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Article

Effects of Guanidinoacetic Acid on Ruminal Fermentation and Greenhouse Gas Production Using Fresh Forage and Silage from Different Maize (Zea mays L.) Genotypes

by
Edwin Rafael Alvarado-Ramírez
1,2,
Verónica Andrade-Yucailla
3,
Mona Mohamed Mohamed Yasseen Elghandour
4,
Néstor Acosta-Lozano
3,
Marco Antonio Rivas-Jacobo
1,
Daniel López-Aguirre
2,
Jonathan Raúl Garay-Martínez
5,
Paulina Vazquez-Mendoza
6,
Marcos Barros-Rodríguez
7,* and
Abdelfattah Zeidan Mohamed Salem
4,*
1
Facultad de Agronomía y Veterinaria, Universidad Autónoma de San Luis Potosí, Soledad de Graciano Sánchez 78321, San Luis Potosí, Mexico
2
Facultad de Ingeniería y Ciencias, Universidad Autónoma de Tamaulipas, Ciudad Victoria 87149, Tamaulipas, Mexico
3
Facultad de Ciencias Agrarias, Universidad Estatal Península de Santa Elena, La Libertad 204102, Santa Elena, Ecuador
4
Facultad de Medicina Veterinaria y Zootecnia, Universidad Autónoma del Estado de México, Toluca 50200, Estado de Mexico, Mexico
5
Campo Experimental Las Huastecas, Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias, Altamira 89610, Tamaulipas, Mexico
6
Facultad Maya de Estudios Agropecuarios, Universidad Autónoma de Chiapas, Catazajá 29980, Chiapas, Mexico
7
Facultad de Ciencias Agropecuarias, Universidad Técnica de Ambato, Cevallos 180103, Tungurahua, Ecuador
*
Authors to whom correspondence should be addressed.
Fermentation 2023, 9(5), 437; https://doi.org/10.3390/fermentation9050437
Submission received: 10 April 2023 / Revised: 28 April 2023 / Accepted: 29 April 2023 / Published: 2 May 2023
(This article belongs to the Special Issue In Vitro Digestibility and Ruminal Fermentation Profile)
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Abstract

:
Guanidinoacetic acid (GAA) is a feed additive that promotes growth in animals, while maize (Zea mays L.) is used for the mitigation of ruminal greenhouse gases. However, it is unknown if GAA affects the efficiency of maize in mitigating gases or if there is synergy between them. Therefore, the objective of this study was to evaluate the in vitro production of total gas, methane (CH4), carbon monoxide (CO), and hydrogen sulfide (H2S), ruminal fermentation characteristics, and the CH4 conversion efficiency of fresh forage and silage of different genotypes (Amarillo, Montesa, Olotillo, Tampiqueño, and Tuxpeño) of maize, with and without the addition of GAA. The silage of the Amarillo genotype without AAG had the highest (p = 0.01) total gas production rate and the lowest (p = 0.044) delay time before gas production. In addition, at 48 h, the Amarillo silage with GAA increased the production of total gas (p = 0.0001) and CH4, as well as the proportion of CH4 (mL CH4 100 mL−1 total gas). The Amarillo and Tuxpeño genotype produced more (p = 0.033) CO in the first 24 h of incubation, while silage and the addition of GAA only increased (p = 0.001) CO at 6 h. The highest (p = 0.02) H2S production was observed with the ensiled Amarillo genotype with GAA. Regarding fermentation characteristics, the silage of the Amarillo and Montesa genotypes presented the highest degradation of dry matter (DMD), short-chain fatty acids (SCFA), and metabolizable energy (ME), and although there was no effect on CH4 efficiency, the Amarillo and Olotillo genotypes produced more SCFA, ME, and OM per unit of CH4. It can be concluded that rumen gas production, fermentation characteristics, and CH4 conversion efficiency are more influenced by the maize genotype and forage condition than by the addition of guanidinoacetic acid, and of the genotypes evaluated, the forage silage from Amarillo showed the best characteristics and efficiency of CH4.

1. Introduction

Ruminant livestock is considered the most efficient in feed conversion compared to non-ruminant livestock [1] because these animals have a digestive system that can take advantage of fibrous forages and transform them into higher-quality products for human nutrition [2,3]. However, during the ruminal fermentation of the forage, they produce gases that, when expelled into the atmosphere, cause the greenhouse effect, thus contributing to global warming [4]. Among these gases is methane (CH4), a gas that is formed from hydrogen (H2) and carbon dioxide (CO2) produced during the fermentation of carbohydrates and which has a higher warming potential than CO2 [5]. Other no less important gases are carbon monoxide (CO), which is a precursor of ozone in the atmosphere [6], and hydrogen sulfide (H2S), which serves as an alternate sink for H2 to reduce CH4 production [7]. However, H2S in high concentrations can affect the digestive system and be toxic to animals [8], and although the production of all these gases is inevitable, high amounts indicate a loss of nutrients and energy [9]. Therefore, reducing the production of greenhouse gases (GHG) without causing an imbalance in the rumen and affecting the productivity of the animals is a challenge [10].
A viable alternative to deal with this problem is to manipulate the diet through the selection of forages of high nutritional quality, in terms of highly digestible protein and carbohydrates [11]. In this sense, maize (Zea mays L.), as a source of forage, provides fibrous carbohydrates and non-fibrous carbohydrates, such as water-soluble forms and starch [12,13,14], which is why it is considered to have high energy value for animal nutrition [15] and is used as a strategy to mitigate GHG emissions in ruminants, especially CH4 [16,17]. In this regard, it has been shown that the silage of whole maize plants reduces the production of ruminal CH4 [18] since the starch it contains promotes the formation of propionate and decreases the H2 available to produce CH4. Furthermore, silage favors post-ruminal digestion [19], which improves performance in cattle and consequently decreases the CH4 per unit of the animal product [20].
Another strategy that has been implemented is the use of feed additives, which have been shown to positively alter ruminal and post-ruminal metabolism, improving nutrient utilization and animal productivity [21,22] and thereby reducing the intensity of greenhouse gas emissions [23]. In this sense, guanidinoacetic acid (GAA; also called glycinamide), a derivative of arginine and glycine, is a natural precursor of creatine biosynthesis [24]. In turn, creatine is part of the energy and protein metabolism of animals [25,26,27], which is why GAA has been used as an additive to improve the performance of animals, including ruminants [28,29,30]. In this regard, it has been reported that in beef cattle, the addition of 0.03 to 0.40% (on DM basis) GAA in the diet increases daily weight gain, dry matter digestibility, and in some cases, feed conversion efficiency, which was reflected by a higher yield and quality of the carcass [31,32,33,34], without negative effects on the serum content of arginine, folate, homocysteine, and methionine [34]. In sheep, studies indicated that with the addition of 0.08 to 0.10% (on DM basis) GAA, an effect similar to that observed in beef cattle is obtained, but of a lesser magnitude [35,36,37,38]. These improvements were not only reflected by the increase in the carcass yield, but also by the increase in glycogen, intramuscular fat, protein, antioxidant capacity, and moisture retention in the meat [35,39]. In addition, when combined with 0.10% (on DM basis) N-carbamyl glutamate, it improved the distribution of body fat and muscle composition in sheep [40]. Meanwhile, in dairy cattle, it has been reported that the addition of 0.03 to 0.09% (on DM basis) GAA to the diet improves feed efficiency and rumen fermentation, which increases yields and milk components [41]. However, it is unknown if, at the rumen level, the addition of GAA negatively affects the efficiency of maize in mitigating greenhouse gases or if there is synergy between maize forage and GAA. Therefore, the objective of this study was to evaluate the in vitro production of total gas, CH4, CO, and H2S, as well as the characteristics of rumen fermentation and the CH4 conversion efficiency of fresh forage and silage from different maize genotypes, with and without the addition of GAA.

2. Materials and Methods

2.1. Experimental Treatments

The treatments consisted of the evaluation of the fresh forage and silage of four genotypes of maize native to Mexico (Amarillo, Olotillo, Tampiqueño, and Tuxpeño) and a commercial hybrid (Montesa), with and without the addition of 0.3% (on DM basis) guanidinoacetic acid (GAA). The addition level was defined according to the GAA percentages and the results reported in previous studies and corresponded to the average of the doses that presented the best results in both sheep and cattle (beef and dairy). The GAA was acquired from the company Evonik México S.A. de C.V. (Mexico City, Mexico) under the trade name of GuanAMINO® for ruminants, and at the time of evaluation the GAA had a purity of 96%.

2.2. Fresh Forage and Silage Production

Forage production was carried out in the city of Aldama, Tamaulipas, México (22°59′09″ N and 98°10′25″ W, at 190 masl) during the rainy season (May–October) of 2021. On the site, the soil is clay–loam in texture, with a high organic matter content, moderate alkalinity, and low salinity. The climate, according to Köppen’s classification, is of the Aw0 type, which corresponds to the driest of the warm sub-humid [42]. The sowing was carried out at a density of 62,500 plants ha−1, and during the growth and development of the crop, pesticides and herbicides were not applied, since pest and weed control was carried out manually. The harvest was carried out when the grain reached the milky-mass stage, and for this, a minimum of 10 plants from different points of each genotype were selected. These plants were crushed with a forage chopper and manually mixed until homogeneous and, later, samples were obtained for the analysis of the chemical composition, which thereafter was called “fresh forage”. Regarding the silages, they were made by placing 5 kg of fresh and chopped forage in black polyethylene plastic bags (30 cm in diameter × 50 cm in height, caliber 500), which were vacuum sealed with the help of a domestic vacuum cleaner and a heat sealer. After 120 days of fermentation, the silages were opened, and “ensiled forage” samples were obtained.

2.3. Chemical Composition

The fresh and ensiled forage of the five genotypes was dehydrated at 60 °C to a constant weight and ground in a hammer mill (Thomas Wiley® Laboratory Mill model 4, Thomas Scientific™, Swedesboro, NJ, USA) with a 2 mm sieve. Organic matter was determined by estimating the ash content using the method of Thiex et al. [43] and subtracting the value obtained from 100. The crude protein (CP) was calculated by multiplying the nitrogen content [44] by the conversion factor of 6.25. The neutral detergent fiber (NDF) and acid detergent fiber (ADF) were analyzed with the methodology described by Van Soest et al. [45] based on the ANKOM200 fiber analyzer (ANKOM Technology Corp., Macedon, NY, USA), while acid detergent lignin was measured through solubilization with a sulfuric acid solution [46]. The ethereal extract (EE) followed the Padmore [47] method, while the non-structural (NSC) and total (TC) carbohydrates were determined with the equations of Mertens [48] and Sniffen et al. [49]:
NSC = 100 − (CP + NDF + EE + Ash)
TC = 100 − (CP + EE + Ash)

2.4. In Vitro Incubation

The nutrient solution was prepared following the methodology of Goering and Van Soest [50] and the rumen fluid was obtained from the ruminal content of four male sacrificed sheep with an average weight of 45 kg. These animals were sacrificed in the municipal slaughterhouse of Toluca, State of Mexico, Mexico, and the rumen content was transferred to the laboratory in a hermetic thermos, where it was filtered with four layers of gauze to extract the rumen liquid. The incubation was carried out in glass vials with a capacity of 160 mL, and to each one, 500 mg of a dehydrated sample of fresh forage and silage of each genotype, 40 mL of nutrient solution, and 10 mL of rumen fluid were added, in addition to 1.5 mg of GAA to 30 vials. The vials were sealed with butyl rubber stoppers and aluminum seals, lightly shaken, and incubated in an incubator (Binder® BD series, BRINDER Inc., Tuttlingen, BaWü, Germany) at 39 °C throughout the evaluation. In total, three incubation runs were performed, and in each one they were incubated in 63 vials, 30 with GAA, 30 without GAA, and 3 blanks (without substrate), since in all cases, samples were incubated in triplicate.

2.4.1. Production of Total Gas, Methane (CH4), Carbon Monoxide (CO), and Hydrogen Sulfide (H2S)

Total gas production (psi) was measured at different incubation times (from 2 to 48 h after incubation—see figures) using a digital pressure gauge with ±2% accuracy (Manometer model 407910, Extech® Instruments, Nashue, NH, USA), following the technique of Theodorou et al. [51]. In addition, the production of CH4, CO, and H2S was evaluated, and for its estimation, the methodology proposed by Acosta et al. [52] was used, which consists of injecting gas extracted from the vials into a portable gas detector (Dräger X-am®, model 2500, Dräger, Lübeck, SH, Germany) using an external pump (Dräger X-am®, Dräger, Lübeck, SH, Germany). At the end of each measurement, the gas accumulated in the upper part of the vials was released with a syringe without a plunger to avoid greater accumulation of gas and the partial dissolution of the gases evaluated [53].

2.4.2. Ruminal Potential of Hydrogen (pH) and Dry Matter Degradability (DMD)

After the in vitro evaluation, the contents of the vials were filtered to retain the residual substrate and collect the liquid in beakers. In the liquid, the pH was measured using a potentiometer with a glass electrode (pH wireless electrode HALO® model HI11102, Hanna® Instruments, Woonsocket, RI, USA), while with the residual substrate, the apparent degradation was estimated using the difference between the weight of the sample at the beginning and the end of incubation; then, it was washed with plenty of water and dehydrated at 60 °C to a constant weight [54].

2.4.3. Calculations

The production kinetics of total gas, CH4, CO, and H2S were estimated by adjusting the volume of gases with the NLIN procedure from SAS [55], according to the model proposed by France et al. [56]:
y = b × [1 − e−c(t−Lag)]
where
y = volume (mL) of TG, CH4, CO and H2S at time t (h).
b = asymptotic TG, CH4, CO and H2S production (mL g−1 DM).
c = rate TG, CH4, CO and H2S production (mL h−1).
Lag = initial delay time before TG, CH4, CO and H2S production begins (h).
The metabolic energy (ME; MJ kg−1 MS) was estimated according to the equation proposed by Menke et al. [57]:
ME = 2.20 + (0.136 × TGP) + (0.057 × CP)
where
CP = crude protein (g kg−1 DM)
TGP = net gas production (mL 200 mg−1 DM) at 24 h of incubation.
The total concentrations of short-chain fatty acids (SCFA; mmol 200 mg−1 DM) were calculated according to Getachew et al. [58] as:
SCFA = (0.0222 × TGP) − 0.00425
where
TGP = total gas production (mL 200 mg−1 DM at 24 h of incubation.
Additionally, the ratio between CH4 and the SCFA (CH4:SCFA; mmol mmol−1), ME (CH4:ME; g MJ−1), and OM (CH4:OM; mL g−1) was calculated.

2.5. Statistical Analysis

The experimental design was completely randomized with a 5 × 2 × 2 factorial arrangement (five maize genotypes × two states of the forage × two levels of guanidinoacetic acid addition) and three repetitions. The data from the three replicates of each treatment in each run were averaged, and the averages obtained were used as the experimental unit of each treatment. Analysis was performed using the GLM procedure of SAS [55] with the following statistical model:
Yijk = µ + Gi + Sj + Ak + (G × S)ij + (G × A)ik + (S × A)jk + (G × S × A)ijk + εijk.
where Yijk is the response variable, μ is the overall mean, Gi is the effect of maize genotype (MG), Sj is the effect of the state of the forage (SF), Ak is the effect of the level of GAA addition, (G × S)ij is the effect of the interaction between the MG and SF, (G × A)ik is the effect of the interaction between the MG and the level of addition of GAA, (S × A)jk is the effect of the interaction between the SF and the level of addition of GAA, and (G × S × A)ijk is the effect of the interaction between the MG, the SF, the level of GAA addition, and εijk, the experimental error. Tukey’s test was used for the comparison of means, and p < 0.05 was the significance threshold.

3. Results

The chemical composition changed in all genotypes after silage, which was to be expected, since silage is a method of conservation that is based on the acidification of the forage through the catabolism of carbohydrates and other compounds (Table 1).

3.1. Ruminal Gas Total Production

The Amarillo genotype presented the highest (p = 0.001) asymptotic total gas production and the shortest (p = 0.0003) time in the lag phase and that Montesa at 24 h and Amarillo at 48 h produced more (p = 0.002) total gas for DM incubated (Figure 1). In addition, the ensiled forage increased (p = 0.042) the asymptotic total gas, the total gas production rate, and the total gas production by DM incubated and, without GAA addition, increased (p = 0.028) the rate and production of total gas only at 24 h, with no effect in the rest of the incubation period. For the interactions, it was observed that MG × SF influenced (p = 0.049) the parameters and the total gas production; MG × GAA influenced the production rate, asymptotic total gas, and total gas production at 6 h; SF × GAA influenced the total gas production rate; and MG × SF × GAA influenced the total gas production rate, the time in the lag phase, and the total gas production at 6 h by DM degraded. The last interaction showed that the Amarillo genotype silage without GAA obtained the highest total gas production rate (p = 0.01) and the shortest (p = 0.0436) time in the lag phase and that this did not influence total gas production during incubation for this and the rest of the genotypes, except at 24 h per DM (Table 2).

3.2. Ruminal Methane (CH4) Production

Although MG did not affect the parameters and the production of CH4 throughout the incubation, it was observed that the silage increased (p = 0.014) the production rate and the production of CH4 per DM incubated at 6 h and that a lack of GAA addition also increased (p = 0.002) the production of CH4 at 6 h (Figure 2). In the interactions, it was observed that MG × SF affected (p = 0.029) the CH4 asymptote and CH4 production throughout the incubation, except at 6 h for CH4 per DM incubated. In contrast, the MG × GAA interaction only affected CH4 production at 6 h, while SF × GAA did not affect the parameters and CH4 production. MG × SF × GAA influenced (p = 0.023) the production rate, time in the lag phase, and CH4 production per DM incubated at 6 h, although this did not impact the production of CH4 in the rest of the incubation period.
The result also showed that the Amarillo genotype, the silage forage, and a lack of GAA addition resulted in the highest (p = 0.033) proportion of CH4 at 6 h but did not affect that at 24 and 48 h, as well as that at 6 h between genotypes. The interaction of MG × SF showed an effect (p = 0.028) on the proportion and of CH4 throughout the incubation, except for at 6 h with the proportion of CH4. Furthermore, MG × GAA and MG × SF × GAA only affected (p = 0.012) the proportion of CH4 at 6 h, and SF × GAA did not influence any parameter (Table 3).

3.3. Ruminal Carbon Monoxide (CO) Production

It was observed that MG, SF, GAA, and their interactions did not have an impact on the parameters of CO production, but that Amarillo and Tuxpeño produced more (p = 0.033) CO per DM incubated and degraded at 24 and 6 h, respectively. Moreover, the silage and the addition of GAA increased (p = 0.001) CO at 6 h. For the interactions, it was found that MG × SF affected (p = 0.025) the production of CO during the entire incubation and that MG × GAA only influenced (p = 0.02) the production of CO by DM incubated at 6 and 24 h, while the rest of the interactions did not show a significant effect on the production of CO (Figure 3 and Table 4).

3.4. Ruminal Hydrogen Sulfide (H2S) Production

It was found that MG, SF, GAA, and their interactions did not have an impact on the H2S production parameters, except for silage, which increased (p = 0.023) the time in the lag phase. In this regard, the highest (p = 0.02) H2S production was observed with the Amarillo genotype, the ensiled forage, and with the addition of GAA. In the interactions, MG × SF affected (p = 0.0001) H2S production at 48 h and MG × GAA influenced (p = 0.041) that throughout the incubation. In addition, for SF × GAA, no significant effect was observed, and MG × SF × GAA showed that Montesa and Amarillo silage, with and without GAA, produced the highest and lowest (p = 0.002) amount of H2S, respectively (Figure 4 and Table 5).

3.5. Fermentation Characteristics and CH4 Conversion Efficiency

The results showed that MG, SF, GAA, and their interactions influenced (p = 0.03) the fermentation characteristics and did not affect the CH4 conversion efficiency, except for the MG × SF interaction (p = 0.028). It was observed that the Amarillo genotype was similar to Montesa and that both were associated with the lowest pH and degradation of dry matter (DMD), short-chain fatty acids (SCFA), and the highest metabolizable energy (ME), while Olotillo resulted in the highest pH and the lowest values (p = 0.0015) of DMD, SCFA, and ME. In addition, the ensiled forage resulted in the lowest (p = 0.03) pH and the highest DMD and SCFA levels and the same ME as the fresh forage, while, without the addition of GAA, the SCFA levels and ME increased (p = 0.0057) without causing variations in the pH and DMD. The MG × SF interaction affected (p = 0.0028) the pH, DMD, SCFA levels, and ME, as well as the CH4 conversion efficiency, while the others only influenced (p = 0.022) the DMD. In particular, the MG × SF × GAA interaction showed that the ensiled forage of all the genotypes and the addition of GAA increased the DMD, except for with the Tampiqueño genotype, where an opposite response was observed (Table 6).

4. Discussion

4.1. Ruminal Gas Total Production

Total gas production is generally associated with greater feed degradability [52] and in turn, with the chemical composition, especially with the concentration of rapidly fermenting carbohydrates [59]. In this study, the highest total gas production and the shortest time in the lag phase were obtained with the Amarillo genotype and the ensiled forage, which was associated with a higher concentration of non-structural carbohydrates in both the genotype and the silage. In this regard, Kholif et al. [60] reported that silage increases non-structural carbohydrates and that this favors the concentration of water-soluble carbohydrates, which represent additional energy for the rumen microbiota and are reflected by greater microbial activity. This coincides with what has been reported in other investigations where they found that in fibrous forages, including maize forage, silage serves as a treatment to reduce structural carbohydrates and that this allows rumen microbes to adhere more easily to the forage and proliferate more rapidly [61]. Contrary to this, forages with a high content of structural carbohydrates promote low rumen fermentation and, therefore, lower total gas production per unit of DM incubated [62]. This statement was corroborated in this study since the Olotillo genotype was associated with the highest percentage of structural carbohydrates, compared to the rest of the genotypes, and showed the longest time in the lag phase and the lowest rate and production of asymptotic total gas. However, it is interesting to note that although the Tuxpeño genotype did not result in more non-structural carbohydrates compared to those with the Amarillo genotype, it was the most efficient in what can be called “degradation efficiency”, since it produced more total gas per unit of DM with the highest total gas production rate. This could be caused by a balance between structural (NDF, ADF, and ADL) and non-structural carbohydrates in this genotype, which allowed ruminal microorganisms to improve digestion and nutrient extraction. Regardless of the state of the forage, in some genotypes, the addition of GAA caused a slight reduction in the production rate and total gas production, which can be attributed to possible momentary inhibition of the growth of some rumen microorganisms while they adapted, since GAA is susceptible to microbial degradation when supplied through the diet and is not protected ruminally [63,64], as in the case of this study.

4.2. Ruminal Methane (CH4) Production

Considering that, in this study, the Amarillo genotype was the one that produced the highest asymptotic gas total, it can be assumed that the CH4 may be split into a larger part of the gas that was produced during digestion. The interaction of genotypes and status showed that both genotype and forage status played an important role in reducing CH4 production. Silage is a conservation method that is based on a fermentation process in an anaerobic environment; thus, it is possible some methanogenic activity would have occurred during this process, resulting in reduced CH4 production during storage incubation [65]. Zhu et al. [66] reported that lactic acid bacteria are important during the silage process since they are the predominant microorganisms in the silage and are in charge of catabolizing carbohydrates for pH reduction [67]. Likewise, this microbe can reduce CH4 due to bioactive compounds, such as b-hydroxy-propionaldehyde (reuterin), an antimicrobial compound resistant to protease (PRA-1), which it produces [68,69,70]. Along the same lines, analysis of another genotype showed that the fresh forage produced less CH4 than the ensiled state, which demonstrated that it is possible that during incubation some genotypes produce fermentation products/metabolites that favor the formation or production of CH4 among themselves. However, it is important to mention that with some genotypes, the addition of GAA decreased the production of CH4, which is consistent because GAA decreases the population of protozoa and methanogens [41], ruminal microorganisms associated with the production of CH4. Furthermore, it has been shown that GAA increases propionate formation and decreases the acetate:propionate ratio [34,36], which translates to a lower availability of metabolic H2 for CH4 production.

4.3. Ruminal Carbon Monoxide (CO) Production

CO is an intermediate product of DM degradation, so its production is associated with incomplete feed degradation or lower microbial activity in the rumen [71]. In addition, it plays an important role in the biological cycle of bacteria and archaea since it represents an essential metabolite and acts as a regulator of some important metabolic pathways [72]. Considering that in this study the trend observed in CH4 production with the genotypes, forage states, and with and without the addition of GAA was similar to that obtained for CO, the variations in production can be attributed to the constant oxidation of CO to carbon dioxide (CO2) and hydroxide (H2) caused by rumen microorganisms, such as methanogenic bacteria and acetogens, for the production of CH4 and acetate [73,74,75]. In addition, it could be assumed that the addition of GAA inhibits the activity of methanogens, and, for this reason, there was a higher accumulated production of CO at 48 h of incubation, since there was no rapid oxidation of CO.

4.4. Ruminal Hydrogen Sulfide (H2S) Production

H2S plays a crucial role in the physiological function and maintenance of the gastrointestinal tract of animals [71], as well as in the reduction in ruminal CH4 production. It has been reported that during the degradation process, amino acids containing sulfur (S) groups, including methionine and cysteine, are catabolized by rumen microbes to produce H2S [76]. Under this assumption, the high production of H2S with the ensiled forage of the Amarillo genotype may be associated with the concentration and degradation of the amino acids that comprise it, especially those amino acids that contain sulfur [77]. Meanwhile, the low H2S production with the Montesa silage may be attributable to the addition of GAA, since there was an interaction between the genotype and the addition of GAA, and it has been reported that the use of additives may influence the reduction in H2S production in the rumen [78].

4.5. Fermentation Characteristics and CH4 Conversion Efficiency

This study showed that the genotype and the state of the forage are the factors that most affect the ruminal fermentation profile and that the silages of the Amarillo and Montesa genotypes were the ones that resulted in the lowest pH and the highest dry matter degradation (DMD), short-chain fatty acids (SCFA), and metabolizable energy (ME). The reduction in pH is rational, taking into account that all forage after silage contains more easily fermentable carbohydrates than in its fresh state [60] and that this favors a decrease in the pH [79]. Consequently, this propitiates a favorable environment for the proliferation of fibrinolytic bacteria, which require a pH between 5.5 and 7.0 [80], thereby increasing microbiological activity and increasing forage degradation. In the particular case of the Amarillo genotype, the DMD of the ensiled forage can also be attributed to the shorter time in the lag phase and the high rate of degradation that resulted, since this helped the forage to ferment for longer. In addition, it has been reported that SCFAs are directly proportional to the degradation, and therefore, the greater the degradation of the forage, the higher the SCFA levels will be, which increases the availability of energy [81,82]. On the other hand, it has been reported that the addition of GAA increases the population of some rumen microbes [34,36], and thus, this cannot be ruled out as the cause of the increase in SCFA and ME in the ensiled forage with the Amarillo and Olotillo genotypes. Similarly, although there was no significant effect of the addition of GAA on the efficiency of CH4, it was observed that efficiency increased quantitatively with all genotypes with GAA, since regardless of the state of the forage, the majority decreased CH4 per unit from SCFA, ME, and OM. However, the state of the forage did affect the efficiency with the Olotillo and Amarillo genotypes; with Olotillo, this can be attributed to the low degradability caused by the high concentration of the detergent acid lignin, while with Amarillo, it was believed to be due to the increase in DMD, which increases gas production [83].

5. Conclusions

It is concluded that the production of total gas, CH4, CO, and H2S, in addition to the ruminal fermentation characteristics and the CH4 conversion efficiency are more influenced by the maize genotype and the state of the forage than by the addition of GAA. Of the genotypes evaluated, Amarillo resulted in the lowest incubation pH and the highest DMD, SCFA levels, and ME, and in both forage states, the addition of GAA improved SCFA levels, ME, and the CH4 conversion efficiency, especially in silage, where the DMD also increased. Therefore, it is suggested that GAA be used as an additive and/or nutritional supplement in ruminant diets that include fresh forage or maize silage, since it does not negatively alter the fermentation profile and the production of ruminal greenhouse gases.

Author Contributions

Conceptualization, E.R.A.-R., A.Z.M.S., V.A.-Y., M.M.M.Y.E., M.A.R.-J., and M.B.-R.; methodology, E.R.A.-R., A.Z.M.S., P.V.-M., D.L.-A., and J.R.G.-M.; software, A.Z.M.S.; formal analysis, E.R.A.-R., A.Z.M.S., M.M.M.Y.E., and M.A.R.-J.; investigation, E.R.A.-R., A.Z.M.S., M.M.M.Y.E., V.A.-Y., and P.V.-M.; resources, E.R.A.-R., A.Z.M.S., M.M.M.Y.E., V.A.-Y., and N.A.-L.; data curation, E.R.A.-R., A.Z.M.S., and M.M.M.Y.E.; writing—original draft preparation, E.R.A.-R., A.Z.M.S., V.A.-Y., M.M.M.Y.E., N.A.-L., and M.B.-R.; writing—review and editing, E.R.A.-R., A.Z.M.S., M.M.M.Y.E., M.A.R.-J., D.L.-A., and J.R.G.-M.; visualization, A.Z.M.S., M.M.M.Y.E., and M.A.R.-J.; supervision, A.Z.M.S., M.M.M.Y.E., and M.A.R.-J.; Project Administration, E.R.A.-R., A.Z.M.S., M.M.M.Y.E., and M.A.R.-J. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by UAEMex, grant number 6612/2022E.

Institutional Review Board Statement

This study did not require ethical review and approval from any Educational Institution because the rumen fluid was extracted from the rumen content of animals that were slaughtered in the municipal slaughterhouse of Toluca, State of Mexico, Mexico. However, it is important to mention that said place is legally regulated by the official Mexican norm NOM-033-ZOO-1995, which establishes the methods for the humane slaughter of domestic and wild animals.

Informed Consent Statement

Not applicable.

Data Availability Statement

The data presented in this study are available on request from the corresponding author.

Conflicts of Interest

The authors declare no conflict of interest.

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Fermentation 09 00437 g001a 550Fermentation 09 00437 g001b 550
Figure 1. Ruminal total gas production kinetics with fresh forage and silage of different maize (Zea mays L.) genotypes at different incubation times, with and without the addition of guanidinoacetic acid (GAA).
Figure 1. Ruminal total gas production kinetics with fresh forage and silage of different maize (Zea mays L.) genotypes at different incubation times, with and without the addition of guanidinoacetic acid (GAA).
Fermentation 09 00437 g001aFermentation 09 00437 g001b
Fermentation 09 00437 g002 550
Figure 2. Ruminal methane (CH4) production of fresh forage and silage of different maize (Zea mays L.) genotypes at different incubation times, with and without the addition of guanidinoacetic acid (GAA).
Figure 2. Ruminal methane (CH4) production of fresh forage and silage of different maize (Zea mays L.) genotypes at different incubation times, with and without the addition of guanidinoacetic acid (GAA).
Fermentation 09 00437 g002
Fermentation 09 00437 g003a 550Fermentation 09 00437 g003b 550
Figure 3. Ruminal carbon monoxide (CO) production of fresh forage and silage of different maize (Zea mays L.) genotypes at different incubation times, with and without the addition of guanidinoacetic acid (GAA).
Figure 3. Ruminal carbon monoxide (CO) production of fresh forage and silage of different maize (Zea mays L.) genotypes at different incubation times, with and without the addition of guanidinoacetic acid (GAA).
Fermentation 09 00437 g003aFermentation 09 00437 g003b
Fermentation 09 00437 g004a 550Fermentation 09 00437 g004b 550
Figure 4. Ruminal hydrogen sulfide (H2S) production of fresh forage and silage of different maize (Zea mays L.) genotypes at different incubation times, with and without the addition of guanidinoacetic acid (GAA).
Figure 4. Ruminal hydrogen sulfide (H2S) production of fresh forage and silage of different maize (Zea mays L.) genotypes at different incubation times, with and without the addition of guanidinoacetic acid (GAA).
Fermentation 09 00437 g004aFermentation 09 00437 g004b
Table
Table 1. Chemical composition (%, on DM basis) of the forage fresh and silage of different maize (Zea mays L.) genotypes (n = 3).
Table 1. Chemical composition (%, on DM basis) of the forage fresh and silage of different maize (Zea mays L.) genotypes (n = 3).
Maize Genotype and State of the Forage 2Component 1
OMCPEENDFADFADLNSCTC
Amarillo
Fresh92.0610.802.3959.7131.683.8419.1678.87
Ensiled92.768.313.6047.5626.233.8933.2980.85
Montesa
Fresh92.7210.542.5960.6330.273.6718.9679.59
Ensiled93.268.293.9050.3526.904.1230.7281.07
Olotillo
Fresh92.7610.292.6466.2336.374.4113.6079.83
Ensiled92.998.353.8059.8236.084.8921.0280.84
Tampiqueño
Fresh92.1210.492.1961.6635.054.2517.7879.45
Ensiled93.628.563.4059.4632.314.8622.1981.66
Tuxpeño
Fresh92.0810.262.4558.8630.523.7020.5079.36
Ensiled91.639.583.6051.4928.884.2126.9578.45
1 OM: organic matter; CP: crude protein; EE: ether extract; NDF: neutral detergent fiber; ADF: acid detergent fiber; ADL: acid detergent lignin; NSC: non-structural carbohydrates; TC: total carbohydrates. 2 The pH of the silages ranged between 3.6 and 3.8.
Table
Table 2. Ruminal total gas kinetics and production, at 4, 24, and 48 h of incubation of fresh forage and silage of different genotypes of maize (Zea mays L.), with (+) and without (−) the addition of guanidinoacetic acid.
Table 2. Ruminal total gas kinetics and production, at 4, 24, and 48 h of incubation of fresh forage and silage of different genotypes of maize (Zea mays L.), with (+) and without (−) the addition of guanidinoacetic acid.
Maize GenotypeState of the Forage (SF)Guanidinoacetic
Acid (GAA)		Total Gas Production
Kinetics Parameters 1mL gas g−1 DM Incubated
bcLag4 h24 h48 h
AmarilloFresh−234.850.02455.8568.04160.52219.71
+277.200.017014.3785.72168.94227.60
Ensiled−307.450.03102.3076.40228.45295.09
+414.900.02302.6386.04241.63368.05
SEM 229.320.00211.937.2216.8923.86
SF0.02300.03960.01660.58000.01410.0106
GAA0.06300.02030.08340.13130.55750.1655
SF × GAA0.32920.91000.10070.60680.89470.2445
MontesaFresh−230.900.02559.7182.65185.99218.70
+230.350.02258.6169.31161.27210.39
Ensiled−327.700.02606.8696.63249.36306.00
+329.750.020010.3396.63211.29290.70
SEM 230.350.00202.203.4013.7024.86
SF0.03190.64830.80940.00370.01440.0280
GAA0.98150.09110.61820.12160.08380.6596
SF × GAA0.96790.50120.35770.12160.65180.8949
OlotilloFresh−226.600.021014.7990.91170.16209.33
+189.600.015020.9882.76121.45165.92
Ensiled−257.950.019510.6276.93164.44225.64
+236.700.02857.2382.23190.59226.28
SEM 29.510.00131.342.098.7310.65
SF0.01460.00930.00260.02540.02210.0228
GAA0.03760.30460.35490.53130.26610.1151
SF × GAA0.45420.00420.02320.03220.01280.1075
TampiqueñoFresh−261.100.02506.3473.12184.56243.74
+278.100.02204.7763.38167.62243.43
Ensiled−253.700.023512.1997.37200.44239.14
+214.400.023011.5077.57163.97199.12
SEM 221.560.00172.353.0214.5820.96
SF0.17450.88870.05550.00310.69640.3081
GAA0.63230.35560.65610.00810.14090.3905
SF × GAA0.26160.49750.85950.17080.53960.3972
TuxpeñoFresh−307.400.02705.0583.50229.71290.59
+281.100.02455.5775.98191.54259.63
Ensiled−263.450.02259.9885.72196.47243.16
+228.200.023011.4291.86167.09216.11
SEM 214.710.00080.607.0811.2814.28
SF0.03020.01920.00080.27040.06280.0334
GAA0.10470.27460.17690.92710.04020.1121
SF × GAA0.77620.13060.48480.38970.71670.8976
Pooled SEM 2 22.600.00161.805.0513.3419.72
p value
Maize genotype0.00090.08280.00030.13360.00150.0008
State of the forage0.00520.04160.18930.00060.00020.0017
Guanidinoacetic acid0.92840.00150.10410.39070.00570.3528
Maize genotype × State of the forage0.00010.0014<0.00010.00510.0003<0.0001
Maize genotype × Guanidinoacetic acid0.02100.00810.33620.01200.14360.1317
State of the forage × Guanidinoacetic acid0.85890.03150.17290.33430.36290.4607
Maize genotype × State of the forage × Guanidinoacetic acid0.46130.00990.04360.23960.13580.3773
1 b is the asymptotic gas total production (mL gas g−1 DM); c is the rate gas total production (mL gas h−1); Lag is the initial delay before gas total production begins (h). 2 SEM, standard error of the mean.
Table
Table 3. Ruminal methane (CH4) kinetics and production, at 4, 24, and 48 h of incubation of fresh forage and silage of different genotypes of maize (Zea mays L.), with (+) and without (−) the addition of guanidinoacetic acid.
Table 3. Ruminal methane (CH4) kinetics and production, at 4, 24, and 48 h of incubation of fresh forage and silage of different genotypes of maize (Zea mays L.), with (+) and without (−) the addition of guanidinoacetic acid.
Maize
Genotype		State of the Forage
(SF)		Guanidinoacetic
Acid (GAA)		CH4 Production
Kinetics Parameters 1mL gas g−1 DM IncubatedmL gas 100 mL−1 Gas
bcLag4 h24 h48 h4 h24 h48 h
AmarilloFresh−38.270.125512.520.34008.9738.130.50005.5017.25
+27.520.06509.860.33154.3636.850.37502.5616.19
Ensiled−58.920.117011.430.382019.1858.550.50008.5020.00
+78.710.096012.720.430012.4478.100.50005.3121.44
SEM 25.200.02761.770.07492.893.600.06251.621.79
SF0.00230.70440.64200.40160.03380.00100.37390.15050.0890
GAA0.43380.21380.71920.80510.12010.06410.37390.13160.9216
SF × GAA0.04260.51370.32620.72530.73020.04440.37390.94220.5233
MontesaFresh−42.970.122511.810.413512.4642.860.50006.6319.50
+22.700.04804.870.25409.6521.700.37506.0610.35
Ensiled−36.450.133012.240.48307.1736.290.50002.8811.63
+34.940.088011.980.24156.5134.690.25003.0611.44
SEM 210.810.02221.250.03651.7910.780.06250.932.82
SF0.80450.31910.03960.47870.07780.78030.37390.02200.2953
GAA0.37070.05460.04530.00540.38610.35060.03990.84980.1733
SF × GAA0.43450.54300.05600.32440.58030.41570.37390.70680.1874
OlotilloFresh−21.710.076010.700.22704.4719.860.25002.639.50
+25.890.118512.370.41405.5625.890.50004.5615.56
Ensiled−58.900.156513.790.38508.4058.880.50004.9425.56
+64.430.128011.520.411021.6764.090.500011.3128.19
SEM 211.370.02031.010.01023.0811.400.00001.584.03
SF0.02910.09100.32610.00160.03130.0276<0.00010.04530.0235
GAA0.69150.74770.77980.00050.08000.6476<0.00010.05790.3413
SF × GAA0.95550.15530.12160.00140.11920.9734<0.00010.23220.6914
TampiqueñoFresh−64.200.109012.290.365518.3863.780.50009.2524.63
+43.670.095511.800.15859.3743.300.25005.5017.75
Ensiled−33.170.108510.910.48706.9232.840.50003.4413.31
+27.720.148513.340.19404.4127.750.25002.6913.94
SEM 218.900.02781.730.01376.8218.780.00003.195.82
SF0.28170.39790.96440.00460.29530.2835<0.00010.24780.2639
GAA0.52960.65810.6047<0.00010.44620.5334<0.00010.51950.6200
SF × GAA0.71030.38980.44820.03470.65870.7029<0.00010.66270.5547
TuxpeñoFresh−60.040.084512.020.417515.6559.330.50007.0020.75
+64.880.166514.250.19005.4464.970.25002.8724.62
Ensiled−29.100.186013.750.42855.9628.660.50003.0611.81
+39.270.121011.390.459012.8939.140.50007.7518.12
SEM 216.990.00850.160.02774.9117.050.00002.345.83
SF0.17140.03030.02230.00720.83060.1728<0.00010.85120.2563
GAA0.68150.37500.69580.02370.75470.6612<0.00010.91030.4318
SF × GAA0.88280.00100.00010.00960.15580.8940<0.00010.13330.8447
Pooled SEM 2 13.560.02241.320.04004.2813.430.03952.084.37
p value
Maize genotype0.48420.10320.10940.11040.96200.36730.03300.94650.2759
State of the forage0.42140.01350.08820.00030.56370.48930.01040.96830.9733
Guanidinoacetic acid0.81970.16760.22780.00010.43470.9645<0.00010.82170.8531
Maize genotype × State of the forage0.00410.88290.22060.43490.02280.00520.01190.02800.0086
Maize genotype × Guanidinoacetic acid0.72630.10110.1933<0.00010.23540.6304<0.00010.16350.4339
State of the forage × Guanidinoacetic acid0.25490.35190.40500.93620.07600.33141.00000.08690.3685
Maize genotype × State of the forage × Guanidinoacetic acid0.94550.01960.02330.01150.53530.97160.00200.57930.8646
1 b is the asymptotic CH4 production (mL gas g−1 DM); c is the rate CH4 production (mL gas h−1); Lag is the initial delay before CH4 production begins (h). 2 SEM, standard error of the mean.
Table
Table 4. Ruminal carbon monoxide (CO) Kinetics and production, at 4, 24, and 48 h of incubation of fresh forage and silage of different genotypes of maize (Zea mays L.), with (+) and without (−) the addition of guanidinoacetic acid.
Table 4. Ruminal carbon monoxide (CO) Kinetics and production, at 4, 24, and 48 h of incubation of fresh forage and silage of different genotypes of maize (Zea mays L.), with (+) and without (−) the addition of guanidinoacetic acid.
Maize
Genotype		State of the Forage (SF)Guanidinoacetic
Acid (GAA)		CO production
Kinetics Parameters 1mL gas g−1 DM Incubated
bcLag4 h24 h48 h
AmarilloFresh−6.17840.02724.900.00080.25360.7511
+3.99890.00032.900.00190.08170.6090
Ensiled−3.11190.00087.550.00190.58381.4391
+3.17280.00205.810.00300.26851.7249
SEM 21.36340.00472.150.00020.03160.1129
SF0.22660.05700.26620.00850.00120.0013
GAA0.48060.05080.43370.01000.00150.5592
SF × GAA0.45740.03930.95510.91980.08550.1311
MontesaFresh−6.39180.00034.200.00100.15990.6604
+2.24040.00035.100.00190.12140.5415
Ensiled−0.69230.01714.310.00210.10260.7018
+0.96130.01096.250.00300.09280.9027
SEM 21.19290.01002.350.00010.03870.3252
SF0.04300.24160.80120.00020.32970.5694
GAA0.17900.76950.57810.00030.56680.9056
SF × GAA0.13760.77220.83590.56140.72980.6487
OlotilloFresh−1.41000.00045.070.00210.07850.2683
+0.27680.04467.950.00220.12660.5658
Ensiled−1.85190.00082.400.00190.18011.1317
+4.31470.01517.280.00280.41821.4497
SEM 21.94490.02280.990.00030.04930.0703
SF0.31360.55900.16790.60410.01630.0002
GAA0.74970.26990.01740.13430.04390.0119
SF × GAA0.40760.54770.37170.20840.12600.8917
TampiqueñoFresh−2.55600.00092.740.00110.23231.0656
+0.95720.00045.900.00150.20121.1168
Ensiled−1.05920.00347.110.00300.09900.6514
+1.50830.00106.140.00230.19150.6809
SEM 20.60030.00141.040.00030.07870.2833
SF0.47500.32660.09020.01320.41490.2079
GAA0.39250.34730.34900.67280.71670.8937
SF × GAA0.16330.51220.11770.17070.47610.9712
TuxpeñoFresh−2.30520.00525.750.00180.29711.0811
+3.14650.00376.150.00210.33301.6444
Ensiled−3.32660.00016.550.00270.14520.4791
+5.01190.01714.200.00270.22380.7056
SEM 21.83810.00240.970.00030.13060.3833
SF0.47620.16140.58640.08700.37420.1148
GAA0.52970.03190.37310.65400.68390.3610
SF × GAA0.82970.01850.23050.65400.87790.6830
Pooled SEM 2 1.46990.01141.620.00030.07510.2648
p value
Maize genotype0.12040.57110.96850.11540.03300.2576
State of the forage0.50620.77230.3500<0.00010.22510.2018
Guanidinoacetic acid0.62170.45140.33900.00040.82900.1638
Maize genotype × State of the forage0.06870.39490.28840.02540.00430.0005
Maize genotype × Guanidinoacetic acid0.55310.14150.14610.01790.01980.8052
State of the forage × Guanidinoacetic acid0.05920.86990.62690.52560.47990.7331
Maize genotype × State of the forage × Guanidinoacetic acid0.92050.43700.63750.18800.60210.8569
1 b is the asymptotic gas production (ppm CO g−1 DM); c is the rate gas production (ppm CO h−1); Lag is the initial delay before gas production begins (h). 2 SEM, standard error of the mean.
Table
Table 5. Ruminal hydrogen sulfide (H2S) kinetics and production, at 4, 24, and 48 h of incubation of fresh forage and silage of different genotypes of maize (Zea mays L.), with (+) and without, (−) the addition of guanidinoacetic acid.
Table 5. Ruminal hydrogen sulfide (H2S) kinetics and production, at 4, 24, and 48 h of incubation of fresh forage and silage of different genotypes of maize (Zea mays L.), with (+) and without, (−) the addition of guanidinoacetic acid.
Maize GenotypeState of the Forage (SF)Guanidinoacetic
Acid (GAA)		H2S Production
Kinetics Parameters 1mL gas g−1 DM Incubated
bcLag4 h24 h48 h
AmarilloFresh−0.02360.00025.900.00010.00580.0458
+0.00411.62326.040.00010.00110.0068
Ensiled−0.05900.00153.800.00010.00930.0768
+0.00570.15315.220.00010.00110.0157
SEM 20.00600.74241.840.00000.00020.0035
SF0.03610.37850.4698<0.00010.00080.0049
GAA0.00370.29800.6930<0.0001<0.00010.0001
SF × GAA0.04750.37780.7464<0.00010.00080.0358
MontesaFresh−0.40360.00273.400.00000.00070.0125
+0.06820.01952.430.00010.00050.0052
Ensiled−0.00300.16105.340.00000.00050.0057
+0.03190.00994.430.00010.00060.0047
SEM 20.15070.02241.310.00000.00010.0005
SF0.22080.02950.2057<0.00010.62130.0022
GAA0.36670.04010.5115<0.00010.62130.0013
SF × GAA0.29320.02000.9843<0.00010.34530.0038
OlotilloFresh−0.00490.18633.590.00000.00170.0062
+0.01710.00011.350.00010.00290.0356
Ensiled−0.02140.30013.930.00010.00110.0060
+0.02390.00069.650.00030.00270.0482
SEM 20.01200.15020.530.00000.00030.0015
SF0.38730.72280.00120.00750.29310.0128
GAA0.57590.18120.03000.00750.0084<0.0001
SF × GAA0.70920.72510.00170.37390.52750.0115
TampiqueñoFresh−0.02550.00095.530.00020.00100.0072
+0.06080.08624.390.00000.00040.0085
Ensiled−0.03910.00446.880.00000.00020.0045
+0.04610.00866.120.00010.00020.0041
SEM 20.02960.04173.260.00010.00040.0009
SF0.98480.42420.66050.75560.30460.0171
GAA0.51480.34340.78560.75560.47660.6489
SF × GAA0.65790.38570.95650.17090.47660.4059
TuxpeñoFresh−0.01440.07251.650.00000.00030.0072
+0.03180.25002.640.00010.00050.0221
Ensiled−0.13630.00674.910.00010.00080.0100
+0.02580.00708.250.00010.00810.0570
SEM 20.02670.12711.820.00000.00290.0055
SF0.09580.29130.07140.37390.23430.0264
GAA0.15650.52280.30000.03990.26500.0048
SF × GAA0.07480.52420.55250.37390.28180.0430
Pooled SEM 2 0.07000.34411.960.00000.00130.0030
p value
Maize genotype0.21860.43360.70790.11380.0026<0.0001
State of the forage0.41280.31420.02300.14730.1077<0.0001
Guanidinoacetic acid0.19940.36650.53120.00340.57320.0716
Maize genotype × State of the forage0.07880.45910.23500.07510.1115<0.0001
Maize genotype × Guanidinoacetic acid0.44960.21230.69850.04110.0004<0.0001
State of the forage × Guanidinoacetic acid0.60440.20610.18620.14730.41670.0561
Maize genotype × State of the forage × Guanidinoacetic acid0.15690.56500.61600.07510.1102<0.0001
1 b is the asymptotic H2S production (ppm gas g−1 DM); c is the rate H2S production (ppm gas h−1); Lag is the initial delay before H2S production begins (h). 2 SEM, standard error of the mean.
Table
Table 6. Fermentation characteristics and methane (CH4) conversion efficiency with fresh forage and silage of different genotypes of maize (Zea mays L.), with (+) and without (−) the addition of guanidinoacetic acid.
Table 6. Fermentation characteristics and methane (CH4) conversion efficiency with fresh forage and silage of different genotypes of maize (Zea mays L.), with (+) and without (−) the addition of guanidinoacetic acid.
Maize Genotype State of the Forage (SF)Guanidinoacetic Acid (GAA)Fermentation Characteristics 1CH4 Conversion Efficiency 2
pHDMDSCFAMECH4:SCFACH4:MECH4:OM
AmarilloFresh−7.0444.093.545.5072.207.549.75
+7.0937.943.735.5933.633.614.74
Ensiled−6.8850.075.055.93111.3915.1020.68
+6.9351.555.346.0869.619.6013.41
SEM 30.091.380.380.1921.242.483.12
SF0.16190.00210.01410.07390.15150.05210.0347
GAA0.61000.16500.55820.55760.13160.12960.1201
SF × GAA1.00000.05020.89500.89520.94340.76710.7352
MontesaFresh−7.1434.124.115.7586.9010.0113.44
+7.1533.883.565.4779.598.2310.40
Ensiled−6.9550.885.516.1737.665.407.69
+6.9751.304.675.7440.155.256.98
SEM 30.061.110.300.1612.181.381.93
SF0.03650.00010.01440.09460.02200.05110.0758
GAA0.84260.93930.08370.08390.85280.52270.3865
SF × GAA0.90500.78130.65260.65320.70830.58670.5796
OlotilloFresh−7.3833.603.765.5434.453.754.82
+7.2535.912.684.9860.015.186.00
Ensiled−7.1040.033.635.2164.807.399.03
+7.0944.534.215.51148.3618.2623.31
SEM 30.030.910.190.1020.692.533.31
SF0.00300.00110.02210.38280.04550.02990.0314
GAA0.11670.01980.26650.26470.05770.07190.0800
SF × GAA0.13830.29310.01280.01280.23360.13570.1193
TampiqueñoFresh−7.2839.984.085.73121.3014.4719.95
+7.2035.663.705.5472.187.8210.17
Ensiled−7.0644.244.435.6545.075.687.39
+7.0841.223.625.2335.283.924.71
SEM 30.090.910.320.1741.825.247.41
SF0.12320.00570.69630.31030.24760.29220.2908
GAA0.74830.01580.14060.14100.52000.46720.4476
SF × GAA0.56270.51530.53960.54090.66270.66490.6567
TuxpeñoFresh−7.0941.145.086.2191.7411.8417.00
+7.1034.134.235.7837.714.395.91
Ensiled−7.0239.364.345.7440.164.846.51
+7.1042.613.695.41101.7211.1014.07
SEM 30.051.230.250.1330.733.815.33
SF0.48390.05260.06270.03060.84960.97130.8375
GAA0.37780.20070.04010.04020.90840.88270.7572
SF × GAA0.42820.01400.71680.71720.13320.14620.1553
Pooled SEM 3 0.071.120.300.1527.283.364.64
p value
Maize genotype0.0015<0.00010.00150.00060.94600.95580.9604
State of the forage0.0301<0.00010.00020.40890.97090.52630.5823
Guanidinoacetic acid0.94750.09540.00570.00570.82440.57050.4344
Maize genotype × State of the forage0.0022<0.00010.00030.00280.02800.02120.0233
Maize genotype × Guanidinoacetic acid0.69260.00240.14360.14390.16300.19650.2400
State of the forage × Guanidinoacetic acid0.86930.00030.36260.36260.08710.07610.0756
Maize genotype × State of the forage × Guanidinoacetic acid0.75680.02200.13580.13640.57940.52690.5363
1 pH is the ruminal pH; DMD is dry matter degradability (%); SCFAs are short-chain fatty acids (mmol g−1 DM); ME is the metabolizable energy at 24 h (MJ kg−1 DM). 2 CH4:ME is the methane:metabolizable energy ratio (g MJ−1); CH4:OM is the methane:organic matter ratio (mL g−1); CH4:SCFA is the methane:short-chain fatty acid ratio at 24 h (mmol mmol−1). 3 SEM, standard error of the mean.
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MDPI and ACS Style

Alvarado-Ramírez, E.R.; Andrade-Yucailla, V.; Elghandour, M.M.M.Y.; Acosta-Lozano, N.; Rivas-Jacobo, M.A.; López-Aguirre, D.; Garay-Martínez, J.R.; Vazquez-Mendoza, P.; Barros-Rodríguez, M.; Salem, A.Z.M. Effects of Guanidinoacetic Acid on Ruminal Fermentation and Greenhouse Gas Production Using Fresh Forage and Silage from Different Maize (Zea mays L.) Genotypes. Fermentation 2023, 9, 437. https://doi.org/10.3390/fermentation9050437

AMA Style

Alvarado-Ramírez ER, Andrade-Yucailla V, Elghandour MMMY, Acosta-Lozano N, Rivas-Jacobo MA, López-Aguirre D, Garay-Martínez JR, Vazquez-Mendoza P, Barros-Rodríguez M, Salem AZM. Effects of Guanidinoacetic Acid on Ruminal Fermentation and Greenhouse Gas Production Using Fresh Forage and Silage from Different Maize (Zea mays L.) Genotypes. Fermentation. 2023; 9(5):437. https://doi.org/10.3390/fermentation9050437

Chicago/Turabian Style

Alvarado-Ramírez, Edwin Rafael, Verónica Andrade-Yucailla, Mona Mohamed Mohamed Yasseen Elghandour, Néstor Acosta-Lozano, Marco Antonio Rivas-Jacobo, Daniel López-Aguirre, Jonathan Raúl Garay-Martínez, Paulina Vazquez-Mendoza, Marcos Barros-Rodríguez, and Abdelfattah Zeidan Mohamed Salem. 2023. "Effects of Guanidinoacetic Acid on Ruminal Fermentation and Greenhouse Gas Production Using Fresh Forage and Silage from Different Maize (Zea mays L.) Genotypes" Fermentation 9, no. 5: 437. https://doi.org/10.3390/fermentation9050437

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