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Original Research

Frontiers in Animal Science ·

Volume 7 - 2026 | https://doi.org/10.3389/fanim.2026.1861221

Substitution with Lentinus edodes chaff enhances the health and growth performance of Altay sheep, and improvement of serum immune function and oxidative resistance

  • Lele Wang1,2
  • Shuzhen Tang1,2
  • Rongyan Qin1,2
  • Abuxiaheman Mubalake1,2
  • Yan Wang1,2
  • Xiangyu Chen3
  • Yanfeng Liu1,2*corresponding author
  • Wenqi Wang1,2*corresponding author
  1. 1Feed Research Institute, Academy of Animal Husbandry of Xinjiang Uygur Autonomous Region, Urumqi, China
  2. 2Xinjiang Key Laboratory of Herbivorous Livestock Feed Biotechnology, Urumqi, China
  3. 3Academy of Animal Husbandry of Xinjiang Uygur Autonomous Region, Urumqi, China
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Abstract

The aim of this experiment was to evaluate the effects of Lentinus edodes chaff on the growth performance, apparent digestibility of nutrients and serum biochemical indicators of male Altay lambs. Forty lambs, weighing 29.9 ± 3.8 kg and approximately 4 months old, were randomly assigned to four treatments, each consisting of ten replicates. The first group was used as a control group and was fed only the basal diet. The other three groups, which were used for comparison, were fed the basal diet, which was supplemented with Lentinus edodes chaff at 10, 20, or 30% of the dry matter, respectively. Compared to the control group, adding Lentinus edodes chaff to the sheep’s diet resulted in a 20% increase in ADG and FI (p < 0.05). Furthermore, a noticeably lower F/G ratio was observed in the 20% group. The Apparent DM digestibility declined in 30% (p < 0.05), whereas CP digestibility rose in 20% (p < 0.05) and fell in 30% (p < 0.01). NDF digestibility was elevated in both 20% and 30% (p < 0.01). In terms of rumen fermentation parameters, the total VFA concentration was higher in the 20% group (p < 0.01). In comparison, the 10% group exhibited lower levels of GLB, ALT, LDH and CK (p < 0.05). In contrast, the levels of IgG and IgA in the 10% group were found to be significantly higher than those in the control (p < 0.05). Concurrently, the levels of LDL, IgG and IgA in the 20% group exhibited higher values in comparison (p < 0.05). Furthermore, the levels of HDL, LDL, IgG and IgA in the 30% group were higher than the levels in the control group (p < 0.05). Furthermore, a substantial increase was observed in the levels of NO, SOD, GSH-Px and T-AOC in the three experimental groups (p < 0.01). whereas IL-2 levels were significantly decreased (p < 0.05), concurrently with a decrease in MDA (p < 0.01). The 20% dietary inclusion level may improve growth, protein utilisation and systemic antioxidant status in growing Altay sheep. The optimal level of Lentinus edodes chaff is 20%.

Keywords

1 Introduction

Global demand for food is rising due to ongoing population growth. It is estimated that agricultural and animal husbandry output will need to increase by 70-100% to meet future global food demand (Yang et al., 2021a; Fytsilis et al., 2024). To meet this growing demand, more animal feed ingredients will need to be produced. However, the sustained increase in roughage and maize prices, coupled with the chronic shortage of high-quality forage, severely limits the sustainable intensification of China’s ruminant production systems. Consequently, identifying and evaluating alternative feed ingredients has emerged as a central research priority in this field (Yang et al., 2025).

Edible mushrooms are currently the fifth largest crop sector in China, after cereals, oilseeds, vegetables and fruit. In 2023, the national output exceeded 43 million tonnes. Projections indicate that by 2026, the global edible mushroom economy will surpass 20 million tonnes (Patrícia et al., 2025). The nutritional value and bioactive components of edible mushrooms render their by-products, particularly spent mushroom substrate, a promising alternative feed source for ruminants. The edible mushroom industry is expanding. A significant volume of spent mushroom substrate (SMS) is produced. SMS is the residue that remains following the cultivation process. SMS represents an intricate composite of organic material and mycelium remnants, a consequence of the cultivation process employed in the production of mushrooms. It is a well-documented fact that the cultivation of commercially important mushrooms, including Oyster mushrooms (Pleurotus Ostreatus) and Lentinus edodes, is frequently undertaken on lignocellulosic materials (Panagiota and Seraphim, 2023; Kousar et al., 2024; Zhang et al., 2024). A common substrate is sawdust from hardwood. As research into waste utilisation increases, a growing body of studies is focusing on the use of cottonseed shells, corn cobs, wheat bran and tea leaves as substrates for mushroom growth (Guo et al., 2022a). The efficiency levels of the culture medium employed in producing edible mushrooms are reported to be between 17-25% (Mahari et al., 2020), which indicates that 3–5 kg of SMS is produced for every 1 kg of mushrooms harvested. SMS is disposed of during mushroom production through a combination of open burning, landfill and composting, often mixed with animal manure. As a result, its full value has not yet been realised because of problems in the environment, including water, soil and air pollution (Gong et al., 2019; Jiang et al., 2017; Lam et al., 2019). Various strategies have been explored to address this challenge and manage SMS effectively. These include using SMS as a biofertiliser (Liu et al., 2014; Waqas et al., 2018; Lou et al., 2017), in biological bogs (Huang et al., 2020), as a soil amendment (Qiao et al., 2011; Siracusa et al., 2017), as a substrate for commercial enzyme fermentation (Economou et al., 2020; Matute et al., 2012), in bioremediation (Faraco et al., 2009), and in biofuel production (Pérez-Chávez et al., 2019; Ribas et al., 2009). SMS is currently being used to an alarming extent in China’s main mushroom-producing areas, with serious environmental problems resulting from this.

As a result, there is an urgent requirement to establish efficient techniques for utilising this used substrate. Significant amounts of mycoprotein and trace elements have been found in SMS, according to research. Furthermore, it is abundant in bioactive substances. These include polysaccharides and triterpenoids. The effects exhibited by the substance are comparable to those of mushrooms. These include immune enhancement, growth promotion and antioxidant properties (Liu et al., 2025). SMS has been demonstrated to enhance the balance of digestive tract flora in livestock and poultry, reduce feeding costs, and enhance economic returns (Paredes et al., 2009). Furthermore, SMS has been found to possess a distinctive mushroom flavour that enhances its palatability (Yuan et al., 2022a). Adding spent substrate to chicken diets has been reported to enhance feed conversion and fat breakdown, improve antioxidant capacity and increase serum immunity levels (Mahfuz et al., 2019; Chuang et al., 2020) SMS has been reported to be an effective functional feed additive for improving body weight, average daily gain, blood profiles, immunoglobulin and inflammation in weaning pigs (Waewaree et al., 2020). In goat diets, supplementing with SMS has been observed to increase nutrient apparent digestibility and decrease rumen pH and glucose concentration with no negative effects on plasma metabolism (Pin and Anusorn, 2018). The findings of these studies suggest that spent substrate has the potential to function as a promising feed additive in the context of livestock and poultry production. However, research on cottonseed hull SMS in sheep diets is scarce (Liu et al., 2025). The present study investigates the effects of feeding sheep diets supplemented with varying doses of SMS on growth performance, nutrient apparent digestibility, rumen fermentation variables and serum biochemical indicators. The objective of the present study was to explore the potential of SMS in ovine production, with the aim of facilitating the discovery of new feed resources, providing a novel strategy to address the issue of competition between humans and animals for food, and contributing to the advancement of environmentally sustainable animal husbandry, while concomitantly reducing feeding costs.

2 Materials and methods

2.1 Ethical approval

The experiments followed the protocols and methods authorised by the Xinjiang Animal Welfare and Ethics Committee of the Academy of Animal Sciences in China (No. 1 20251009). All animal treatments and experiments were carried out in accordance with the guidelines for ethical review of animal welfare in the national standards of the People’s Republic of China.

2.2 Animals and the administration

A substrate consisting of 79% cottonseed hulls, 20% wheat bran and 1% lime, all of which were sourced from an edible fungus planting cooperative on the outskirts of Hutubi County in Xinjiang, was used to cultivate the Lentinus edodes chaff. Table 1 describes the chemical composition of the Lentinus edodes chaff used in the experiment.

Table 1. The chemical composition* of Lentinus edodes chaff used in the experiment.

Nutrient levels†Content
Dry matter48.78
Crude protein13.61
Ether extract0.26
Organic matter88.46
Neutral detergent fiber62.17
Acid detergent fiber57.06
Calcium1.65
Total phosphorus0.27

*The chemical composition was based on dry matter basis.

†Nutrient levels were obtained from chemical analysis.

The Altay sheep used in this study were four months old and weighed an average of 29.7 kilograms. They were selected and purchased from the Xinjiang Zhongxin Qi Agricultural and Animal Husbandry Professional Cooperative Association. A complete feed mixture was created by evenly mixing coarse feed (e.g. Lentinus edodes chaff) with concentrate feed and other raw materials. The basal diet (Table 2) was designed to meet the requirements for growing sheep. These requirements are set out in the Meat Sheep Feeding Standard (NY/T 816-2004).

Table 2. Composition and nutrient levels of diets.

IngredientsLevels %Nutrient composition %†Levels %
01020300102030
Corn34.030.627.223.8Metabolic energy (MJ/kg)9.929.439.619.18
Bran3.02.72.42.1Dry matter91.2190.5690.6789.86
Soybean meal10.09.08.07.0Crude protein14.0613.5013.1312.27
NaCl1.00.90.80.7Ether extract3.253.433.923.41
Calcium hydrogen phosphate1.00.90.80.7Crude ash9.188.998.918.24
Premix*1.00.90.80.7Neutral detergent fiber38.2642.7441.0945.01
Alfalfa30.027.024.021.0Acid detergent fiber28.3030.1530.0532.73
Rice straw20.018.016.014.0Calcium0.880.930.880.90
Lentinus edodes chaff010.020.030.0Total phosphorus0.540.510.550.50
Total100100100100

*Premix provided the following per kg in basic diets: 30000 IU of vitamin A, 5000 IU of vitamin D, 200 IU of vitamin E, 55 mg of iron, 70 mg of manganese, 70 mg of zinc, 0.2 mg of cobalt, 12 mg of copper, 0.2 mg of selenium, 0.5 mg of iodine.

†The metabolic energy in the nutritional level were calculated, and the rest were measured values.

The Xinjiang Zhongxin Qi Agricultural and Animal Husbandry Professional Cooperative Association was the site of the animal testing. Prior to the purchase of the test animals, the animal house was thoroughly cleaned and disinfected. A 1.0×1.2 m pen was used to house each animal individually, with each sheep constituting one experimental unit, with food being given to them twice a day at 10:00 and 19:00. The animals were free to eat and drink during these times.

2.3 Trial design

The sheep were divided into four dietary treatments at random, utilising the equivalent body weight principle, with ten sheep per group. Consequently, the initial group functioned as a control group and was provided solely with the basal diet. The remaining three groups, which constituted the experimental set, ingested basal diets in which 10, 20 or 30% of the basal diet had been substituted with Lentinus edodes chaff. The feeding experiment ran for a total of 40 days, including a 10-day pre-experimental phase and a 30-day experimental phase.

2.4 Measurements

2.4.1 Growth performance

The quantity of feed consumed was meticulously recorded on a daily basis, with the amount of feed being adjusted every three to four days to allow for a residual amount of five to ten per cent. The residual feed from the previous day was collected prior to morning feeding, then dried and weighed to determine dry matter intake (DMI). The weight gain of each sheep was measured on days 1, 30 and at the conclusion of the experiment. The calculation of average daily gain (ADG) and feed-to-gain (F/G) ratios was performed based on data pertaining to ADG and daily dry matter intake.

2.4.2 The measurement of apparent digestibility

The provision and refusal of feed was meticulously documented on a daily basis. During the experiment’s concluding week, a selection of six sheep from each treatment group exhibiting comparable DMI and BW were subjected to a seven-day digestibility test. The apparent digestibility of nutrients was determined by means of the acid-insoluble ash method. The faeces of each sheep were collected on two occasions per day, using the total manure collection method. Following the mixing process, 10% of the feces were meticulously collected as composite samples and stored in a zip-lock bag. A total of 10 millilitres of nitrogen fixation solution (10% diluted sulphuric acid) was added to every 100 g of fresh faeces over the course of five days of continuous faeces collection. Each sheep’s faecal sample was then combined individually and frozen at -20 °C.

The ANKOM 200i fibre analyser (ANKOM Technologies, Inc., Fairport, NY, USA) was employed to analyse the contents of the DM, organic matter (OM), and CP in accordance with AOAC (2000). The neutral detergent fibre (NDF) and acid detergent fibre (ADF) were analysed using the ANKOM filter bag technique. The model employed for the purpose of determining digestion was as follows: nutrient digestibility=100-100x(% indicator in feed x% nutrient in feces)/(% indicator in feces x% nutrient in feed).

2.4.3 Rumen fluid collection, rumen fermentation parameters testing

At the 30-day mark of the experiment, rumen fluid samples were collected. Rumen fluid samples of six sheep from each group were collected at a fixed time point (prior to the morning feeding) to minimise diurnal variation and ensure consistent comparison across treatment groups. The fluid was then filtered through four layers of cheesecloth. Subsequently, 100 mL of rumen fluid was collected and thoroughly amalgamated. The samples were then divided into 10 mL centrifuge tubes, stored in liquid nitrogen, and finally stored at -20 °C. The ruminal pH was determined by means of a digital pH meter (Testo205 type). The test utilised a solution of phenol-hypochlorite to determine the concentration of NH3-N. The volatile fatty acids (VFA) in the rumen were measured using a gas chromatography (Agilent 6890 N Gas Chromatograph) that was fitted with a capillary column (19091N-213; Agilent).

2.4.4 Serum biochemical indicators

The experiment was conducted over a period of 30 days. Blood samples were obtained from the jugular vein prior to the commencement of feeding. A total of 10 mL of blood was collected from six sheep per treatment group. The serum was separated by a low-speed centrifuge, with the upper layer transferred to 1.5 mL tubes and stored in a -20 °C refrigerator until testing. The following indicators were measured using an automatic biochemical analyser (Mindray BS-420, Shenzhen Mindray Biomedical Electronics Co., Ltd.): total protein (TP), albumin (ALB), globulin (GLB), urea (UREA), total cholesterol (TC), triglyceride (TG), high-density lipoprotein (HDL), low-density lipoprotein (LDL), glucose (GLU), immunoglobulin G (IgG), immunoglobulin M (IgM), immunoglobulin A (IgA), aspartate aminotransferase (AST), alanine aminotransferase (ALT), creatine kinase (CK), lactate dehydrogenase (LDH), alkaline phosphatase (ALP). We detected the levels of interleukin-2 (IL-2), malondialdehyde (MDA), nitric oxide (NO), superoxide dismutase (SOD), glutathione peroxidase (GSH-Px) and total antioxidant capacity (T-AOC) in serum using an enzyme-labelled analyser (Huawei Delang DR-200 BS, Wuxi Huawei Delang Instrument Co., Ltd.).

2.5 Statistical analysis

The statistical software SPSS (2019) was used to analyse growth performance, serum biochemical parameters, rumen fermentation parameters and digestibility data. A one-way analysis of variance (ANOVA) was conducted with dietary treatment as the fixed effect. Because the treatments represented increasing concentrations of Lentinus edodes chaff (0, 10, 20, and 30%), orthogonal polynomial contrasts were used to test the linear, quadratic, and cubic effects of LEC inclusion level. Statistical significance was evaluated at p ≤ 0.05.

3 Results

3.1 Growth performance and apparent digestibility

Table 3 shows the effects of Lentinus edodes chaff in the diet on final body weight (FBW), body weight gain (BWG), average daily gain (ADG), feed intake (FI) and feed gain ratio (FGR). The 4-month-old sheep’s initial body weights showed no significant differences across the treatment groups (averaging 29.7 ± 2.5 kg), indicating that the animals were randomly assigned to the experimental groups. No significant variations in FBW and FGR were revealed by ANOVA between the experimental groups, but substantial differences (p < 0.05) were found in BWG, ADG and FI. The 20% group had higher BWG, ADG and average daily FI than the 0% group (p < 0.05). Furthermore, the total and average daily FI were considerably higher than in the control, with a significant linear increase (p = 0.025 and P = 0.026) as Lentinus edodes chaff inclusion level increased.

Table 3. Effects of Lentinus edodes chaff on the growth performance in Altay sheep.

ItemsLevels %p-value
0102030LinearQuadraticCubic
Initial body weight (kg)29.35 ± 2.5129.11 ± 2.5330.38 ± 2.8030.01 ± 2.280.4280.9430.451
Final body weight (kg)37.38 ± 3.2837.86 ± 3.1040.21 ± 3.4838.69 ± 2.990.2290.3960.285
Body weight change (kg)8.03 ± 1.59b8.75 ± 1.11ab9.83 ± 1.49a8.68 ± 1.48ab0.1940.0830.284
Average daily gain(g/d)267.50 ± 53.08b291.66 ± 37.06ab327.50 ± 49.57a289.38 ± 49.46ab0.1940.0830.284
Total FI (kg)45.17 ± 8.37b49.35 ± 9.79ab50.36 ± 10.09ab50.78 ± 10.24a0.0250.2880.744
Average daily FI(kg/d)1.50 ± 0.27b1.64 ± 0.32ab1.67 ± 0.33a1.69 ± 0.34a0.0260.2840.749
Feed/gain ratio5.84 ± 1.185.73 ± 0.745.25 ± 0.885.98 ± 0.930.9480.2290.313

Means within the same row with different superscripts are significantly different (P<0.05).

The effect of Lentinus edodes chaff on DM, OM, EE, CP, ADF and NDF is shown in Table 4. Analysis of variance showed that apparent digestibility of DM, CP, and NDF differed significantly among treatments. Compared with the control group, sheep fed 10% or 20% Lentinus edodes chaff had significantly higher apparent digestibility of DM, CP, and NDF, whereas the 30% group was significantly lower. Orthogonal polynomial contrasts further indicated significant linear (p = 0.028) and quadratic (p = 0.001) effects for DM; significant linear (p = 0.024), quadratic (p = 0.001), and cubic (p = 0.023) effects for CP; and a significant quadratic effect (p = 0.001) for NDF. The apparent digestibility of OM, EE and ADF was shown to be the same across the four groups.

Table 4. Effects of Lentinus edodes chaff on apparent digestibility of nutrients in Altay sheep.

ItemsLevels %p-value
0102030LinearQuadraticCubic
Dry matter (%)63.59 ± 4.33ABa67.27 ± 2.12Aa68.71 ± 4.93Aa57.01 ± 2.68Bb0.0280.0010.203
Organic matter (%)61.92 ± 4.1663.42 ± 6.3462.47 ± 6.8957.89 ± 4.080.2270.1600.765
Ether extract (%)74.45 ± 2.1373.44 ± 4.3475.40 ± 2.7172.38 ± 4.240.5140.4880.226
Crude protein (%)66.41 ± 2.34Ab68.46 ± 2.97Aab71.53 ± 2.95Aa60.64 ± 4.12Bc0.0240.0010.023
Neutral detergent fiber (%)44.75 ± 7.94Bb55.82 ± 4.35Aa56.42 ± 6.09Aa45.41 ± 4.19Bb0.8110.0010.918
Acid detergent fiber (%)34.83 ± 6.2739.49 ± 6.8042.10 ± 6.1440.00 ± 5.140.1200.1900.815

A, BMeans within the same row with different superscripts are extremely significantly different (P<0.01); a, bMeans within the same row with different superscripts are significantly different (P<0.05).

3.2 Rumen fermentation

Table 5 displays the findings for Lentinus edodes chaff used in meals, along with its impact on pH, NH3-N, total volatile fatty acids (VFA), acetate, propionate, isobutyrate, butyrate, isovalerate, valerate, and acetate/propionate. The results of the ANOVA revealed that there were no statistically significant differences in ruminal pH or ammonia nitrogen concentration between the experimental groups of fattening sheep. The proportions of various individual volatile acids, such as propionate, acetate, isobutyrate, valerate, isovalerate and acetate/propionate, remained unaltered by the chaff. Nevertheless, the percentage of butyrate in the 30% group was significantly higher than in the 10% group (p < 0.05). Total volatile fatty acid content was significantly increased in the 20% group compared with the control (p < 0.01), with a significant quadratic effect (P = 0.026) across the inclusion levels.

Table 5. Effects of Lentinus edodes chaff on rumen fermentation parameters in Altay sheep.

ItemsLevels %p-value
0102030LinearQuadraticCubic
pH6.93 ± 0.536.82 ± 0.367.00 ± 0.496.91 ± 0.300.8960.9600.545
NH3-N (mg/dL)12.56 ± 2.4812.89 ± 3.3514.37 ± 2.9912.03 ± 3.110.3400.7730.645
TVFA (mmol/L)56.17 ± 1.52Bb59.39 ± 3.89ABb65.58 ± 3.37Aa58.74 ± 7.32ABb0.1710.0260.081
Molar percentages of individual volatile fatty acids (%)
Acetate63.60 ± 1.6465.02 ± 0.9364.38 ± 0.7564.32 ± 1.790.5930.2280.305
Propionate20.77 ± 1.8420.72 ± 1.8620.18 ± 0.8619.68 ± 1.990.3210.7760.872
Iso-butyrate0.56 ± 0.100.58 ± 0.240.48 ± 0.030.43 ± 0.050.1500.5970.571
Butyrate13.54 ± 0.80ab12.10 ± 1.77b13.61 ± 0.69ab14.42 ± 0.61a0.1300.0570.131
Iso-valerate0.65 ± 0.140.69 ± 0.380.50 ± 0.050.44 ± 0.030.1270.6760.435
Valerate0.88 ± 0.220.89 ± 0.170.85 ± 0.080.71 ± 0.220.1700.3840.917
A/P3.08 ± 0.333.16 ± 0.323.20 ± 0.163.30 ± 0.460.3510.9230.858

A, BMeans within the same row with different superscripts are extremely significantly different (P<0.01); a, bMeans within the same row with different superscripts are significantly different (P<0.05).

NH3-N, ammonia nitrogen; TVFA, total volatile fatty acids; A/P, acetic acid/propionic acid.

3.3 Serum biochemical

As shown in Table 6, the serum protein metabolism indexes of the Altay sheep revealed that the Lentinus edodes chaff had no significant impact on the serum concentrations of TP, ALB, or GLB (p > 0.05). However, serum UREA concentration in the 30% group was significantly lower than that in the control group (p < 0.05), with a significant linear decrease (p = 0.018) as Lentinus edodes chaff inclusion level increased.

Table 6. Effects of Lentinus edodes chaff on serum protein metabolism indexes in Altay sheep.

ItemsLevels %p-value
0102030LinearQuadraticCubic
TP(g/L)58.20 ± 7.8555.47 ± 9.5156.65 ± 5.6557.19 ± 2.010.8850.5660.721
ALB(g/L)25.35 ± 2.4124.28 ± 2.6725.98 ± 2.3226.51 ± 1.810.2390.4070.364
GLB(g/L)34.88 ± 5.2231.10 ± 1.5932.38 ± 3.0532.10 ± 2.690.3700.3330.420
UREA(mmol/L)6.75 ± 0.91a6.57 ± 0.30ab6.24 ± 0.85ab5.74 ± 0.52b0.0180.5960.997

a, bMeans within the same row with different superscripts are significantly different (P<0. 05).

TP, total protein; ALB, albumin; GLB, globulin; UREA, urea.

As shown in Table 7, the addition of Lentinus edodes chaff to the sheep’s feed has an impact on the serum energy metabolism indicators of Altay sheep. TC differed significantly among treatments by ANOVA, although pairwise comparisons did not detect significant differences between individual groups. Orthogonal polynomial contrast analysis revealed a significant linear trend (p = 0.044) as Lentinus edodes chaff inclusion increased. TC showed no significant treatment effects (p > 0.05). Serum HDL concentration exhibited a significant linear increase (p = 0.019); the 30% group was numerically higher than the control. LDL levels showed a significant linear increase (p = 0.010), with the 20% and 30% groups being significantly higher than the control (p < 0.01 and p < 0.05, respectively). GLU levels in the 10% group were significantly lower than those in the control and 20% groups (p < 0.05), with a significant cubic effect detected across treatments (p = 0.008).

Table 7. Effects of Lentinus edodes chaff on serum energy metabolism indexes of sheep.

ItemsLevels %p-value
0102030LinearQuadraticCubic
TC (mmol/L)0.95 ± 0.250.96 ± 0.141.16 ± 0.121.13 ± 0.180.0440.8380.237
TG (mmol/L)0.21 ± 0.030.21 ± 0.070.20 ± 0.050.19 ± 0.040.4280.8490.880
HDL(mmol/L)0.61 ± 0.11b0.63 ± 0.13ab0.69 ± 0.11ab0.76 ± 0.09a0.0190.5680.904
LDL (mmol/L)0.40 ± 0.22Bb0.59 ± 0.12ABab0.73 ± 0.16Aa0.65 ± 0.16ABa0.0100.0610.540
GLU(mmol/L)3.39 ± 0.67a2.63 ± 0.26b3.31 ± 0.49a3.00 ± 0.23ab0.5410.2460.008

A, BMeans within the same row with different superscripts are extremely significantly different (P<0.01); a, bMeans within the same row with different superscripts are significantly different (P<0.05).

TC, total cholesterol; TG, triglyceride; HDL, high-density lipoprotein; LDL, low-density lipoprotein; GLU, glucose.

The serum enzyme indexes of the tested sheep in Table 8 showed no differences in AST or ALP throughout the experiment (p > 0.05). Compared with the control group, serum ALT levels were significantly lower in the 10% and 30% groups (p < 0.05), with a significant cubic effect detected across treatments (p = 0.034). CK levels were also significantly lower in the 10% and 30% groups (p < 0.05), with a significant linear decrease observed (p = 0.044). In addition, LDH level was significantly higher in the 10% group than in the control (p < 0.01) and was also significantly elevated in the 20% group (p < 0.05), exhibiting a significant quadratic effect (p = 0.017).

Table 8. Effects of Lentinus edodes chaff on serum enzyme indexes of sheep.

ItemsLevels %p-value
0102030LinearQuadraticCubic
AST (U/L)76.36 ± 3.4868.49 ± 7.9175.54 ± 9.6576.04 ± 3.160.8120.2370.130
ALT (U/L)36.31 ± 2.99a29.41 ± 4.77b35.81 ± 4.96a35.87 ± 2.17a0.5190.0720.034
LDH (U/L)468.61 ± 33.73Aa387.16 ± 41.57Bb408.45 ± 61.63ABb424.17 ± 23.83ABab0.1720.0170.225
CK (U/L)218.16 ± 43.55a164.31 ± 18.00b182.23 ± 29.47ab169.43 ± 11.57b0.0440.1450.114
ALP (U/L)234.45 ± 76.88269.23 ± 76.09279.78 ± 92.10304.51 ± 10.380.1160.8730.794

A, BMeans within the same row with different superscripts are extremely significantly different (P<0.01); a, bMeans within the same row with different superscripts are significantly different (P<0.05).

AST, aspertate aminotransferase; ALT, alanine aminotransferase; LDH, lactate dehydrogenase; CK, creatine kinase; ALP, alkaline phosphatase.

Table 9 shows the serum immune index parameters. In contrast to the control group, Lentinus edodes chaff had no significant impact on serum IgM content in sheep (p < 0.05). IgG and IgA levels were significantly higher in the 10% group than in the control group (p < 0.05) and extremely significantly higher in the 30% group (p < 0.01). Additionally, serum IgG and IgA concentrations in the 20% group were significantly higher than those of the control (p < 0.01 and p < 0.05, respectively), both exhibiting significant linear increases (p = 0.001). IL-2 levels in all three test groups were significantly lower than those in the control (p < 0.01), accompanied by a significant linear decrease (p = 0.001).

Table 9. Effects of Lentinus edodes chaff on serum immune indexes of sheep.

ItemsLevels %p-value
0102030LinearQuadraticCubic
IgG (μg/mL)16.99 ± 0.65Bb18.70 ± 0.83ABa19.43 ± 1.98Aa20.14 ± 0.74Aa0.0010.3130.652
IgM (μg/mL)1.08 ± 0.081.13 ± 0.141.20 ± 0.311.23 ± 0.100.1300.9440.871
IgA (μg/mL)0.52 ± 0.18Bb0.67 ± 0.08ABa0.66 ± 0.02ABa0.74 ± 0.02Aa0.0010.3410.175
IL-2 (pg/mL)321.88 ± 9.62Aa285.46 ± 8.07Bb261.93 ± 15.58Cc241.88 ± 7.73Dd0.0010.0760.635

A, BMeans within the same row with different superscripts are extremely significantly different (P<0.01); a, bMeans within the same row with different superscripts are significantly different (P<0.05).

IgG, Immunoglobulin G; IgM, Immunoglobulin M; IgA, Immunoglobulin A; IL-2, Interleukin-2.

As shown in Table 10, analysis of variance revealed significant differences in serum antioxidant indexes (T-AOC, SOD, GSH-Px, MDA and NO) between treatment groups. Levels in the sheep fed 10%, 20% or 30% Lentinus edodes chaff were significantly higher than in the control group. The T-AOC content in the 30% group increased significantly (p < 0.01) when compared with the control group, as did the content in the 20% group (p < 0.05). There was a significant increase in SOD content in the 20% and 30% groups (p < 0.05), as well as in the GSH-Px content of the 20% and 30% groups (p < 0.01). The 10% group showed a significant increase in GSH-Px content (p < 0.05), as did the NO content in the 10%, 20% and 30% groups (p < 0.01). The MDA levels in the three groups that were put through the experiment decreased significantly (p < 0.01). Orthogonal polynomial contrast analysis revealed significant linear effects for all these parameters (p = 0.001).

Table 10. Effects of Lentinus edodes chaff on serum antioxidant indexes of sheep.

ItemsLevels %p-value
0102030LinearQuadraticCubic
T-AOC (U/mL)6.63 ± 0.83Bc8.13 ± 1.04ABbc9.24 ± 2.80ABab10.49 ± 0.84Aa0.0010.8470.860
SOD (U/mL)59.27 ± 6.91b66.5 ± 12.05ab72.74 ± 11.28a74.88 ± 4.61a0.0050.5010.867
GSH-Px (U/mL)226.58 ± 9.097Bc255.0 ± 9.52ABb275.1 ± 30.69Aab281.6 ± 25.41Aa0.0010.2140.893
MDA (nmol/mL)5.07 ± 0.27Aa4.59 ± 0.35Bb4.12 ± 0.27Cc3.56 ± 0.19Dd0.0010.7090841
NO(umol/L)33.99 ± 6.68Cc42.21 ± 3.27Bb47.88 ± 6.34ABab53.77 ± 2.12Aa0.0010.5720.762

A, BMeans within the same row with different superscripts are extremely significantly different (P<0.01); a, bMeans within the same row with different superscripts are significantly different (P<0.05).

T-AOC, Total antioxidant capacity; SOD, Superoxide dismutase; GSH-Px, Glutathione peroxidase; MDA, Malondialdehyde; NO, Nitric oxide.

4 Discussion

In the breeding process, animal growth performance has a significant impact on economic benefits, so ADG (average daily gain) and FI (feed intake) are key research indicators. The ingestion of feed by sheep can be influenced by both physiological factors and external factors. The factors under discussion include dietary habits, lifestyle behaviours, physiological factors, environmental factors and animal learning abilities (Yuan et al., 2022). It was found by Xin et al. (2025) that the ADG of Yanbian cattle fed with 50% fermented SMS from Pleurotus eryngii was increased by 25.74%. The experiment revealed no major differences in the FBW and FGR of the sheep in each group. However, the 20% group showed significantly higher BWG, ADG and Average Daily FI compared to the control group. Consequently, it can be inferred that the utilisation of Lentinus edodes chaff has a favourable impact on the growth and development of ovines. Collectively, these results indicate that dietary supplementation with mushroom bran, particularly at the 20% level, may enhance feed efficiency in Altay rams. The observed improvement in ADG, which exceeded the proportional increase in ME intake, further supports the notion that Altay sheep possess a breed-specific metabolic advantage in converting dietary energy into body tissue. This utilisation has been shown to enhance feed utilisation and concomitantly reduce breeding costs.

As proposed by (Yang et al., 2021b), a positive correlation exists between the digestibility of feed and the absorption rate of nutrients by animals. This, in turn, affects the quality of feed, both of which are vital to the growth and development of ruminants. The study suggests that the biodegradation of straw with Pleurotus ostreatus enhances its nutritional value and digestibility in ruminant diets (Yuan et al., 2022). In comparison with the control, the Lentinus edodes chaff exhibited no significant impact on the apparent digestibility of OM, EE and ADF. Concurrently, the incorporation of 20% SMS into the diets of Altay male sheep resulted in enhanced apparent digestibility of CP and NDF. The observed phenomenon may be attributable to the nutritional qualities and distinct composition of the feed raw materials (i.e. bacterial residue), which have been demonstrated to directly enhance apparent digestibility (Pin and Anusorn, 2018). However, the findings of this study indicated that replacing 30% of the feed with SMS resulted in a reduction in digestibility of DM. This phenomenon may be attributable to the DM content of the SMS in comparison with the mixed concentrate feed, and it is evident that further research is required in order to ascertain the underlying causes. Furthermore, the administration of 30% SMS led to a substantial decline in apparent digestibility of CP. This decline may be attributed to the elevated fibre content of SMS, which has the capacity to prolong the feed retention time in the digestive tract and reduce nutrient digestibility. Huang et al. (2023) also ascertained that the use of SMS results in a reduction of the digestibility of DM, CP and fibre in sheep. In summary, the present study has demonstrated that SMS can enhance the growth performance of sheep, thereby generating economic benefits. However, it is crucial to acknowledge that as the proportion of SMS substituted rises, the apparent digestibility of sheep may decline.

It is an established phenomenon that fluctuations in the pH value of the rumen are a reliable indication of the fermentation process and the overall stability of the internal environment. The normal range is 6.0-7.0 (Guo et al., 2022b). The pH values of the rumen samples obtained from both groups were found to be within the normal range. The level of NH3-N is dependent on the rates at which nitrogenous substances in the diet are degraded, and on the rate at which NH3-N is utilised by microorganisms and rumen epithelial cells (Liang et al., 2024). Increasing SMS addition to feed was found to provide NH3 concentrations that sustained rumen microorganisms, enabling better feed utilisation, but at optimal levels. The findings of the present study demonstrated a high degree of similarity with those results that had been previously documented by Pin and Anusorn (2018). It is evident that the rumen of ruminants possesses the capacity to degrade dietary nutrients and to produce acetic and propionic acids. It has been demonstrated that the process of propionic acid conversion to glucose is more efficient than that of acetic acid. It has been demonstrated that elevated levels of propionic acid can enhance an organism’s capacity to utilise energy effectively, thereby promoting growth (Zhao et al., 2025). It is frequently the case that the rumen fermentation pattern is characterised by the A:P ratio. It has been demonstrated that an increase in the A:P ratio results in a shift towards the production of acetic acid during the fermentation process (Li et al., 2025, 2024). The present study examined the effects of dietary SMS on A:P concentrations and the isobutyrate ratio. The findings of the study demonstrated that the incorporation of dietary SMS at a proportion of 20% into the sheep’ diets led to a 7.1% increase in A:P concentrations and a concurrent reduction of the isobutyrate ratio by 23.2% (from 0.56 to 0.43) compared to the control. Although this finding suggests a shift in the rumen fermentation pattern towards an acetate-type fermentation, it is important to note that the observed changes in VFA concentrations were relatively modest and may not fully account for the substantial improvement in ADG. Therefore, the growth-promoting effect of SMS supplementation may also involve mechanisms beyond rumen fermentation, such as the bioactive compounds present in cottonseed hull mushroom bran, which may exert direct effects on intestinal health, nutrient absorption, or systemic metabolic regulation. These observations align with the findings of prior research (Pin and Anusorn, 2018; Fruge et al., 2025).

It is evident that ALB and TP play pivotal regulatory roles within the bloodstream. These include the binding of hormones, fatty acids and bilirubin, the modulation of vascular permeability and the maintenance of osmotic pressure (Narinx et al., 2022). In ruminants, fluctuations in blood UREA levels can be influenced by the quantity of dietary protein, the level of feed intake, and the degradability of rumen protein (Yuan et al., 2022b). However, the present study demonstrated a progressive decline in serum UREA levels in response to SMS. The potential for mediation by SMS modulation of gastrointestinal microbial balance and improved hepatic function is a hypothesis that merits further investigation (Chen et al., 2023). In a series of studies, Wang et al. (2023) reported a consistent finding that the dietary inclusion of fermented sawdust mushroom bran resulted in a reduction of serum UREA concentrations in sheep. Moreover, no substantial disparities were detected in serum TP and ALB concentrations across the treatment groups, indicating that SMS substitution did not exert a deleterious effect on protein synthesis or metabolism in sheep.

The GLU is employed as a marker of the energy status in ruminants, a factor which is of significance for the health and productivity of these animals. In the present study, the substitution of 10% of Altay male sheep with SMS resulted in a reduced glucose concentration, likely reflecting an enhanced flux of glucose into anabolic pathways that support protein synthesis and lipid deposition. Elevated levels of HDL have been shown to be associated with a reduced risk of mortality from atherosclerosis (Mackness and Mackness, 2010; Mineo et al., 2006). In this study, serum concentrations of HDL and LDL were elevated in both experimental groups following SMS supplementation, in comparison with the control. It has been hypothesised that SMS exerts immunostimulatory properties, a phenomenon that is probably attributable to bioactive substances generated during the fermentation process in the rumen (Wang, 2020; Fan et al., 2021).

ALT and AST have been identified as significant reference indicators for evaluating hepatic function. AST and ALT activity rises rapidly when liver damage occurs or protein metabolism increases (Yang et al., 2022; Rotaru et al., 2025). As demonstrated in previous studies, the administration of SMS has been shown to reduce serum ALT and AST levels in sheep (Feng et al., 2025; Long et al., 2024). The present study established that the provision of 10% SMS diets resulted in a decline in ALT content. This finding indicates that the provision of 10% SMS diets has the potential to enhance the immune function of Altay male sheep and promote liver health.

IgG is vital to the immune system and is the main antibody present in the blood after food is digested. In comparison, IgA’s primary function is to neutralise pathogenic bacteria by interacting with non-specific innate defence mechanisms (Liu et al., 2015). The present investigation established that providing sheep with diets consisting of 20% and 30% SMS increased the concentrations of IgA and IgG in their serum. This suggests a potential enhancement of humoral immune capacity. However, these changes were relatively modest and alone are unlikely to fully explain the substantial improvement in ADG. The growth-promoting effect of SMS is therefore likely attributed to multiple mechanisms, rather than immune enhancement alone.

The detrimental effects of oxidative stress on animal growth and production highlight the importance of increasing the body’s antioxidant capacity. This approach is instrumental in ensuring the safe production of animals and maintaining their overall health. A plethora of significant indicators, encompassing GSH-Px, T-AOC, SOD, NO and MDA, are frequently employed to assess an animal’s antioxidant stress resistance (Wang et al., 2023). In this study, the concentrations of GSH-Px, T-AOC, NO and SOD increased in proportion to the rise in the proportion of SMS in the diet, while the opposite trend was exhibited by MDA levels. This finding indicates that the provision of SMS to sheep may enhance their antioxidant capacity and promote optimal growth.

In summary, Altay sheep exhibited a significant enhancement in growth performance, nutrient digestibility, and serum immunity, as well as antioxidant capacity, when their intensive feed was supplemented with SMS. The results of this study show that a daily dietary supplement containing 20% SMS effectively promotes the growth of male Altay sheep. The same regime promotes Altay male sheep development and immune function. Consequently, further refinement of SMS dosages in subsequent research endeavours is anticipated to yield even more pronounced beneficial effects.

5 Conclusion

The findings of the present study corroborated the viability of SMS as a feed ingredient in Altay male sheep diets, a finding that is of significance to the research community. When administered at 20% DM, SMS was found to enhance average daily gain and reduce the feed-to-gain ratio. The 20% and 30% treatments demonstrated increased immune function and antioxidant capacity. Based on a comprehensive evaluation, the optimal supplementation level was 20%. From a practical perspective, SMS supplementation at 20% DM reduces feeding costs, alleviates human-livestock feed competition, and promotes sustainable animal husbandry by recycling agricultural by-products.

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Footnotes

Edited by: Soumaya Boukrouh, Mohammed VI Polytechnic University, Morocco

Reviewed by: Fanfan Zhang, Shihezi University, China

Cuiling Wu, Xinjiang Normal University, China

Statements

Conflict of interest

The author(s) declared that this work was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Funding

The author(s) declared that financial support was received for this work and/or its publication. This work was financially supported by the Xinjiang Uygur Autonomous Region Key Research and Development Project of China (2022B02029), and the National Key Research and Development Program of China (2021YFD1600701-1), and Fund of the Ministry of Finance and the Ministry of Agriculture and Rural Ministry-National Modern Agricultural Industry Technology System (CARS-38).

Data availability

The raw data supporting the conclusions of this article will be made available by the authors, without undue reservation.

Ethics approval

The animal studies were approved by Xinjiang Animal Welfare and Ethics Committee of the Academy of Animal Sciences in China. The studies were conducted in accordance with the local legislation and institutional requirements. Written informed consent was obtained from the owners for the participation of their animals in this study.

Author contributions

LW: Data curation, Writing – review & editing, Methodology, Software, Formal analysis, Conceptualization, Writing – original draft. ST: Conceptualization, Writing – original draft, Writing – review & editing, Data curation. RQ: Validation, Writing – original draft, Writing – review & editing, Methodology, Conceptualization. AM: Conceptualization, Writing – review & editing, Writing – original draft, Investigation. YW: Software, Writing – review & editing, Writing – original draft, Data curation. XC: Writing – review & editing, Investigation, Writing – original draft, Conceptualization. YL: Formal analysis, Writing – review & editing, Data curation, Conceptualization, Writing – original draft, Methodology. WW: Writing – review & editing, Conceptualization, Writing – original draft, Validation, Data curation.

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Citation

Wang, L., et al. (2026). Substitution with Lentinus edodes chaff enhances the health and growth performance of Altay sheep, and improvement of serum immune function and oxidative resistance. Frontiers in Animal Science, Volume 7. https://doi.org/
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7 - 2026

Copyright

Copyright © 2026 Wang, Tang, Qin, Mubalake, Wang, Chen, Liu and Wang.

License: https://creativecommons.org/licenses/by/4.0/

*Correspondence: Yanfeng Liu; Wenqi Wang

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