Eggs are a valuable source of protein and fat in the human diet. Due to continuous improvement in the production performance of laying hens, the requirements regarding the feed energy of laying hens are increasing. Oils, which are the main energy sources in feed, exert a substantial effect on the production performance and egg quality of laying hens.
1. Introduction
Oils are the most commonly applied sources of energy in feed diets for laying hens and exert multiple effects, such as improving palatability, feed intake, animal immunity, and reducing morbidity
[1][2][3]. Compared with that included in the diet of broilers, the amount of oil added to hen feed is low because laying hens have a unique physiological state and are more prone to lipid metabolism disorders than broilers. Therefore, the appropriate proportion and type of oil addition is particularly important for the production performance, lipid metabolism, and egg quality of laying hens.
Eggs have an important economic value, are an excellent source of animal protein, and have become an important consumer product worldwide due to their low cost
[4][5]. With the continuous improvement in people’s living standards in developed and developing countries, interest in the internal nutritional composition of eggs, such as the content and proportion of omega-3 and omega-6 unsaturated fatty acids in the egg yolk, is increasing. Eggs consist of the albumen, yolk, and shell. The albumen is composed of 88% water, 11% protein, 0.2% lipids, and 0.8% minerals, and egg yolks comprise 48% water, 17.5% protein, 32.5% lipids, and 2% minerals
[6]. The lipids in the egg yolk mainly derive from the oils in the feed, and thus oils play a crucial role in the production performance and egg quality of laying hens. Due to continuous improvements in livestock and poultry production performance, the important role of dietary oils in livestock and poultry feed has attracted increasing attention from breeders and producers. Hence, methods to improve egg quality and enhance egg production performance to meet the diverse needs of consumers have become one of the focuses of the poultry industry.
Currently, the main oils used in the diets of laying hens are vegetable oils and animal oils. The vegetable oils commonly used in hen feeds are rich in a variety of unsaturated fatty acids (UFAs), which are important for the growth and development of laying hens, particularly linolenic acid and linoleic acid, which are essential fatty acids (EFAs) for birds. Linolenic acid deficiency hinders the development of hens and reduces production performance, whereas linoleic acid is deposited directly into the egg yolk and thereby increases the weight of the egg by increasing the weight of the yolk
[7]. Animal oils (mainly triglycerides (TGs)) are extracted from animal fat tissue and, when added to the diet, these fats increase the UFA contents of eggs
[8][9]. The addition of oils to the diets of laying hens has become an effective method to promote animal growth, increase the FCR, and improve poultry health. The fatty acid composition and physicochemical properties of different types of oils and their physiological effects on laying hens vary substantially
[10][11]. As an example, the addition of 6% canola oil will decrease egg production and significantly increase thiobarbituric acid reactive substance (TBARS) levels in eggs on the 21st day of the addition trial (corn–soybean meal compared with 6% rapeseed oil)
[12]. Some studies have shown that the addition of 3% and 5% rapeseed oil to the basal diet of 24-week-old Hyland brown chickens has no significant effect on the egg weight
[13], but the addition of 2%, 4%, and 6% rapeseed oil to the diet of 40-week-old brown chickens reduces feed intake, egg production, and egg weight
[12]. Compared with the results obtained with the addition of 3% fish oil, olive oil, grape seed oil, or soybean oil, the addition of equal amounts of canola oil to the diets of laying hens does not significantly change the egg production, egg weight, feed intake, or FCR but increases the linolenic acid content in the egg yolk
[14]. In addition, oils have adhesion properties that reduce dust in hen feed production, promote better particle aggregation, reduce wear on machinery, and result in both low waste and feed savings. The unique aroma of oils may improve the palatability of feeds and enhance their flavor, which leads to increased animal intake. The addition of oils with low heat gain to feed serves to effectively reduce heat stress, which improves feed utilization and reduces mortality
[3].
2. An Overview of Oils
As already mentioned, feed oils are divided into vegetable oils and animal oils. The commonly used vegetable oils in feeds mainly include soybean oil, rapeseed oil, palm oil (PO), and linseed oil, and the animal oils commonly used in feeds mainly include lard, poultry fat, tallow, and fish oil
[15]. Different oils have different ratios of saturated fatty acids (SFAs) to UFAs. In general, vegetable oils contain more UFAs than animal oils, which contain more SFAs
[16]. Therefore, vegetable oils are currently used at higher levels than animal oils in egg production, but no major difference in energy has been observed. The energy and nutrients that animals obtain from feed depend on the species and age of the animal and vary depending on the quality and chemical composition of the fat
[17].
The quality, fatty acid composition, total energy, and price of various oils differ
[18]. Various countries and organizations worldwide have established corresponding standards for quality measurements. The main indicators are divided into sensory, technical, and microbial characteristics
[19]. The microbial indicators are the total number of colonies and the number of colonies of coliforms,
Salmonella,
Shigella,
Staphylococcus aureus,
Streptococcus hemolyticus, and other common pathogenic strains
[20].
3. Mechanism Underlying the Effects of Dietary Oils on the Production Performance and Egg Quality of Laying Hens
3.1. Mechanism Underlying the Effect of Oils on Lipid Metabolism in Laying Hens
During the egg-laying period, the amount of lipid mobilization and synthesis in laying hens is in dynamic equilibrium, and the amount of fatty feed added to the diet of laying hens is generally less than 10%. The amount of lipids that laying hens obtain from feed is approximately 3 g per day, and 5–6 g of lipids is needed for the formation of each egg. Therefore, some lipids are synthesized endogenously by laying hens and then deposited in the ovaries through transport via the blood and other circulatory systems to finally form egg yolk material
[21]. The fatty acid composition of the diet directly affects the composition of the egg yolk. This observation also serves as the theoretical basis for the inclusion of lipid additives to regulate the egg production performance and egg quality of laying hens. The main organ in which poultry synthesize lipids is the liver, and the fat needed to maintain egg production performance is also mainly synthesized by the liver
[22]. The raw materials for synthesizing lipids are fatty acids, glycerol, and cholesterol. These raw materials can be directly supplied by exogenous feed or converted from protein and glucose in the feed. EFAs cannot be synthesized in the body of poultry and can only be obtained from feed
[23]. The liver assembles synthetic TGs, cholesterol, and apolipoproteins to form lipoproteins with different diameters and densities, which are responsible for the targeted transport of lipids. Lipoproteins are generally divided into five types according to their density: chylomicrons (CMs), very low-density lipoprotein (VLDL), intermediate-density lipoprotein (IDL), low-density lipoprotein (LDL), and high-density lipoprotein (HDL)
[24]. Laying hens have a very strong lipid metabolism in their bodies, particularly during the peak period of egg production. As one of the essential nutrients for animal growth and one of the main components of egg yolk, oils play an important role in regulating the production performance of laying hens and their egg quality. Therefore, the production performance and egg quality of laying hens may be feasibly improved in theory and practice by supplementing the diet with different oils. The development and production of high-quality eggs through dietary oil nutrition regulation technology provides people with foods with balanced fatty acid nutrition, which is not only beneficial to health but also provides a new approach to the production of high value-added poultry products.
3.2. Providing Essential Fatty Acids and Affecting Lipid Metabolism
During the peak period of laying hens, the intensity of lipid metabolism in the body increases, and the oils in feed further enter the eggs through the ovaries in the body. The commonly used oils in feed, particularly vegetable oils, are rich in PUFAs (polyunsaturated fatty acids). Most of these fatty acids exhibit an important relationship with the growth and production performance of laying hens. Therefore, fatty acids have become highly valued raw materials in compound feed for laying hens. As hens have physiological requirements for laying eggs, poultry, particularly laying hens, are significantly more dependent on EFA than pigs and ruminants. In particular, linoleic acid and linolenic acid are EFAs that play a decisive role in the growth and reproduction of laying hens
[25]. Insufficient dietary EFAs will significantly affect growth and development, affect ovarian development, and reduce the body weight, egg weight, and fertilized egg hatchability
[26][27]. Dietary supplementation with linoleic acid increases the egg weight
[28][29]. In addition, when the diet of laying hens meets the demand for linoleic acid, the addition of exogenous oil significantly increases the egg weight independently without affecting the dietary metabolizable energy and the linoleic acid content
[30]. Vegetable oils rich in PUFAs reduce the liver fat content or alleviate fatty liver syndrome in laying hens, which is conducive to lipid metabolism in laying hens, whereas excessive consumption of animal oils or starch aggravates fatty liver syndrome in laying hens or significantly increases hepatic lipid deposition in laying hens
[31][32].
3.3. Effect of Oils on the Nutritional Composition of the Yolk
Due to improvements in people’s living standards, people are currently paying increasing attention to the nutritional content of eggs. The UFA content is an important basis for determining the nutritional value of poultry eggs. A certain amount of UFAs in the diet exerts a positive effect on animal health
[33]. The lipids of the egg contain SFAs (30–35%) and UFAs (30–33%) and, among these, 0–45% and 20–25% of UFAs are MUFAs and PUFAs, respectively
[34]. In addition, the fatty acid composition of the egg yolk substantially affects the flavor of the egg
[35]. The use of feed rich in n − 3 fatty acids increases the content of n − 3 fatty acids in eggs. Therefore, the fatty acid composition and content of egg yolk can be adjusted through the feed formula to produce eggs rich in PUFAs
[36]. Eicosapentaenoic acid (EPA, C20:5n − 3) and docosahexaenoic acid (DHA, C22:6n − 3) are both n − 3 PUFAs, and supplementation with EPA and DHA significantly improves brain and cardiovascular function
[37][38]. Eggs rich in n − 3 PUFAs have been sold in the United States, Canada, Australia, and other countries and have shown excellent nutritional benefits
[39]. Published studies have confirmed the effects of oils on the fatty acid composition of egg yolks and their promotion of human health
[40][41][42][43].
After the addition of suitable oils, the composition and proportion of FAs in the yolk will significantly change. FAs mainly contain SFAs, MUFAs, and PUFAs. Egg yolks are rich in PUFAs, which are divided into ω − 3 and ω − 6 PUFAs. Among ω − 3 PUFAs, those that exert the most significant effects on the human body are DHA and EPA
[44]. Studies have shown that both ω − 3 and ω − 6 fatty acids exert anticancer effects, and the anticancer effect of ω − 3 polyunsaturated fatty acids is significantly better than that of ω − 6 polyunsaturated fatty acids
[45]. In addition, the ideal intake ratio of ω − 6 PUFAs and ω − 3 PUFAs would be between 4:1 and 10:1
[46]. The method for increasing the content of specific unsaturated fatty acids in eggs by adding unsaturated fatty acids to the diet of laying hens has become increasingly mature. People use laying hens as a fatty acid converter to achieve nutritional optimization. For example, the addition of 5% rapeseed oil to the diet of laying hens significantly increases the weight of the egg yolk and the contents of DHA and n − 3 FAs in the egg yolk
[13]. Han et al.
[47] showed that the addition of 1.5 to 4.5% sesame oil to the diet increases the UFA content in egg yolks and reduces the blood lipid levels of laying hens.
3.4. Immunomodulatory Effect of Oils
Fatty acids in oils play an important role in maintaining poultry health. The classic indices include ∑SFAs, ∑MUFAs, ∑PUFAs, ∑n − 6 PUFAs, ∑n − 3 PUFAs, and n − 6 PUFAs/n − 3 PUFAs
[48]. When the body is exposed to external antigens, the secretion of lymphokines and antibodies and the production of new immune cells depend on the participation of fat
[49]. PUFAs in the diet are involved in the synthesis of biofilm structures and are precursors for a variety of physiologically active substances, such as eicosanoids and leukotrienes
[50]. These substances are particularly important for poultry and may participate directly or indirectly in physiological processes, particularly immune system processes
[51]. PUFAs significantly improve the inflammatory response in the body and reduce the serum TNF-α and IL-1β levels
[52]. The intake of PUFAs by laying hens significantly increases their serum lysozyme activity and improves immune function
[53]. Peroxisome proliferator-activated receptors (PPARs) are nuclear hormone receptors and transcription factors. Long-chain UFAs may mediate lipid metabolism and the immune inflammatory response in laying hens through the PPAR pathway
[54]. In addition, the FAs in the egg yolk are mainly synthesized by the liver, which is substantially affected by the fatty acid composition of the diet. The addition of linoleic acid and linolenic acid to the diet of laying hens significantly increases the contents of arachidonic acid and DHA in the egg yolk
[55]. Various PUFAs, such as linolenic acid and arachidonic acid, are components of the cell membrane structure and are essential for maintaining the integrity of the cell membrane structure and function. In addition, EFAs are precursors of eicosanoids, such as prostaglandins, prostacyclins, thromboxane, and leukotrienes, and these substances are used for blood coagulation, nerve signal transduction, embryonic development, reproduction, immune response, and bones in poultry. These compounds play an important role in physiological processes such as embryonic development and are also involved in the transport of body fluids, the activation of certain enzymes, and the metabolism of lipids, particularly cholesterol, which are very important for maintaining and improving the production performance and egg quality of laying hens
[1][56].
In modern egg production, laying hens are usually under high-intensity production pressure. Heat stress and oxidative stress (OS) are common factors that affect the production performance of laying hens
[57][58]. The digestion and absorption efficiency of oil is outstanding, and its heat increment is significantly lower than that of carbohydrates and protein. This property may substantially reduce the heat dissipation burden of the animal’s body in summer, particularly in high-density laying hen production, and thereby alleviate heat stress and improve the FCR
[59]. The double bond in oil compounds exerts a protective effect on easily oxidized substances in the body (such as vitamin E), which might improve the body’s antioxidant capacity and relieve OS. Therefore, the benefits of oils for animal health are likely attributed to their effects on the immune and antioxidant systems in the body
[60].
3.5. Effects of Dietary Oils on the Structure and Function of Intestinal Microbes
The structure and function of the gut microbiota are critical to the health and production performance of laying hens. The acquisition and establishment of the intestinal microbiota throughout the poultry production cycle exerts a substantial effect on the development and physiological regulation of the intestine and maintains intestinal homeostasis (i.e., nutrition, metabolism, immunity, and integrity of the intestinal barrier), which results in the optimization of the host’s energy absorption and use efficiency
[61]. The processes of the digestion and absorption of nutrients are closely related to the intestinal microbiota, and the nutrient absorption, FCR, and production performance of the host are affected by the composition and diversity of the microbiota
[62][63]. The microbial composition of the poultry intestine is affected by many internal and external factors, such as the host and environment. Feeding and nutrition management are important factors that affect the composition and function of intestinal microbes in poultry. Therefore, the intestinal microbiota can be adjusted by regulating the feed composition
[64]. At the phylum level, the gut microbiota of chickens includes hundreds of species, including microbial species dominated by
Firmicutes, Bacteroides,
Proteobacteria, and Actinomycetes [65][66]. Oil additives such as α-linolenic acid (ALA), DHA, and glycerin and its derivatives (such as glycerol monocaprylate and glycerol monolaurate) exert a positive effect on the intestinal microbial composition, egg production performance, egg quality (e.g., egg weight and egg yolk FA composition), reproductive performance, and body health of laying hens, which indicates the potential role of dietary oils in improving the microbial community, lipid metabolism, and health of laying hens
[67][68][69][70]. For example, studies have found that the addition of linseed oil and algae oil to the diet increases the types of Firmicutes (such as
Faeculus,
Clostridium, and
Ruminococcus) microorganisms, and these microorganisms are closely related to FA metabolism
[68][69]. Different types and amounts of lipids affect the health of the host to different extents, which may be due to their varying effects on the composition of the gut microbiota
[71][72].
4. Relationship between Fatty Acids in Human Diets and Health Indicators
Studies have found that eating the right amount of egg every day can reduce the risk of metabolic syndrome in adults over the age of 40 and has a significant positive effect on blood glucose and triglyceride levels in men. The reason for this may be that fatty acid composition has an immense impact on the dietary factors of fats and oils. The quality of dietary fat mainly depends on the ratio of n − 6 to n − 3 fatty acids
[73]. The fat content in the human diet has related effects on common diseases such as the risk of coronary heart disease (CHD) and type 2 diabetes
[74]. The thrombogenic index (TI) and atherogenicity index (AI) are the most commonly used indices to assess the composition of FAs. The AI and TI were proposed by Ulbritcht and Southgate in 1991
[48], where the AI represents the sum of SFAs and the relationship with the sum of unsaturated fatty acids (UFAs), which also represents the atherosclerotic potential of saturated fatty acids; the TI describes the thrombotic potential of FAs, predicts the tendency of thrombus formation in blood vessels, and indicates the functions of different FAs. At present, AI indicators have been widely used to evaluate the quality of meat, eggs, dairy products, the nutrient composition of oil, and other livestock products because they are not only crucial for the nutritional value of eggs but they also affect shelf life. In addition, the TI was used due to the correlation between FAs and human health
[49][75].
5. Effect of Different Oils in Diets on the Production Performance and Egg Quality of Laying Hens
The use of specific dietary oils depends on the price and availability of feed ingredients. As mentioned above, different dietary oils affect the production performance and egg quality of laying hens by regulating the lipid metabolism, immune function, and intestinal microbial composition, and this phenomenon provides a theoretical basis for the oil-mediated regulation of the laying performance and egg quality of laying hens. However, due to differences in the physical and chemical properties, lipid composition, main functional substances, quality, and added amount of different types and sources of oils, their effects on the health, laying performance, and egg quality of laying hens are also different. In general, for a single oil, the digestion and absorption of vegetable oils is significantly better than that of animal oils, but balanced and combined oils are better than single oils. Table 1 lists the effects of animal and vegetable oils on egg quality, production performance, and nutrients.
Table 1. Studies showing the effects of different oils on egg quality and production performance.
Supplements
|
Results (Production Performance/Egg Quality/Nutritional Content)
|
Author/s
|
Soybean oil
|
Its addition increases the egg production rate and feed conversion rate; increases the amount of calcium deposited in eggshells and significantly reduces the rate of broken or soft eggs; improves the glycolipid metabolism of laying hens; improves the antioxidant capacity of hens; improves the nutritional value of eggs, reduces the content of cholesterol in eggs, and reduces the TI and AI.
|
[10][11][76][77][78][79][80]
|
Rapeseed oil
|
The addition of 1–5% rapeseed oil has no significant effect on production performance. The addition of 2–6% rapeseed oil reduces egg production, and the addition of 5% rapeseed oil significantly increases the egg yolk weight and contents of DHA and n − 3 FAs in the egg yolk.
|
[11][12][13][81][82][83][84][85]
|
Linseed oil
|
The addition of 2–3% linseed oil does not significantly alter production performance; the addition of 5% or higher concentrations of linseed oil significantly reduces the body weight and egg production rate due to a high n − 3 PUFA deposition efficiency.
|
[86][87][88][89][90][91][92][93][94]
|
Palm oil
|
With increases in the level of red palm oil added (1–3%), the production performance of laying hens improves, the color of the egg yolk significantly improves, and the nutritional value of the egg yolk and the flavor of the egg also improve. The addition of this oil also reduces the TG levels in the egg yolk and increases the MUFA contents in the egg yolk.
|
[95][96][97][98][99][100][101]
|
Cottonseed oil
|
The addition of this oil significantly reduces the egg production rate, average egg weight, and FCR of laying hens, hardens the egg yolk, causes rubbery eggs, and changes the composition of the protein in the yolk granules and plasma.
|
[102][103][104]
|
Microalgae oil
|
The addition of this oil does not negatively affect the production performance and egg quality, increases the ratio of n − 3/n − 6 PUFAs in the egg yolk, and optimizes the FA composition of the egg yolk.
|
[92][105][106][107][108][109][110][111][112][113][114][115][116][117][118][119][120][121]
|
Fish oil
|
Excess fish oil significantly reduces the production performance and egg quality of laying hens and increases the deposition of EPA and DHA in the egg yolk.
|
[108][122][123][124][125]
|
Lard
|
Excess lard reduces the egg production rate and induces fatty liver development but increases the yellow color of the egg and the FA content of the egg yolk.
|
[32][126][127][128]
|
Cod liver oil
|
By promoting the absorption of calcium, it will increase the proportion of unsaturated fatty acids in the egg yolk; poor quality cod liver oil will reduce the quality of eggs, adding too much cod liver oil will make the eggs have a significant fishy smell.
|
[129][130][131]
|
Oil oxidation
|
The addition of oxidized oil reduces the production performance of hens, destroys the integrity of the yolk globules of cooked eggs, and makes animals more susceptible to oxidative stress.
|
[132][133][134]
|