Animal Feed Science and Technology 299 (2023) 115622

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Animal Feed Science and Technology

journal homepage: www.elsevier.com/locate/anifeedsci

Effects of substitution of millet straw for corn silage and alfalfa

hay on lactation performance, ruminal fermentation, and blood
metabolites in late-lactation Holstein dairy cows
Meimei Wang a, c, 1, Yan Li b, c, 1, Jianfang Feng a, c, 1, Yizhao Shen a, c, Yufeng Cao a, c,
Qiufeng Li a, c, Yanxia Gao a, c, *, Jianguo Li a, c, *
a
College of Animal Science and Technology, Hebei Agricultural University, Baoding, Hebei 071000, People’s Republic of China
b
College of Veterinary Medicine, Hebei Agricultural University, Baoding, Hebei 071000, People’s Republic of China
c
Key Laboratory of Healthy Breeding in Dairy Cattle (Co-construction by Ministry and Province), Ministry of Agriculture and Rural Affairs, Baoding,
Hebei 071000, People’s Republic of China

A R T I C L E I N F O A B S T R A C T

Keywords: The study was conducted to evaluate the effect of replacing dietary corn silage (CS) and alfalfa
Byproduct hay (AH) with millet straw (MS) on the production performance and the ruminal fermentation
Dairy cows patterns of Holstein cows in late lactation. For this purpose, forty-eight multiparous Holstein
Millet straw
dairy cows blocked by milk yield (32 ± 3.2 kg/d) and days in milk (179 ± 19 d) were randomly
Nutrient digestibility
allocated to one of four treatments. The diets were formulated with a forage-to-concentrate ratio
of 48:52 [dry matter (DM) basis] and contained the same concentrate mixtures, with different
forage sources (on a DM basis): (1) Control (60% CS, 40% AH, and 0% MS; on 100% roughage);
(2) LMS (54% CS, 36% AH, and 10% MS), (3) MMS (48% CS, 32% AH, and 20% MS), and (4)
HMS (42% CS, 28% AH, and 30% MS). The dry matter intake and milk yield were lower in the
HMS diet than those in the Control, LMS, and MMS diets (P < 0.01), whereas there was no dif­
ference in the Control, LMS, and MMS diets. Compared with the Control diet, 4% FCM, milk
protein yield, and N conversion ratio decreased in the MMS and HMS diets (P < 0.01), whereas no
difference between the Control and LMS diets was found. Besides, cows fed the HMS diet had
lower milk fat yield, lactose percentage, and feed efficiency than those in the Control and LMS
diets (P < 0.05), with no difference between the Control, LMS, and MMS diets. The percentage of
milk fat and protein was not affected by the diets. Compared with the Control diet, the total VFA
concentration was decreased, whereas the ammonia-N concentration was increased in the HMS
diet (P < 0.05), with no difference between the Control, LMS, and MMS diets. The molar pro­
portion of acetate and the ratio of acetate to propionate were lower (P < 0.01), while the molar
proportion of propionate was higher in the Control diet than that in the MMS and HMS diets (P <

Abbreviations: ADL, acid detergent lignin; AH, alfalfa hay; ALT, alanine aminotransferase; AST, aspartate aminotransferase; BHBA, β-hydroxy­
butyric acid; BUN, blood urea nitrogen; CP, crude protein; CS, corn silage; DM, dry matter; DMI, dry matter intake; FCM, fat corrected milk; HMS, a
diet containing 42% corn silage, 28% alfalfa hay, and 30% millet straw; LMS, a diet containing 54% corn silage, 36% alfalfa hay, and 10% millet
straw; MS, millet straw; MMS, a diet containing 48% corn silage, 32% alfalfa hay, and 20% millet straw; NH3-N, ammonia-nitrogen; TVFA, total
volatile fatty acid; VFA, volatile fatty acid.
* Corresponding authors at: College of Animal Science and Technology, Hebei Agricultural University, Baoding, Hebei 071000, People’s Republic
of China.
E-mail addresses: yanxia.gao@hebau.edu.cn (Y. Gao), jgli@hebau.edu.cn (J. Li).
1
These authors contributed equally to this work.

https://doi.org/10.1016/j.anifeedsci.2023.115622
Received 22 November 2022; Received in revised form 19 February 2023; Accepted 22 February 2023
Available online 24 February 2023
0377-8401/© 2023 Elsevier B.V. All rights reserved.
M. Wang et al. Animal Feed Science and Technology 299 (2023) 115622

0.01), with no difference between the LMS and MMS diets. Compared with the Control, the
apparent digestibility of DM, CP, NDF, and ADF was decreased, while the blood urea nitrogen
concentration was increased in the HMS diet (P < 0.05), with no differences in the Control, LMS,
and MMS diets. Thus, it was concluded that it is feasible to use MS as a fiber source for late-
lactation dairy cows. However, considering the milk production performance and the feed di­
gestibility, the replacement of CS and AH with MS should not exceed 20% of the roughage.

1. Introduction

Corn silage (CS) and alfalfa hay (AH) are commonly used roughage resources for dairy cows due to their high digestibility, which
helps to maintain the intake and milk production (Ferraretto et al., 2015; Fustini et al., 2017), but the supply of these high-quality
forages is sometimes limited by adverse weather, lack of land or water, or price issues. Meanwhile, numerous crop byproducts are
produced annually worldwide, most of which are burned, resulting in environmental issues (Seglah et al., 2020; Wang et al., 2021).
Crop residues like straws could provide a certain amount of protein and energy for ruminants, thus, could be used in ruminants’ diets
(Coelho da Silva et al., 2019; Adugna et al., 2020). Using these crop residues as feed can not only reduce environmental pollution but
also alleviate feed shortage. However, the low digestibility of most crop residues, like wheat straw or rice straw, limits their utilization.
Millet straw (MS) is the byproduct of the finger millet crop, which is widely grown in arid and semi-arid areas of the world (Sood
et al., 2019). Compared with that in wheat straw or rice straw, the higher DM digestibility indicated that MS should be an optimal feed
source for ruminants (Wilman et al., 1999). However, previous studies found that the utilization of MS was limited due to its

Table 1
Ingredients and chemical composition of the experimental diets.
Items Dietsa

Control LMS MMS HMS

Ingredient, % of DM
Corn silage 28.6 25.8 22.9 20.1
Alfalfa hay 19.1 17.2 15.3 13.4
Millet straw 0 4.77 9.55 14.3
Cracked corn 5.56 5.56 5.56 5.56
Steam-flaked corn 13.9 13.9 13.9 13.9
Cottonseed meal 1.01 1.01 1.01 1.01
Soybean meal 9.22 9.22 9.22 9.22
Extruded soybean 1.84 1.84 1.84 1.84
Wheat bran 2.00 2.00 2.00 2.00
Soybean hull 2.00 2.00 2.00 2.00
Whole cottonseed 6.45 6.45 6.45 6.45
Rapeseed meal 0.68 0.68 0.68 0.68
Corn DDGSb 1.73 1.73 1.73 1.73
Beet grain 4.09 4.09 4.09 4.09
Dicalcium phosphate 0.69 0.69 0.69 0.69
MgO 0.19 0.19 0.19 0.19
Limestone 0.39 0.39 0.39 0.39
NaCl 0.22 0.22 0.22 0.22
NaHCO3 0.61 0.61 0.61 0.61
Rumen protected methionine 0.14 0.14 0.14 0.14
Fat powder 0.73 0.73 0.73 0.73
Premixc 0.86 0.86 0.86 0.86
Total 100 100 100 100
Nutrition levelsd
NEL, MJ/Kg 6.70 6.68 6.65 6.62
CP, % 16.5 16.1 15.8 15.4
NFC, % 40.7 39.6 38.6 37.6
NDF, % 31.8 33.3 34.8 36.3
ADF, % 20.1 20.9 21.8 22.6
Ca, % 0.83 0.83 0.82 0.82
P, % 0.41 0.41 0.40 0.39
a
Control = 60% corn silage, 40% alfalfa hay, and 0% millet straw (on 100% roughage); LMS = 54% corn silage + 36% alfalfa hay + 10% millet
straw; MMS = 48% corn silage + 32% alfalfa hay + 20% millet straw; HMS = 42% corn silage + 28% alfalfa hay + 30% millet straw.
b
DDGS = distillers dried grains with solubles.
c
Premix contained (per kg of DM): 700,000 IU of vitamin A; 250,000 IU of vitamin D3; 10,000 IU of VE; 920 mg of vitamin B1; 10 mg of vitamin B12;
100 mg of D-Biotin; 270 mg of D-pantothenic acid; niacinamide of 30,000 mg; 1860 mg of Fe; 1700 mg of Cu; 4030 mg of Mn; 7512 mg of Zn; 62 mg of
Co; 79,200 mg of Mg; 52,000 mg of K.
d
NEL, data calculated from NRC (2001); CP, crude protein; NFC, non-fiber carbohydrates; NDF, neutral detergent fiber; ADF, acid detergent fiber;
Ca, calcium; P, phosphorus

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M. Wang et al. Animal Feed Science and Technology 299 (2023) 115622

unbalanced nutrient profile (Darshan et al., 2007; Alemu et al., 2020). Moreover, other studies demonstrated that MS, as sole forage,
might compromise DMI, leading to a poor production performance of animals (Chandrasekharaiah et al., 2011a, 2011b; Kumar et al.,
2021). Therefore, it is necessary to explore an optimal utilization method of MS.
Kahyani et al. (2019) reported no treatment effect on DMI and milk yield when substitution wheat straw for CS and AH. Gowda and
Prasad (2005) found that cows fed with MS exhibited higher DMI, milk yield, and nutrient digestibility than those fed with rice straw.
Considering the higher nutrient digestibility, better nitrogen and cell-soluble nutrients of MS than wheat straw and rice straw (Wilman
et al., 1999; Gowda and Prasad, 2005), the substitution of MS for CS and AH should be optimal for MS utilization in dairy cows.
Therefore, we hypothesized that MS substituted for CS and AH would not compromise DMI and milk production of dairy cows. The
objectives of this study were to investigate the effects of the substitution of MS for CS and AH on lactation performance, ruminal
fermentation, and blood metabolites of dairy cows, and to explore the optimal substitution ratio.

2. Materials and methods

This study was carried out at Hongda Animal Husbandry Co., Ltd. (Baoding, China) from March 2021 to May 2021. The experi­
mental protocol (JGL 1718) was approved by the Institutional Animal Care and Use Committee of Hebei Agricultural University
(Baoding, China).

2.1. Cows, diets, and experimental design

Forty-eight multiparous late-lactation Holstein dairy cows (parity = 2.8 ± 0.83, milk production = 32 ± 3.2 kg/d, days in milk =
179 ± 19 d; mean ± SD) were blocked by milk production, day in milk, and parity, and randomly allocated to one of four dietary
treatments (Table 1). Diets were formulated with a forage-to-concentrate ratio of 48:52 (DM basis) and contained the same concentrate
mixtures, with different forage sources (on a DM basis): (1) Control, a diet containing 60% CS, 40% AH, and 0% MS (100% roughage);
(2) LMS, a diet containing 54% CS, 36% AH, and 10% MS; (3) MMS, a diet containing 48% CS, 32% AH, and 20% MS; (4) HMS, a diet
containing 42% CS, 28% AH, and 30% MS. The chemical composition of MS, CS and AH is listed in Table 2. The diets were formulated
according to the NRC (2001) and were offered for cows as TMR. The experiment lasted for 63 days, with 7 days for adaption.
Dairy cows were housed individually in tie stalls (1.5 m × 2.8 m) with automatic water troughs and fed three times daily at 0630,
1330, and 2030 h with ad libitum feeding (ensuring at least 5% refusals). Orts were collected and weighed daily. The cows had free
access to drinking water, and they were milked three times daily at 0530, 1230, and 1930 h.

2.2. Sampling and data collection

The diets offered and refused by each cow were recorded daily. Feed offered and orts were collected daily, pooled weekly, and
mixed to present a representative sample for chemical composition analysis. The TMR and orts were dried at 55 ℃ in the oven for 48 h
according to Beauchemin and Yang (2005) and Stahl et al. (2020), then ground through a 1-mm screen (stand model 4 Wiley Mill,
Arthur H. Thomas, Philadelphia, PA) for further analysis. The daily DMI was calculated as the difference between DM offered and DM
refused. The DMI data were averaged weekly for statistical analysis.
Milk yields were recorded daily at each milking and averaged per week for data analysis. Milk was sampled weekly at each milking
time from each cow on d 42, 49, and 56 during the sampling period. In each period, about 60 mL- aliquot of milk were pooled and
mixed based on the milk production by volume, further mixed with 2-bromo-2-nitropropane-1,3-diol, and stored (− 20 ℃) until milk
composition analysis. The milk composition (milk fat, milk protein, milk lactose, and nonfat solids) was determined by a near-infrared
milk analyzer (Foss-4000, Foss Electric A/S, Hillerød, Denmark; AOAC International, 2006; method 972.16). The milk component
yield was calculated by multiplying the measured concentration of a specific milk component by the corresponding daily milk yield.
Six cows from each group (blocked by milk production and day in milk) were randomly selected for feces, blood, and ruminal fluid
sample collection. The fecal samples were collected directly from the rectum every 6 h on d 50 to d 53, to represent a 24-h period of
each digestibility period. And then, the samples were pooled and mixed for each cow and stored at − 20 ℃ until analysis. After being

Table 2
Chemical composition of the roughage sources.
Itema, % of DM Corn silage Alfalfa hay Millet straw

DM 32.8 94.5 93.5

OM 95.2 90.9 94.7
CP 8.0 14.8 4.7
NDF 37.1 50.0 74.7
ADF 20.9 39.9 46.4
ADL 3.13 7.82 8.25
EE 3.4 1.7 1.2
Ash 4.8 7.9 5.3
NFC 46.7 25.6 14.1
a
DM, dry matter; OM, organic matter; CP, crude protein; NDF, neutral detergent fiber; ADF, acid detergent fiber; EE, ether
extract; ADL, acid detergent lignin (ADL); NFC (non-fiber carbohydrates) = 100% － (%NDF + %CP + %EE + %Ash).

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M. Wang et al. Animal Feed Science and Technology 299 (2023) 115622

dried in the oven at 55 ℃ for 72 h, they were ground through a 1-mm screen (stand model 4 Wiley Mill; Arthur H. Thomas) for further
chemical analysis.
About 40 mL of blood samples were collected from the jugular vein using gel vacuum tubes without any additives (Na heparin)
before morning feeding on d 42, 49, and 56 during the sampling period. Then the blood samples were centrifuged at 3000g (4 ℃) for
15 min to get the serum, the samples of which were transferred into 2 mL centrifuge tubes and stored at − 20 ℃ for the analysis of
blood urea nitrogen (BUN), albumin, total protein, cholesterol, triglyceride, alanine aminotransferase (ALT), aspartate aminotrans­
ferase (AST), β-hydroxybutyric acid (BHBA), and non-esterified fatty acid (NEFA).
On d 42, 49, and 56, the rumen fluid samples were collected with an oral stomach tube as described by Shen et al. (2012) before
morning feeding. The first 100 mL of rumen fluid was discarded to prevent the contamination of saliva. Then another 100 mL of rumen
fluid was collected and filtered through four layers of cheesecloth. Ruminal pH was determined immediately using a portable pH meter
(Starter 300; Ohaus Instruments Co. Ltd., Shanghai, China). After pH determination, five mL of samples were acidified with 1 mL of
25% (wt/vol) HPO3 and stored at − 20 ℃ for VFA analysis, and 5 mL was preserved with 1 mL 1% (wt/vol) H2SO4 for
ammonia-nitrogen (NH3-N) concentration analysis.

2.3. Sample analysis

The DM (method 934.1), crude protein (CP, method 990.03), ether extract (EE, method 2003.05), organic matter (OM, method
942.05), calcium (Ca, method 927.02), and phosphorus (P, method 965.17) content in TMR, feed refusals, and feces were determined
as outlined in AOAC (2006). Neutral detergent fiber (NDF) of the TMR, feed refusals, and feces were determined using heat-stable
α-amylase and sodium sulfite according to Van Soest et al. (1991), and the acid detergent fiber (ADF) was determined as outlined
in AOAC (2006). Both NDF and ADF were expressed including residual ash (Mertens et al., 2002). Acid detergent lignin (ADL) was
determined based on the AOAC (2006) method 973.18, and modified to use a 1.0 g of sample in Ankom F57 bags (Ankom Technology).
Non-fiber carbohydrate (NFC) was calculated as 100 – (CP + NDF + EE + ash). The concentration of ruminal NH3-N was determined
using a UV spectrophotometer (UV-2600, Unico, Shanghai, China) as described by Broderick and Kang (1980). Ruminal VFA con­
centration was measured using gas chromatography (Agilent Technologies 7890a system; a fused silica column, 30 m × 0.32 mm ×
0.25 mm; column temperature, 150 ℃; injector temperature, 200 ℃; and detector temperature, 250 ℃) as described by Shen et al.
(2019).
The concentrations of BUN, albumin, total protein, cholesterol, triglyceride, ALT, AST, BHBA, and NEFA were measured using
commercial kits (Nanjing Jiancheng Bioengineering Institute, Nanjing, China). The results showed that the intra-assay coefficients of
variation were lower than 9%, and the inter-assay coefficients of variation were lower than 15%.

2.4. Calculations and statistical analysis

The apparent total-tract digestibility of nutrients was determined with the ADIA as the internal marker (Stahl et al., 2020), and the
equation was as follows:

Table 3
Effects of substitution of millet straw for corn silage and alfalfa hay on DMI, milk production, and composition in late-lactating Holstein cows.
Item Treatment1 SEM P-value2

Control LMS MMS HMS T W T×W

DMI, kg/d 24.1a 23.9a 23.7a 22.8b 0.15 < 0.01 < 0.01 0.46
Yield, kg/d
Milk 29.2a 28.5a 28.3a 26.3b 0.71 0.04 < 0.01 0.53
4% FCM 28.4a 27.6ab 27.0bc 25.6c 0.47 < 0.01 0.05 0.61
Milk fat
% 3.79 3.84 3.82 3.84 0.071 0.95 0.93 0.99
kg/d 1.11a 1.08a 1.06ab 1.01b 0.023 0.03 0.15 0.77
Milk Protein
% 3.24 3.21 3.21 3.19 0.031 0.76 0.79 0.66
kg/d 0.95a 0.90ab 0.89bc 0.85c 0.019 < 0.01 0.03 0.45
Lactose
% 4.61a 4.56ab 4.55ab 4.47b 0.035 0.05 0.22 0.84
kg/d 1.35a 1.30ab 1.27b 1.18c 0.025 < 0.01 < 0.01 0.33
SNF3, % 8.60 8.62 8.63 8.69 0.050 0.63 0.52 0.36
Feed efficiency4 1.20a 1.18a 1.14ab 1.09b 0.019 < 0.01 0.16 0.20
N conversion5, % 25.1a 23.9ab 23.8bc 22.6c 0.50 0.01 0.03 0.40
a-c
Different lowercase letters within a row indicate significant differences.
1
Control = 60% corn silage, 40% alfalfa hay, and 0% millet straw (on 100% roughage); LMS = 54% corn silage + 36% alfalfa hay + 10% millet straw;
MMS = 48% corn silage + 32% alfalfa hay + 20% millet straw; HMS = 42% corn silage + 28% alfalfa hay + 30% millet straw.
2
T = treatment effect; W = week effect; T × W = the interaction between treatment and week.
3
SNF = non-fat solids.
4
Feed efficiency = 4% FCM/DMI.
5
N conversion = milk protein yield/CP intake.

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M. Wang et al. Animal Feed Science and Technology 299 (2023) 115622

Nutrient digestibility = 100－[100 × (% ADIA in DM consumed / % ADIA in feces) × (% nutrient in feces/nutrient consumed DM)].
Data on lactation performance, DMI, feed efficiency, blood metabolites, and rumen fermentation variables were analyzed utilizing
the PROC MIXED procedure of SAS (SAS Institute Inc., Cary, NC). The statistical model was as follows:

Yijk = μ ＋ Ti ＋ Dj(i)＋ Wk ＋ (TW)ik ＋ Eijk,

where Yijk is the response variable; μ is the overall mean; i = diet, j = cow; Ti is the effect of ith diet; Dj (i) is the random effect of the jth
cow within diet; Wk is the effect of the kth week; TWik is the interaction between ith diet and kth week; Eijk is the error term. The model
included diet, week, and interaction between diet and week as fixed effects, cow within diet as a random effect, and the week as a
repeated measure.
Data for nutrient digestibility was analyzed by the following model:

Yijk = μ ＋ Ti ＋ Dj(i) ＋ Eijk,

where Yijk is the response variable; μ is the overall mean; i = diet, j = cow; Ti is the effect of ith diet; Dj(i) is the random effect of the jth
cow with diet; Eijk is the error term. The model included diet as a fixed effect and cow within diet as a random effect. All data were
shown as least square means.
The repeated measures analysis of results was subjected to 5 covariance structures: AR, UN, CS, SP, and VC. The covariance
structure with the smallest Schwarz-Bayesian criterion was used owing to the most desirable and reliable analysis (Littell et al., 1998).
Least squares means were calculated and separated using the PDIFF option of SAS. Differences between diets were detected using
Duncan’s adjustment. Statistical significance was defined at P ≤ 0.05, and trends were declared at 0.05 < P ≤ 0.1.

3. Results

The results of feed intake and milk production performance are listed in Table 3. As shown in the table, the feed intake and milk
yield were affected by week and diet (P < 0.01), but the interaction of diet by week was not found. The HMS diet had lower feed intake
(P < 0.01), milk yield (P < 0.05), and yield of milk lactose (P < 0.01) than the other diets (Control, LMS, and MMS diets), while there
were no differences in feed intake and milk yield between the Control, LMS, and MMS diets. Lower lactose yield was observed for the
MMS diet than the Control (P < 0.05), with no difference relative to the LMS diet. The percentages of milk fat, protein, and SNF were
similar among the diets, whereas the yields of 4% FCM (P < 0.01), milk fat (P < 0.05), and milk protein (P < 0.01) were lower for HMS
versus Control or LMS diets, with no difference between Control and LMS diets. The Control and LMS diets had higher feed efficiency
than the HMS diet (P < 0.01), whereas no difference was found among Control, LMS, and MMS diets. Compared with the Control diet,
N conversion was lower for MMS and HMS diets (P = 0.01), with no difference relative to the LMS diet. The effect of week was
significant in 4% FCM (P = 0.05), milk protein yield (P = 0.03), milk lactose yield (P < 0.01), and N conversion (P = 0.03).
The rumen fermentation characteristics are listed in Table 4. As illustrated in the table, the rumen pH, ruminal butyrate con­
centration, and its molar proportion were not affected by the diets. Total VFA concentration was lower in HMS than in the Control and
LMS diets (P < 0.05), with no difference among the Control, LMS, and MMS diets. The concentration of NH3-N and the ruminal acetate
were higher for the HMS diet than all the other diets (P < 0.05), which had similar concentrations. The cows fed the HMS diet had a
higher molar proportion of ruminal acetate and a higher ratio of acetate to propionate than those fed all the other diets (P < 0.01), with
no difference for the Control versus LMS diet, or LMS versus MMS diet. And the molar proportion of ruminal acetate and the ratio of
acetate to propionate in the MMS diet were higher than those in the Control diet (P < 0.01). However, the ruminal propionate con­
centration and its molar proportion were lower in HMS than in the Control, LMS, and MMS diets (P < 0.01), with no difference between

Table 4
Effects of substitution of millet straw for corn silage and alfalfa hay on ruminal fermentation patterns in late-lactating Holstein cows.
Item3 Treatment1 SEM P-value2

Control LMS MMS HMS T W T×W

pH 6.43 6.52 6.54 6.63 0.063 0.27 0.93 0.59

Total VFA, mM 96.1a 95.9a 95.2ab 94.1b 0.49 0.04 0.25 0.80
Acetate, mM 64.6b 65.0b 65.5b 67.2a 0.41 < 0.01 0.38 0.96
Propionate, mM 21.2a 20.7ab 19.9b 17.7c 0.42 < 0.01 0.72 0.92
Butyrate, mM 10.2 10.2 9.79 9.40 0.22 0.14 0.67 0.94
Molar proportion, mM/100 mM
Acetate 67.1c 67.7bc 68.7b 71.7a 0.46 < 0.01 0.92 0.96
Propionate 22.1a 21.6ab 20.9b 18.7c 0.37 < 0.01 0.97 0.97
Butyrate 10.6 10.6 10.3 9.99 0.22 0.24 0.89 0.95
Acetate: propionate 3.05c 3.15bc 3.28b 3.83a 0.077 < 0.01 0.92 0.97
NH3-N, mg/dL 11.9b 12.5b 13.0b 14.6a 0.52 0.02 0.37 0.75
a-c
Different lowercase letters within a row indicate significant differences.
1
Control = 60% corn silage, 40% alfalfa hay, and 0% millet straw (on 100% roughage); LMS = 54% corn silage + 36% alfalfa hay + 10% millet straw;
MMS = 48% corn silage + 32% alfalfa hay + 20% millet straw; HMS = 42% corn silage + 28% alfalfa hay + 30% millet straw.
2
T = treatment effect; W = week effect; T × W = the interaction between treatment and week.
3
Total VFA = total volatile fatty acid; NH3-N = ammonia-N.

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Control and LMS diets, and LMS and MMS diets.

The total-tract apparent nutrient digestibility is shown in Table 5. According to the table, the OM digestibility was lower for the
HMS diet than for the other diets (P < 0.01), while no difference between the Control and LMS diets was found. The total-tract apparent
digestibility of DM, CP, NDF, and ADF decreased when feeding the HMS-based diet (P < 0.05), while it maintained a similar level in
other diets. The total-tract apparent digestibility of EE and NFC was not affected by diets.
All the blood metabolites (Table 6) were not affected by the diets except BUN concentration which was higher for the HMS diet than
the other diets (P < 0.05), with no difference among the Control, LMS, and MMS diets.

4. Discussion

In the present study, the DMI was not affected in the LMS and MMS groups, whereas it was lower in the HMS group, illustrating that
the 30% substitution ratio might be too much when replacing CS and AH with MS in dairy cows. The decreased DMI in the HMS group
could be partly due to the higher NDF content. Hayirli et al. (2002) and Mertens (1994) reported a negative association between DMI
and NDF content in dairy cows, and Dado and Allen (1995) found that DMI was limited when the dietary NDF levels exceeded 35%. In
the present study, the NDF level in the HMS diet was 36.3%, exceeding 35%, so the decreased DMI in the HMS group should be
acceptable. Moreover, Allen (1996) believed that DMI was bound by physical distension of the gastrointestinal effect. The lower DMI
was consistent with lower DM digestibility in the HMS group. This result indicated that the lower DMI in the HMS group also might be
attributed to the greater fill effect caused by lower DM digestibility. Oba and Allen (1999) demonstrated that DMI could be greatly
affected by NDF digestibility, which was higher and might have a shorter retention time, allowing greater DMI. Thus, the lower DMI in
the HMS group also might be explained by the lower NDF digestibility. Besides, the reduction in DMI in the HMS group was also
possibly caused by the poorer palatability of MS in comparison to AH and CS, so the substitution ratio at 30% might be too much. Our
results were supported by Muller et al. (1999), who reported that the palatability of roughages had a great effect on the DMI in dairy
cows. Based on the similar DMI among the Control, LMS, and MMS groups, whereas a lower DMI in the HMS group, the substitution of
MS for CS and AH at 20% is feasible in dairy cows.
Oba and Allen (1999) reported a positive association between DMI and milk yield and pointed out that the increased DMI was the
crucial factor for increased milk yield. In the present study, the changes in milk yield were consistent with the DMI among the
treatments, illustrating that the lower milk yield in the HMS group could have resulted from the lower DMI. Since milk yield is mainly
dependent on lactose concentration, the lower milk yield associated with lower lactose yield in the HMS group could be expected. In
this study, changes in lactose yield were in line with the blood concentration of glucose and ruminal propionate concentration (21.2a,
20.7ab, 19.9b, 17.7c). In dairy cows, propionate can convert into glucose in the liver, and after being transferred into the mammary
gland, glucose converts into lactose and is involved in milk synthesis (Lin et al., 2016). Besides, as propionate is mainly produced by the
fermentation of NFC (Wei et al., 2018), the lower milk yield in the HMS group also could be explained by the lower NFC content.
Additionally, the feed efficiency was also lower in the HMS group, which was associated with the DM digestibility, indicating that the
lower milk yield also might be attributed to the lower DM digestibility in the HMS group. A similar decrease in milk yield and feed
efficiency when substituting high-quality forage like AH with cereal straw was reported by Wang et al. (2014).
Milk fat is a crucial indicator to evaluate the production performance of dairy cows. It can be synthesized by acetate and butyrate,
and the greater concentration of acetate and butyrate usually results in a higher milk fat yield (Urrutia et al., 2019). In this study,
although the butyrate concentration (10.2, 10.2, 9.79, 9.40) and its molar proportion were not affected by treatment, the ruminal
acetate concentration (64.6b, 65.0b, 65.6b, 67.2a) increased significantly in the HMS group, indicating that cows in the HMS group
should produce higher milk fat. However, interestingly, the yield of milk fat was lowest in the HMS group. It is well known that the
rumen fermentation patterns are largely affected by the nutrient composition of feed, and acetate is mainly produced by NDF
fermentation (Doane et al., 1997). Thus, in this study, the ruminal acetate concentration increased in the HMS group could be
attributed to the higher NDF content. However, although the ruminal acetate concentration and its molar proportion were higher in

Table 5
Effects of substitution of millet straw for corn silage and alfalfa hay on the total-tract apparent nutrient digestibility in late-lactating Holstein cows.
Treatment1 SEM P-value
2
Item Control LMS MMS HMS

Apparent digestibility, %
DM 76.2a 75.6a 74.9a 70.9b 0.68 < 0.01
OM 77.4a 76.6ab 75.5b 71.9c 0.58 < 0.01
CP 78.1a 77.4a 76.5a 72.7b 0.54 < 0.01
NDF 58.4a 57.7a 56.3a 52.1b 1.06 < 0.01
ADF 55.9a 54.9a 54.1a 50.6b 1.07 0.02
EE 85.7 85.2 85.6 84.1 0.64 0.32
NFC 94.7 93.5 93.4 92.8 1.14 0.69
a-c
Different lowercase letters within a row indicate significant differences.
1
Control = 60% corn silage, 40% alfalfa hay, and 0% millet straw (on 100% roughage); LMS = 54% corn silage + 36% alfalfa hay + 10% millet straw;
MMS = 48% corn silage + 32% alfalfa hay + 20% millet straw; HMS = 42% corn silage + 28% alfalfa hay + 30% millet straw.
2
DM = dry matter; OM = organic matter; CP = crude protein; NDF = neutral detergent fiber; ADF = acid detergent fiber; EE = ether extract; NFC (non-
fiber carbohydrates) = 100% － (%NDF + % CP + % EE + % Ash).

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M. Wang et al. Animal Feed Science and Technology 299 (2023) 115622

Table 6
Effects of substitution of millet straw for corn silage and alfalfa hay on blood metabolites in late-lactating Holstein cows.
Item3 Treatment1 SEM P-value2

Control LMS MMS HMS T W T×W

b b b a
BUN, mM 2.13 2.27 2.37 2.79 0.10 0.02 0.33 0.98
Albumin, g/L 25.1 25.6 25.9 26.0 0.71 0.59 0.85 0.48
Total protein, g/L 75.3 75.5 76.7 73.1 1.77 0.57 0.86 0.30
Glucose, mM 2.74 2.73 2.78 2.51 0.072 0.09 0.88 0.40
Cholesterol, mM 6.88 7.22 7.33 7.28 0.26 0.63 0.98 0.90
Triglyceride, mM 0.41 0.44 0.41 0.41 0.017 0.38 0.69 0.98
ALT, U/L 31.5 29.7 34.0 34.1 1.42 0.12 0.54 0.83
AST, U/L 66.2 65.3 70.8 71.3 2.42 0.29 0.52 0.33
BHBA, mM 0.578 0.565 0.563 0.564 0.021 0.95 0.25 0.91
NEFA, μM 700 701 707 680 9.96 0.31 0.63 0.48
a-b
Different lowercase letters within a row indicate significant differences.
1
Control = 60% corn silage, 40% alfalfa hay, and 0% millet straw (on 100% roughage); LMS = 54% corn silage + 36% alfalfa hay + 10% millet straw;
MMS = 48% corn silage + 32% alfalfa hay + 20% millet straw; HMS = 42% corn silage + 28% alfalfa hay + 30% millet straw.
2
T = treatment effect; W = week effect; T × W = the interaction between treatment and week.
3
BUN = blood urea nitrogen; AST = aspartate aminotransferase; ALT = alanine aminotransferase; BHBA = β- hydroxybutyric acid; NEFA = non-
esterified fatty acid.

the HMS group, the lower total VFA concentration due to the lower DM digestibility (Beckman and Weiss, 2005) might ultimately lead
to limited differences in acetate production among treatments. Additionally, a previous study demonstrated that the ratio of acetate to
propionate positively affected milk fat yield (Razzaghi et al., 2020). In the present study, due to the greatest NDF and lowest NFC
content in the diet of the HMS group, the greater ratio of acetate to propionate was predictable. However, the milk fat yield was lower
in the HMS group, which was inconsistent with a previous study. A similar negative association between milk fat yield and acetate to
propionate ratio was also reported by Sandri et al. (2020), who believed that the milk fat yield should be mainly affected by total VFA
production, so the lower milk fat yield in the HMS group might be explained by the lower VFA concentration. Besides, since rumen
fluid collection before morning feeding might only reflect the rumen fermentation patterns at that time, whereas the yield of milk fat
was affected by the rumen fermentation throughout the day, this may lead to the inconsistency between the ruminal acetate con­
centration and milk fat yield. Therefore, although the NDF content was higher in the HMS group, the lower milk fat yield was
reasonable due to the lower DMI and NDF digestibility.
Milk protein yield and N conversion rate are also primary indicators to evaluate the production performance of dairy cows. Ac­
cording to the meta-analysis reported by Hristov et al. (2004), milk protein yield was positively associated with DMI and dietary CP
content. Thus, the lower milk protein yield in the HSM group might be caused by the lower CP content, DMI, and CP digestibility.
Besides, milk protein secretion in dairy cows is generally dependent on the supply of metabolizable protein (NRC, 2001), which mainly
consists of dietary rumen undegradable protein and microbial protein. Rumen undegradable protein is mostly provided by concentrate
(Schor and Gagliostro, 2001; Putri et al., 2019). But the concentrate composition was the same among the treatments in this study,
indicating limited changes in rumen undegradable protein content. However, Wang et al. (2014) found that microbial protein syn­
thesis was lower in substituting cereal straw for AH. Although microbial protein synthesis was not detected in this study, the greater
rumen concentration of NH3-N in the HMS group suggested that microbial protein synthesis might be lower (Herrera-Saldana et al.,
1990). In addition, microbial protein synthesis needs fermentable carbohydrate, which is mainly derived from NFC and NDF (Wang
et al., 2014). In the study, although the NFC digestibility was not affected, NDF digestibility was lowest in the HMS group, which could
partly prove that microbial protein synthesis was lower in the HMS group, ultimately resulting in the lower milk protein yield in the
HMS group.
Another interesting finding in the present study was that the N conversion was lower in the HMS group. Besides, the variation
tendency of N conversion was mostly consistent with CP digestibility, suggesting that the lower N conversion could be partly explained
by the lower CP digestibility. Additionally, as reported by Nousiainen et al. (2004), greater concentration of milk urea nitrogen, BUN,
and rumen NH3-N usually indicated less nitrogen balance and a lower N efficiency in dairy cows. The hypothesis was supported by
Wang et al. (2014), who reported an increase in milk urea nitrogen and rumen NH3-N, whereas a decrease in N conversion when
substituting cereal straw for AH. Conversely, a lower ruminal NH3-N concentration corresponded to a lower BUN concentration in the
control diet, indicating that the control-fed cows had better energy and nitrogen synchronization on rumen microbes (Landau et al.,
2005). Although the milk urea nitrogen was not measured in the present study, considering the greater concentration of rumen NH3-N
and BUN in the HMS group, a lower N conversion rate should be expected. And in this study, the milk yield, the yield of milk fat and
milk protein, feed efficiency, and N conversion were similar among the Control, LMS, and MMS groups, which only decreased in the
HMS group, suggesting that 20% should be the upper limit when substituting of MS for CS and AH in dairy cows.
In the present study, the total-tract apparent digestibility of DM, OM, CP, NDF, and ADF in the HMS group was all lower, which was
consistent with the results of Wang et al. (2014). In ruminants, the rumen plays a primary role in the digestion of feed, especially fiber
(Wang and McAllister, 2002), and the rumen digestion mainly depends on microbial degradation. As mentioned before, Wang et al.
(2014) found that microbial protein synthesis was limited when substituting cereal straw for AH, so the decreased digestibility of DM,
OM, CP, NDF, and ADF could be explained by the less microbial protein synthesis in the HMS group.

7
M. Wang et al. Animal Feed Science and Technology 299 (2023) 115622

5. Conclusions

Though microbial protein synthesis was not measured in the present study, the HMS treatment might decrease microbial protein
synthesis, resulting in lower nitrogen balance, production performance, and feed efficiency. Since the DMI, production performance,
and feed digestibility were similar among the Control, LMS, and MMS groups, whereas lower in the HMS group, substituting MS for CS
and AH is feasible in late-lactation dairy cows, but the substitution ratio should not exceed 20%.

CRediT authorship contribution statement

Meimei Wang: Data curation, Formal analysis, Methodology, Software, Writing – original draft. Yan Li: Data curation, Meth­
odology, Validation, Visualization. Jianfang Feng: Data curation, Investigation, Methodology, Software, Validation. Yizhao Shen:
Data curation, Formal analysis, Funding acquisition, Resources, Software, Writing – review & editing. Yufeng Cao: Investigation,
Methodology, Supervision, Validation, Visualization. Qiufeng Li: Conceptualization, Investigation, Resources, Validation, Visuali­
zation. Yanxia Gao: Funding acquisition, Methodology, Software, Supervision. Jianguo Li: Funding acquisition, Investigation, Project
administration, Resources, Supervision.

Declaration of interest

The authors declare that there is no conflict of interest.

Acknowledgments

This research was supported by the earmarked fund for CARS-36 (CARS-36), Hebei Dairy Cattle Innovation Team of Modern Agro-
industry Technology Research System (HBCT2018120203), Key Research and Development Project of Hebei (20326606D), the Top
Talent Project of Hebei Province (6012018), and Precision Animal Husbandry Discipline Group Construction Project of Hebei Agri­
cultural University (1090064).

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