1. Introduction
Taurine was first isolated as a component from
ox (Bos taurus, from which its name is derived) bile in 1827 [
1], and its chemical structure was determined in 1850. Since then, considerable efforts have been devoted to its chemical properties, functionalities, absorption and metabolism. Taurine is a sulfur-containing amino acid (AA) that accounts for approximately 0.1% of the animal body [
2]. Taurine cannot be used for protein synthesis [
3] but plays a wide range of pivotal roles in humans and animals, including functions in muscle growth [
4], energy metabolism [
5], bile acid conjugation [
6], immune response, antioxidation [
7], osmotic adjustment, regulation of blood pressure, management of the retinal and cardiac function, modulation of neuroendocrine activity [
8], and therapy of fatty liver disease [
9]. Taurine deficiency may cause weak energy metabolism and energy metabolism dysfunction [
5]. Taurine has been widely used as a feed additive in aquatic and poultry animals to enhance feed efficiency, promote growth, and prevent diseases. Previous studies showed that supplementing the diet with taurine at 2.5 or 5.0 g/kg improved the feed intake, egg production, and apparent digestibility of quails [
10] and the addition of taurine at 12.3 g/kg and 16.7 g/kg improved the growth and health status of belugas [
11]. It was also reported that adding taurine to the diet of chicken improved antioxidant activity and growth performance [
12,
13].
In mammals, taurine is mainly synthesized in the liver using methionine as the precursor [
14]. However, the amount of taurine synthesized in the animal body can only meet 30–40% of their requirement [
15]. Animal products, especially aquatic products, contain a high level of taurine, whereas plants almost do not contain taurine [
16]. As herbivores, beef cattle cannot be fed with feeds of animal products. Therefore, adult beef cattle are quite possibly in a taurine deficiency status. Supplementing their diets with taurine could possibly improve the growth performance and feed utilization efficiency of beef cattle. To the best of our knowledge, however, no data are available regarding the ruminal degradation of taurine and its effects on rumen fermentation. Free amino acids (AA), such as methionine and lysine, are extensively degradable by ruminal microorganisms [
17]. As a sulfur-containing AA, it could be hypothesized that taurine is also degradable and plays an important role in rumen fermentation. Therefore, the objectives of the present study were to investigate the effects of taurine on rumen fermentation and the ruminal degradation of taurine using
in vitro rumen fermentation techniques to clarify the usefulness of taurine as a feed additive for beef cattle.
3. Results and Discussion
Dietary carbohydrates can be fermented to produce VFA, and CP can be degraded into peptides, AA and NH
3 to some extent by rumen microorganisms. Amino acids such as methionine and lysine are easily hydrolyzed to NH
3 and CO
2 by rumen microorganisms [
27]. Many
in vitro studies showed that supplementing N sources (urea, casein or ovalbumin) or sulfur-containing substances (sodium sulfate) increased the concentration of NH
3-N [
28,
29]. The results in
Table 3 showed that the ruminal degradability of taurine increased linearly with incubation time, and the 2 h
in vitro degradability of taurine was found to be as high as 99 %, indicating that taurine was quickly and highly degradable for
in vitro rumen fermentation. The results in
Table 4 show that taurine significantly increased the 48 h-pH value (
p = 0.018) and decreased the 48 h-DMD (
p = 0.008). Ruminal microorganisms should have played important roles in hydrolyzing taurine. However, the actual reactions of the hydrolyzing process are unclear and need to be investigated in further research.
The results in
Table 5 showed that taurine increased the NH
3–N concentration of incubation liquid in a linear manner (
p < 0.001), which was in accordance with taurine degradability. Previous
in vitro rumen fermentation studies showed that adding methionine increased the ruminal MCP concentration [
30] and confirmed that, in cattle, supplementing with methionine or sodium sulfate improved the ruminal MCP synthesis [
31]. As a semi-essential AA, taurine contains N and sulfur, which are essential nutritional elements for MCP synthesis. The results in
Table 5 showed that taurine quadratically affected the ruminal MCP synthesis (
p < 0.001) and that low levels of taurine improved the ruminal MCP concentration, whereas a high level of taurine decreased the ruminal MCP concentration, suggesting that the effect of taurine on improving MCP synthesis was on a dose-dependent basis. The beneficial impact of low taurine on increasing ruminal MCP synthesis could possibly be resulted from the increased NH
3–N concentration and sulfur from taurine degradation. However, the effect should not be attributed to the direct use of taurine for MCP synthesis because taurine cannot be incorporated into protein [
3]. The decreasing impact of high taurine on the ruminal MCP concentration could be a result of the inhibitive effect of a high level of NH
3–N [
32,
33]. The normal concentration of rumen NH
3–N ranges from 6.3 to 27.5 mg/100 mL, and the optimum ruminal concentration of NH
3-N required to support the maximum synthesis of MCP is 12.8 mg/100 mL [
34]. The MCP synthesis can be inhibited when the rumen NH
3-N reaches or exceeds 27.5 mg/100 mL, as indicated in the present study (
Table 5).
Dietary carbohydrates can be extensively degraded and fermented by rumen microorganisms to produce VFA. The results in
Table 5 also showed that taurine decreased the total VFA concentration (
p = 0.014), but it did not affect the molar proportions of individual VFA (
p > 0.10) of the incubation liquid. The results were in accordance with the decreased 48 h total gas production (
p < 0.001), the potential gas production (
a +
b) (
Table 6) (
p < 0.001) and the 48 h-DM degradability (
Table 4) (
p = 0.008). The results suggested that taurine inhibited the ruminal fermentation of feed carbohydrates. Since taurine decreased the gas production of the slowly degradable fraction of feeds (
b) (
p < 0.001), which is mainly cellulose and hemicellulose, taurine should have an inhibitive impact on the activity of ruminal cellulolytic microorganisms. However, the hypothesis needs to be clarified by investigating the effects of taurine on ruminal microbiota in the future. The results in
Table 6 showed that taurine quadratically affected the CH
4 (
p = 0.026) and CO
2 (
p = 0.019) production, suggesting that a suitable level of taurine had the potential to decrease ruminal CH
4 production.
Ruminal pH is mainly affected by the NH
3–N from feed CP degradation and the VFA from ruminal fermentation of feed carbohydrates [
35,
36]. The results in
Table 4 and
Table 5 showed that taurine increased the ruminal pH. The results could be attributed to the increased NH
3–N and the decreased total VFA concentrations.