动物营养学报 >

2025 , Vol. 37 >Issue 3: 1503 - 1515

DOI: https://doi.org/10.12418/CJAN2025.129

综述

黑水虻幼虫的营养价值及其在动物生产中的应用

  • 姜强 , 1, 2 ,
  • 曲扬华 2 ,
  • 孙玉丽 2 ,
  • 吕增鹏 , 1, *
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  • 1 中国农业大学动物科学技术学院,畜禽营养与饲养全国重点实验室,北京 100193
  • 2 中粮营养健康研究院有限公司,北京市畜产品质量安全源头控制工程技术研究中心,营养健康与食品安全北京市重点实验室,北京 102209
*吕增鹏,副教授,硕士生导师,E-mail:

姜 强(2001—),男,湖南常德人,硕士,从事动物营养与饲料科学研究。E-mail:

Copy editor: 武海龙

收稿日期: 2024-08-22

  网络出版日期: 2025-03-13

基金资助

现代农业产业技术体系(CARS-41)

现代农业产业技术体系(CARS-40)

中粮集团战略前瞻研发项目“昆虫蛋白关键技术开发及应用”(2024-C1-F001)

中国农业大学2115人才培育发展支持计划项目

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Nutritional Value of Black Soldier Fly Larvae and Its Application in Animal Production

  • JIANG Qiang , 1, 2 ,
  • QU Yanghua 2 ,
  • SUN Yuli 2 ,
  • LYU Zengpeng , 1, *
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  • 1 State Key Laboratory of Animal Nutrition and Feeding, College of Animal Science and Technology, China Agricultural University, Beijing 100193, China
  • 2 Beijing Key Laboratory of Nutrition, Health and Food Safety, Beijing Engineering Research Center of Livestock Product Quality and Safety Source Control, China Oil and Foodstuffs Corporation Nutrition and Health Research Institute, Beijing 102209, China
*associate professor, E-mail:

Received date: 2024-08-22

  Online published: 2025-03-13

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摘要

黑水虻幼虫具有繁殖快、适应性强、生物转化效率高等生物学特性,含有丰富的蛋白质、脂肪、微量元素、维生素等营养物质,以及几丁质、抗菌肽等活性物质,是一种优质饲用资源型昆虫,在替代豆粕等蛋白质饲料原料上有着巨大潜力。此外,由于黑水虻幼虫独特的腐生生活方式,在解决有机废弃物和资源再利用方面具有重要应用价值。本文综述了黑水虻幼虫的生物学特性、营养价值、基质优化策略及其在动物生产中的应用进展,推荐了黑水虻幼虫在各类动物饲粮中的适宜用量,并探讨了产业面临的挑战和未来需要深入研究的方向,旨在为黑水虻幼虫饲料化的开发及其在动物生产中的应用提供参考。

关键词： 黑水虻幼虫; 营养价值; 饲料化; 动物生产

本文引用格式

姜强 , 曲扬华 , 孙玉丽 , 吕增鹏 . 黑水虻幼虫的营养价值及其在动物生产中的应用[J]. 动物营养学报, 2025 , 37(3) : 1503 -1515 . DOI: 10.12418/CJAN2025.129

Abstract

The black soldier fly larvae have the biological characteristics of rapid reproduction, strong adaptability and high bioconversion efficiency, rich in protein, fat, trace elements, vitamins and other nutrients, as well as active substances such as chitin and antimicrobial peptides. It is a kind of high-quality forage resource-based insect, which has great potential to replace soybean meal and other protein raw materials. In addition, due to the unique saprophytic lifestyle of the larvae of the black soldier fly, it has important application value in solving organic waste and resource reuse. This article reviews the biological characteristics, nutritional value, substrate optimization strategies and application progress of black soldier fly larvae in animal production, recommends the appropriate amount of black soldier fly larvae in various animal feeds, and discusses the challenges faced by the industry and directions for future in-depth research, aiming to provide a reference for the development of black soldier fly larvae in feed utilization and its application in animal production.

Key words： black soldier fly larvae; nutritional value; feed utilization; animal production

随着近年来我国畜牧业的高速发展以及居民消费水平的不断提高,养殖规模逐渐增大,饲料需求量不断增加,饲料产业面临巨大压力。蛋白质饲料是饲料生产的核心原料之一,而我国豆粕、鱼粉等蛋白质饲料原料大部分依赖进口[1],相关数据显示,2023年我国进口大豆9 941万t,鱼粉162万t[2]。因此,寻找新型蛋白质饲料资源成为畜牧行业可持续发展面临的严峻挑战。当下,人工养殖的昆虫已成为解决蛋白质饲料资源匮乏问题的关键突破口。其中,黑水虻幼虫拥有独特的腐生生活方式,能够高效利用有机废弃物,转化率最高可达到40%[3-4]。
黑水虻幼虫营养物质丰富,是猪、家禽和水产动物优质的蛋白质饲料来源[5]。Håkenåsen等[6]研究表明,饲粮中添加5%~20%黑水虻幼虫对断奶仔猪出栏重、采食量和料重比没有负面影响。赵燕等[7]通过添加2%~6%黑水虻幼虫干发现,肉鸡回肠绒毛高度以及蛋白质等营养物质在胸肌的沉积率明显提高。黑水虻幼虫替代25%~75%的鱼粉对罗非鱼生长性能、血清生化指标以及体成分组成没有负面影响[8]。2017年,欧盟正式颁布法规,明确黑水虻幼虫作为水产动物饲料中一种合法且可持续的蛋白质饲料来源,并在2021年扩展至猪和家禽饲料,填补了绿色循环经济中后端资源再利用的缺口[欧盟委员会法规(EU)2017/893;欧盟委员会法规(EU)2021/1372]。鉴于黑水虻幼虫具有较高的营养价值,在动物饲料生产中的应用前景广阔,本文对其营养价值及在动物生产中的应用进行了综述,以期为挖掘黑水虻幼虫饲料化潜力提供参考。

1 黑水虻概述

1.1 黑水虻生物学特性

亮斑扁角水虻,俗称黑水虻,属昆虫纲、双翅目、短角亚目、水虻科、扁角水虻属。该物种广泛分布于全球各地的温带、热带以及大部分亚热带区域。在我国黑水虻生长地集中在华南、西南以及中部地带,是一种重要的资源型昆虫[7]。黑水虻的生活史可分为卵、幼虫、预蛹、蛹、成虫5个时期。黑水虻幼虫营腐生生活,生长速度快,适应性极强,能够将各种有机废弃物转化为优质的蛋白质和脂质,在处理城镇餐厨垃圾、畜禽粪便等方面具有极高的应用价值[9⇓-11]。黑水虻幼虫经过预蛹和蛹羽化为成虫,在成虫阶段可不进行采食,只需饮水就能生存,降低了与食物源相关病原体的接触机会,从而大幅度减少了饲养过程中生物风险[12]。

1.2 黑水虻幼虫的营养价值

黑水虻幼虫具有较高的营养价值,其体内含有丰富的蛋白质、油脂、矿物质以及维生素等营养物质,此外还能产生几丁质、抗菌肽等活性物质。近年来,有关黑水虻幼虫营养物质含量以及应用方式的研究陆续开展。陈柏宇等[13]报道了黑水虻的老熟幼虫的营养物质含量,其中粗蛋白质含量占干重比例为40%~45%,粗脂肪含量占干重比例为23%~36%,粗灰分含量变化幅度较大,占干重比例为3.10%~19.70%。有研究对黑水虻幼虫蛋白质的生物学价值进行了综合评估,表明其蛋白质含量高,氨基酸组成平衡,生物学价值高,是潜在的、持久性的蛋白质来源[14-15]。
黑水虻幼虫的饲料化应用形式一般为脱脂与全脂2种,不同的加工处理方式对黑水虻幼虫的营养成分有影响。表1总结了不同脱脂程度的黑水虻幼虫及主要蛋白质饲料原料的营养物质组成,脱脂黑水虻幼虫的粗蛋白质含量在51.81%~55.30%,进行高度脱脂粗蛋白质含量可达到65.50%,高于豆粕粗蛋白质含量(49.44%),低于鱼粉粗蛋白质含量(67.53%);全脂黑水虻幼虫的粗蛋白质含量在41.88%~46.38%,低于豆粕和鱼粉。脱脂黑水虻幼虫的粗脂肪含量在4.60%~18.00%,远高于豆粕粗脂肪含量(1.40%),其平均值(12.12%)高于鱼粉粗脂肪含量(10.36%);全脂黑水虻幼虫粗脂肪含量26.28%~38.60%,远高于豆粕和鱼粉。黑水虻幼虫的粗灰分含量2.70%~12.98%,其平均值(7.45%)高于豆粕粗灰分含量(7.19%),但远低于鱼粉粗灰分含量(17.15%)。黑水虻的粗纤维含量(4.10%)介于豆粕(7.43%)和鱼粉(0.26%)之间。
表1 黑水虻幼虫及主要蛋白质饲料原料的营养物质组成(干物质基础)

Table 1 Nutrient composition of black soldier fly larvae and major protein feed ingredients (DM basis) %

项目
Items		 黑水虻幼虫Black soldier fly larvae 豆粕
SBM		 鱼粉
FM
全脂
FF		 脱脂
DF		 部分脱脂
PDF		 高度脱脂
HDF
粗蛋白质
CP		 46.38
±1.441)		 43.10
(0.6)2)		 41.88
±1.32		 42.05
±1.64		 54.40
±1.02		 51.813) 55.30 65.50 49.44 67.53
粗脂肪
EE		 32.28
±1.78		 38.60
(2.3)		 26.28
±1.80		 30.34
±2.13		 11.15
±1.49		 14.71 18.00 4.60 1.40 10.36
粗灰分
Ash		 8.02
±0.49		 2.70
(0.3)		 12.98
±0.65		 6.36
±0.14		 8.66
±0.29		 7.27 — — 7.19 17.15
粗纤维
CF		 — — — — — — — — 7.43 0.26
必需氨基酸Essential amino acid
赖氨酸
Lys		 2.49
±0.03		 2.30 — 2.38
±0.43		 2.98
±0.36		 1.96 2.10 2.52 3.11 4.87
蛋氨酸Met 0.68
±0.01		 0.71 — 0.37
±0.12		 0.52
±0.08		 0.62 0.65 0.86 0.68 1.85
异亮氨酸
Ile		 2.11
±0.11		 1.91 — 1.54
±0.43		 2.12
±0.20		 2.24 1.85 2.40 2.21 2.73
亮氨酸
Leu		 2.75
±0.09		 3.06 — 2.55
±0.33		 3.48
±0.46		 3.30 2.86 3.67 3.86 4.77
组氨酸
His		 1.75
±0.15		 1.38 — 2.91
±0.73		 4.04
±1.12		 0.47 1.23 1.63 1.42 1.54
精氨酸
Arg		 2.66
±0.03		 1.99 — 1.61
±0.41		 2.01
±0.57		 1.64 2.15 2.70 3.57 4.10
缬氨酸
Val		 3.18
±0.05		 2.82 — 2.41
±0.38		 2.98
±0.53		 3.58 2.72 3.45 2.17 3.27
苯丙氨酸
Phe		 1.97
±0.20		 1.64 — 1.55
±0.27		 1.93
±0.32		 1.69 1.66 2.18 2.55 2.64
苏氨酸Thr 1.39
±0.02		 1.62 — 1.43
±0.09		 1.94
±0.26		 1.93 1.72 2.18 1.98 2.75
色氨酸
Trp		 0.72
±0.01		 0.54 — 2.49
±0.79		 3.01
±1.21		 0.02 — — 0.66 0.67
钙Ca 1.60
±0.03		 0.10 — — — — — — — —
磷P 0.99
±0.01		 0.40 — — — — — — — —
几丁质
Chitin		 — 6.70
(1.3)		 6.24
±1.96		 — — — 5.00 6.90 — —
参考文献
References		 [16] [17] [18] [21] [21] [20] [19] [19] [22]

1):平均值±标准差;2):平均值(变异系数);3):平均值。

1): mean±standard deviation; 2): mean (coefficient of variation); 3): mean。

黑水虻幼虫拥有丰富的氨基酸谱,含有动物生长发育所需的全部必需氨基酸,其中赖氨酸、异亮氨酸、亮氨酸、精氨酸、缬氨酸含量最丰富,其平均值分别为2.39%、2.02%、3.10%、2.11%、3.02%;缬氨酸平均含量高于豆粕(2.17%),接近鱼粉(3.27%),异亮氨酸平均含量与豆粕、鱼粉较接近;赖氨酸、亮氨酸、精氨酸平均含量均低于豆粕、鱼粉;组氨酸平均含量为1.92%,高于豆粕(1.42%)、鱼粉(1.54%);蛋氨酸平均含量为0.63%,与豆粕(0.68%)相当,但低于鱼粉(1.85%);苯丙氨酸、苏氨酸平均含量分别为1.80%、1.74%,均低于豆粕(2.55%、1.98%)、鱼粉(2.64%、2.75%);色氨酸平均含量为1.36%,高于豆粕(0.66%)、鱼粉(0.67%)。黑水虻幼虫的氨基酸谱与豆粕、鱼粉相接近,并且其组成和比例不会因为脱脂工艺而变化[21]。
黑水虻幼虫的粗脂肪含量能达到26%~38%,黑水虻预蛹的粗脂肪含量亦能达到39.70%[23]。表2总结了黑水虻幼虫的脂肪酸组成,其中饱和脂肪酸含量最高,最丰富的是月桂酸(32.96%~45.97%)。已有研究报道,月桂酸能在动物体内转化为月桂酸甘油酯,具有抗菌、抗病毒活性,但过高的月桂酸可能会对营养物质的消化利用有一定的不利影响,例如黑水虻幼虫油完全替代罗非鱼饲料中的豆油会使脂质的表观消化率降低30%,能量沉积率降低12%[24⇓⇓⇓-28]。棕榈酸(12.21%~15.08%)、肉豆蔻酸(6.72%~8.70%)、硬脂酸(2.53%~3.90%)含量也较高;单不饱和脂肪酸中最丰富的是油酸(11.24%~22.13%),其次是棕榈油酸(1.91%~2.75%);多不饱和脂肪酸中最丰富的是亚油酸(12.59%~14.07%),其次是亚麻酸(1.51%~1.65%)[29⇓-31]。但各脂肪酸丰度并不是稳定的,这与饲养基质密切相关。胡俊茹等[32]探讨了餐厨垃圾和鸡粪对黑水虻幼虫脂肪酸组成的影响,发现以餐厨垃圾为基质饲养黑水虻幼虫,不饱和脂肪酸含量高于饱和脂肪酸,并且油酸含量高达30.70%,月桂酸含量仅为9.70%;相反,鸡粪饲养的黑水虻幼虫以饱和脂肪酸为主,月桂酸含量为41.30%。黑水虻幼虫脂肪酸组成依赖于饲养基质组成,餐厨垃圾和鸡粪脂肪含量具有明显差异,这使得生产出的黑水虻幼虫脂肪酸组成不同。因此,通过调节基质营养物质成分可有效影响黑水虻幼虫营养物质的含量,从而优化其营养价值。
表2 黑水虻幼虫脂肪酸组成(占总脂肪酸比例)

Table 2 Fatty acid composition of black soldier fly larvae (proportion of total fatty acids) %

项目Items 全脂FF 脱脂DF
月桂酸C12∶0* 45.971) 32.96±0.672) 37.55 6.21±0.55
肉豆蔻酸C14∶0* 8.70 6.72±1.50 6.73 1.25±0.05
葵酸C10∶0* 0.86 1.01±0.02 — 0.19±0.01
十五碳酸C15∶0* 0.15 0.02±0.00 — 0.01±0.00
棕榈酸C16∶0* 12.21 15.08±0.50 15.60 3.85±0.08
硬脂酸C18∶0* 2.53 3.20±0.08 3.90 0.57±0.09
花生酸C20∶0* 0.10 0.05±0.00 — 0.01±0.01
肉豆蔻油酸C14∶1△ — 0.21±0.01 — 0.04±0.02
棕榈油酸C16∶1△ 1.91 2.75±0.11 2.53 0.54±0.06
油酸C18∶1△ 11.24 22.13±0.38 17.88 4.16±0.28
亚油酸C18∶2n-6# 14.07 12.59±0.27 12.72 2.43±0.02
亚麻酸C18∶3n-3# 1.65 1.64±0.01 1.51 0.29±0.02
花生四烯酸C20∶4# 0.14 0.37±0.01 0.07 0.06±0.01
二十碳一烯酸C20∶1△ 0.06 0.11±0.01 0.21 0.02±0.01
二十碳二烯酸C20∶2# 0.01 0.11±0.00 0.08 0.01±0.00
二十碳五烯酸C20∶5n-3# — 0.21±0.01 — 0.05±0.01
二十二碳六烯酸C22∶6n-3# — 0.05±0.00 — 0.01±0.01
饱和脂肪酸Saturated fatty acids 70.72 59.22±0.27 65.01 12.14±0.49
单不饱和脂肪酸Monounsaturated fatty acids 13.41 25.33±0.42 20.62 4.78±0.33
多不饱和脂肪酸Polyunsaturated fatty acids 15.87 15.03±0.27 14.37 2.87±0.07
其他脂肪酸Other fatty acids — 0.43±0.27 1.09 0.01±0.00
参考文献References [31] [21] [29] [21]

*:饱和脂肪酸;△:单不饱和脂肪酸;#:多不饱和脂肪酸。1):平均值;2):平均值±标准差。

*: saturated fatty acid; △: monounsaturated fatty acid; #: polyunsaturated fatty acid. 1): mean; 2): mean±standard deviation.

黑水虻幼虫含有丰富的矿物质,其中钙、铁、锌、铜等微量元素的含量较高,铬、镉、铅和砷等重金属元素和有害元素在虫体内有少量积累[33]。此外,幼虫含有丰富的胆碱、B族维生素,但维生素A含量较少[34]。为适应有害环境,幼虫体内可产生几丁质、抗菌肽等多种活性物质,具有重要的应用价值。研究表明,黑水虻幼虫的活性物质对革兰氏阴性菌和革兰氏阳性菌具有广谱抗菌活性,在抵抗病原体上具有重要作用[35⇓-37]。

1.3 黑水虻幼虫营养基质优化策略

挖掘黑水虻幼虫的高价值潜能,改善黑水虻幼虫营养品质,营养基质优化是核心路径。各类有机废弃物复配平衡基质营养成分是最简单的优化策略,如与单一牛粪相比,鸡粪复配可显著增加黑水虻幼虫产量,基质的利用效率大大提高[38]。Rehman等[39]使用牛粪与豆腐渣按2∶3复配,黑水虻幼虫生长性能得到明显改善,对有机废弃物的利用率也得以提高。Nyakeri等[40]将粪便污泥与餐厨垃圾、啤酒糟、香蕉皮等按照7∶3混合,与Wang等[41]和Spranghers等[17]所报道的结果相比,黑水虻幼虫粗蛋白质含量分别提高了6.94%和9.28%,粗脂肪含量分别降低了43.95%和32.28%,虫体营养结构得到改善。幼虫脂肪受基质油脂和糖类含量的影响较大,通过调控混合物蛋白质和非纤维类碳水化合物水平,限制脂肪和纤维素含量,可提高黑水虻幼虫转化能力并减少营养参数的差异,营养基质中碳氮比为14∶1~18∶1,有利于幼虫对有机质的利用与转化[38,42-43]。
基质中应用添加剂可辅助黑水虻幼虫生长,提高其营养价值。郝建伟等[44]研究显示,猪粪中添加4%的紫马铃薯皮粉能够有效促进黑水虻幼虫生长,可使虫体粗蛋白质含量提升30%以上。Mazza等[45]利用黑水虻幼虫与其伴生细菌联合处理鸡粪,发现幼虫的生长速度有明显提高,虫体营养成分如蛋白质、脂肪等含量更加丰富,且鸡粪的残留量更少。Gu等[46]研究发现,在微藻中添加枯草芽孢杆菌、酵母菌和植物乳杆菌可提高黑水虻幼虫生物转化率、虫体氨基酸和饱和脂肪酸含量。此外,控制饲喂速率[200~250 mg/(d 只)]和调整基质粒径(0.5 mm)等同样是有效的基质优化策略[40,47]。综合运用上述营养基质优化策略,有利于进一步挖掘黑水虻幼虫作为优质生物资源的潜力。

2 黑水虻幼虫在动物生产中的应用

2.1 黑水虻幼虫在猪生产中的应用

目前,黑水虻幼虫的猪饲料化主要应用在仔猪、生长育肥猪上[15,48⇓-50]。研究表明,饲粮中添加黑水虻幼虫粉对仔猪生长性能、血清生化指标、饲料消化率以及肠道健康状况等无负面影响[48-49]。有研究报道了黑水虻幼虫粉对断奶仔猪肠道菌群和黏膜免疫应答的影响,发现使用黑水虻替代50%的鱼粉能够显著提高回肠和盲肠中的乳杆菌、双歧杆菌等益生菌丰度,增加乳酸盐以及短链脂肪酸含量,并能上调肠道黏膜屏障功能、发育及抗炎因子相关基因表达,抑制炎症通路,保护仔猪肠道健康[51]。
Zhu等[52]研究报道,生长猪饲粮中添加黑水虻幼虫粉可提高日增重、背膘厚度和肌内脂肪含量,降低肌肉滴水损失,改善生长性能以及肉品质。在育肥猪上的研究显示,饲粮中使用黑水虻幼虫粉替代50%~100%的鱼粉能够明显提高育肥猪的生长性能,改善胴体性状和肉品质[50]。Yu等[53]研究表明,饲粮中添加4%的黑水虻幼虫除了能够提高育肥猪的生长性能和胴体性状外,还能通过上调脂肪合成基因表达显著增加肌内脂肪沉积。张放等[54]通过在育肥猪饲粮中添加黑水虻幼虫粉替代鱼粉,发现对干物质和粗蛋白质的消化率无不利影响。Yu等[55]研究发现,饲粮中添加4%和8%的黑水虻幼虫粉对改善育肥猪的肠道微生物菌群结构与代谢产物以及黏膜免疫功能具有积极作用,4%组乳酸菌、罗斯氏菌属和梭状芽孢杆菌群ⅩⅣa的数量显著增高,8%组丁酸盐浓度增加,这有益于育肥猪的健康生长。此外,猪对黑水虻幼虫呈现出更高的偏好性,相较于普通饲料更倾向于采食黑水虻幼虫[56]。这可能与黑水虻幼虫独特的气味、适口性以及高含量的蛋白质、油脂有关[56⇓-58]。因此,黑水虻幼虫在猪生产中可替代部分豆粕或鱼粉,建议在仔猪饲粮中替代50%的鱼粉,生长育肥猪饲粮中可替代50%~100%的鱼粉,以降低饲料成本。

2.2 黑水虻幼虫在蛋鸡生产中的应用

鸡天生具备捕食虫子的习性,能够有效地将昆虫中的蛋白质转化为营养丰富的鸡蛋等产品,从而实现资源的高效利用与转化。国内外对黑水虻幼虫作为蛋鸡饲料原料的研究较多。Maurer等[59]使用脱脂黑水虻幼虫替代蛋鸡饲粮中50%或100%的豆粕,发现对生产性能以及蛋品质没有负面影响。Heuel等[60]使用黑水虻幼虫蛋白质与油脂分别替代饲粮中的豆粕和豆油,发现对蛋鸡的采食量、产蛋性能没有显著性影响。饲粮中添加3%和6%全脂黑水虻虫干对46~53周龄蛋鸡生产具有积极作用,能够改善蛋品质,降低次品蛋率,同时提高免疫能力[61]。波文英等[62]研究表明,补饲黑水虻幼虫活虫、冻虫和虫干能够改善鸡蛋的哈氏单位和蛋壳厚度,但提高了料蛋比、采食量等生产性能指标。饲粮中使用黑水虻幼虫替代20%、30%的鱼粉,同样能够提高鸡蛋的蛋白高度和哈氏单位,改善蛋品质[63]。
对蛋鸡抗氧化和免疫指标方面的研究显示,饲粮中添加2%~6%的黑水虻幼虫粉能够线性提高蛋鸡血清中总抗氧化能力(T-AOC)、总超氧化物歧化酶(T-SOD)活性及免疫球蛋白A(IgA)、免疫球蛋白M(IgM)、白细胞介素-4(IL-4)以及肝脏中T-AOC、T-SOD和谷胱甘肽过氧化物酶(GSP-Px)活性[64]。使用黑水虻幼虫替代饲粮中25%和50%的豆粕可降低蛋鸡血液中的胆固醇、甘油三酯含量,提高球蛋白含量,50%替代组的干物质、有机物和粗蛋白质消化系数降低,可能是由于几丁质所造成的负面影响[65]。使用黑水虻幼虫完全替代饲粮中的豆粕能够提高24~45周龄蛋鸡的十二指肠绒毛高度、盲肠丁酸含量以及麦芽糖酶、蔗糖酶活性,但降低了蛋白质消化率和生产性能[66]。黑水虻幼虫含有大量的月桂酸、几丁质和抗菌肽等活性物质,以上研究结果提示了其作为饲料原料具有益生功能,在抗菌、抗氧化以及提升免疫力等方面可发挥重要作用。但黑水虻幼虫高含量的几丁质可能会阻碍家禽对营养物质的消化吸收,影响生产性能[67]。因此,黑水虻幼虫替代饲粮中的部分豆粕或鱼粉对蛋鸡的生产性能有不同的影响,推荐蛋鸡饲粮中添加剂量为3%~6%。

2.3 黑水虻幼虫在肉鸡生产中的应用

贾友刚等[68]报道,使用黑水虻幼虫粉替代饲粮中2%~8%的豆粕,对肉鸡的生产性能、屠宰性能均无显著影响,6%豆粕替代组胸肌滴水损失最低,肠道厚度高、弹性好,肠道黏膜脱落与出血性肠炎的发生率均为最低水平,在一定程度上提升肉品质以及肠道健康水平。许多研究得到相似的研究结果,Pieterse等[69]研究发现,在肉鸡饲粮中添加15%的黑水虻幼虫蛋白粉,对屠宰性能和肉品质没有负面影响。Hartinger等[70]研究发现,黑水虻幼虫粉替代饲粮中15%的豆粕对肉鸡的生产性能以及健康状况没有影响。因此,在肉鸡饲粮中使用黑水虻幼虫替代豆粕是可行的,但以上研究的黑水虻幼虫添加或替代量均在较低水平。肉鸡不同于蛋鸡,其肠道和免疫功能发育尚未发育成熟,抵抗外界刺激的能力相对较差。随着黑水虻幼虫添加量的增加,可能会对肉鸡性能产生不利影响。有研究报道,使用全脂黑水虻幼虫粉替代50%~100%的豆粕,会显著降低肉鸡的生长性能,并对胴体性状以及鸡肉风味和品质产生不利影响[71]。Hartinger等[72]亦发现,在饲粮中添加15%的脱脂黑水虻幼虫粉会降低肉鸡的出栏重、蛋白质和部分氨基酸的表观回肠消化率,增加死亡率。亦有研究报道,使用脱脂黑水虻幼虫粉替代饲粮中15%的豆粕会降低肉鸡的饲料转化率以及肠道的绒毛高度,并提高隐窝深度,对肠道形态产生不利影响[72]。因此,黑水虻幼虫在饲粮中高剂量使用可能会对肉鸡生产造成不利影响,其在肉鸡饲粮中最佳的添加或替代量应在10%以下,以确保对肉鸡的生长性能、屠宰性能和肉品质不会产生负面影响。

2.4 黑水虻幼虫在水产动物生产中的应用

鱼粉是水产饲料的核心原料,然而随着海洋渔业资源的日益枯竭和环保意识的提升,传统鱼粉的供应面临着严峻挑战。通过昆虫蛋白质替代鱼粉是推动水产饲料行业可持续性发展的有效策略。Rawski等[73]研究表明,黑水虻幼虫可刺激鱼类采食,提高生产性能,饲料中添加30%的黑水虻幼虫粉可使西伯利亚鲟鱼体重、增重率和特定生长率分别提高30.19%、36.46%和15.18%。Magalhães等[74]研究报道,欧洲鲈鱼饲料中45%的鱼粉可被黑水虻预蛹粉替代,对其生长性能、血清生化指标以及消化酶活性没有影响,并且提高了精氨酸、组氨酸和缬氨酸的表观消化率以及蛋白质利用率。Belghit等[75]研究亦发现,饲料中添加脱脂黑水虻幼虫粉替代33%~100%的鱼粉,对大西洋鲑鱼的生长性能和营养物质消化率没有显著影响,且对鱼体营养成分以及肝脏功能没有负面作用。Li等[76]研究表明,在大西洋鲑鱼饲料中使用黑水虻幼虫粉完全替代鱼粉,会使鱼体近端肠道上皮细胞脂肪变形程度降低,远端肠道相对重量增加,对改善肠道健康有一定的积极作用。但是,Sangsawang等[6]研究发现,黑水虻幼虫完全替代鱼粉会引起罗非鱼肝脏和肠道组织损伤。这可能是由于不同来源的黑水虻幼虫体内几丁质含量存在差异所致,高含量的几丁质可能会引发肝脏组织出血坏死以及肠道炎症[6,23]。Zhou等[77]研究发现,建鲤饲料中使用黑水虻幼虫替代鱼粉达到50%~100%时,肌肉内氨基酸组成较为平衡;鱼体、肌肉和器官内饱和脂肪酸含量升高,多不饱和脂肪酸含量降低;但是,血清T-AOC显著下降,这可能与脂肪酸组成发生变化有关,影响了建鲤的抗氧化能力。此变化与黑水虻幼虫脂肪酸组成有关,黑水虻幼虫含有丰富的饱和脂肪酸(如月桂酸),在鱼类体组织中极易富集,使饱和脂肪酸和不饱和脂肪酸的比例失衡,进而影响鱼肉的营养结构[78-79]。此生产问题可以通过调整黑水虻幼虫的饲养基质来调控虫体脂肪酸的组成,进而实现肉品中饱和脂肪酸与不饱和脂肪酸比例的均衡[80-81]。
在虾、蟹等其他水产动物的研究中同样证明了采用黑水虻幼虫替代饲料中鱼粉的可行性。采用脱脂黑水虻幼虫粉替代饲料中30%、50%和70%的鱼粉,可线性提高凡纳滨对虾的体增重、特定生长率,降低饲料转化率,改善生长性能[82]。张文锐等[83]研究表明,青蟹饵料中添加黑水虻幼虫替代25%和50%的鱼粉能够使体重提高69%以上;但替代量达到75%时,会降低青蟹部分必需氨基酸和多不饱和脂肪酸含量,对营养品质造成不利影响。类似研究表明,采用黑水虻幼虫替代饲料中60%以上的鱼粉时,凡纳滨对虾的体增重率和特定生长率会明显降低[84]。因此,饲料中高替代比例的黑水虻幼虫同样会对虾、蟹等水产动物来带负面影响,这可能与黑水虻幼虫含有的几丁质有关,饲料中适量的几丁质可提高甲壳类动物的生长性能以及抗应激能力,但几丁质含量过高可能会对肠道组织结构造成损伤,影响肠道对营养物质的消化和吸收,最终抑制甲壳类动物的生长[85]。因此,黑水虻幼虫在鱼类饲料中100%替代鱼粉具有可行性,但在实际生产中仍需关注完全替代引起的负面作用。对于虾、蟹等甲壳类动物饲料中,建议替代25%~50%的鱼粉。

2.5 挑战与展望

目前,黑水虻幼虫饲料化应用面临着诸多挑战。对于不同的动物,黑水虻幼虫使用尚无明确的标准,并且由于饲养基质的差异导致虫体营养成分不稳定,从而使其应用受到限制。因此,需通过进一步研究黑水虻幼虫对各种动物生产性能、肠道健康、免疫与抗氧能力等指标的综合影响,确定最佳需要量,建立饲料应用标准;并通过营养基质优化策略提高虫体营养价值与稳定性,保证可持续性生产。黑水虻幼虫成本价格高昂,远高于豆粕,并且养殖规模化不足,每年产出数量不能满足饲料产业的大规模需求。因此,需加速培育优质虫源,扩大人工养殖规模;并通过评估不同有机废弃物的饲养潜力来扩大基质来源,因地制宜地完善供应链,从而达到降本、提质、增量的目的。此外,通过把控虫体品质,推广黑水虻应用于仔猪精料、宠物饲料等高端产品来提高其商品价值。黑水虻幼虫生产应用中还存在一些瓶颈,如全脂黑水虻幼虫在原料储存、加工和运输上存在不足,脱脂虫粉终端销售价格与成本脱节,虫沙、油脂、几丁质和抗菌肽等副产物的利用缺乏深入研究与应用标准。因此,未来研究需要深入探讨加工方法、应用标准以及市场策略,以推动其在动物生产中的广泛应用。

3 小 结

综上所述,黑水虻幼虫营养价值高,是一种新型的蛋白质饲料资源,在饲料产业中有着巨大的可持续应用潜力,能够替代动物饲料中部分豆粕和鱼粉,从而有效缓解蛋白质饲料资源匮乏的问题。然而,在实际生产过程,黑水虻幼虫的应用仍存在诸多瓶颈。为此,需要持续进行深入研究与技术创新,逐步构建绿色、高效、可持续的产业链。

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