲线

Fig.9 Fitted curve of lactose and PUFA contents

为了进一步探究乳糖与乳蛋白和乳脂含量之间的关系,我们另外收集了大量其他泌乳动物的乳成分数据[65⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓-80],参见表4。同样对其进行整理分析得出的相关系数如表5所示。结果显示:乳糖与乳脂含量呈极显著负相关,拟合曲线如图10所示;乳糖与乳蛋白含量呈极显著负相关,拟合曲线如图11所示。该结论与上述分析结果一致。
表4 其他哺乳动物乳成分数据

Table 4 Data on milk composition in other mammals[65⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓-80]mg/mL

项目
Items		 人
Human		 狐猴
Lemur		 奶牛
Cow		 马
Mare		 山羊
Goat		 小鼠
Mouse		 大鼠
Rat		 兔
Rabbit		 狗
Dog		 迷你猪
Min-pig		 水牛
Buffalo		 绵羊
Sheep		 骆驼
Camel		 驴
Donkey
乳糖
Lactose		 70.0 81.0 50.0 63.7 41.0 26.0 26.0 18.0 34.5 49.5 40.5 50.0 43.0 66.0
乳脂
Milk fat		 36.0 18.0 43.5 12.1 40.0 205.0 134.5 152.0 79.0 88.5 71.5 70.0 40.0 10.5
乳蛋白
Milk protein		 13.5 26.0 36.0 21.4 41.0 114.0 93.0 103.0 70.5 62.5 37.0 57.5 33.0 17.0
表5 乳糖与各乳成分含量之间的相关系数

Table 5 Correlation coefficients between lactose and each milk component contents

项目 Item 乳脂 Milk fat 乳蛋白 Milk protein
乳糖 Lactose 0.006 6** 0.000 1**
图10 乳糖与乳脂含量拟合曲线

Fig.10 Fitted curve of lactose and milk fat contents

图11 乳糖与乳蛋白含量拟合曲线

Fig.11 Fitted curve of lactose and milk protein contents

以上结果显示乳糖与乳中水分含量呈正相关关系,即乳糖含量上升,水分含量相应增多(产奶效率提高),这是由于乳糖增加了分泌囊泡渗透压,导致乳中水分含量提高;另外乳糖含量和乳蛋白、乳脂含量均呈负相关关系,这是由于水分含量的增加稀释了乳中其他固形物成分。

4 小结与展望

乳糖不仅是乳汁中重要的营养物质,对泌乳过程中调控乳汁中其他营养物质也起着重要的作用。本文综述了乳糖合成与分泌的相关机制,总结了乳糖与渗透压的关系,阐明了乳糖调控产奶量及奶品质的重要作用,挖掘了乳糖合成效率[乳糖合成效率=乳糖量/(葡萄糖量×时间)][81]是评价奶牛产奶性能的新潜在靶标。
目前,奶牛养殖业面临的重要挑战之一是如何提高奶牛的产奶效率。奶牛产奶效率受宿主遗传因素、微生物、饲粮供应模式等多因素的共同调控[82⇓-84]。因此,运用现代生命科学与分子生物学等原理、方法和技术手段,与传统奶牛营养理论和技术密切结合,全面揭示乳糖合成效率的调控机制,建立乳糖合成效率和宿主、微生物组及营养素的精准联系,并构建提升奶牛乳糖合成效率的精准干预策略,是推进奶牛高效生产,保障我国“奶瓶子”稳定供给的核心。

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