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Revista mexicana de ciencias pecuarias

versão On-line ISSN 2448-6698versão impressa ISSN 2007-1124

Rev. mex. de cienc. pecuarias vol.7 no.2 Mérida Abr./Jun. 2016

 

Articles

Yield, and ruminal dry matter and energy degradability of ten tropical grasses harvested at four ages

Braulio Valles de la Moraa 

Epigmenio Castillo Gallegosa 

Hugo Bernal Barragánb  * 

aUniversidad Nacional Autónoma de México, Facultad de Medicina Veterinaria y Zootecnia, CEIEGT. Martínez de la Torre, Veracruz, México.

bUniversidad Autónoma de Nuevo León, Facultad de Agronomía, Gral. Escobedo, N.L., México.


Abstract

The objective of the study was to determine dry mater yield (DMY), and in situ ruminal degradable dry matter and energy of ten grasses of the genera Brachiaria: B. brizantha (Insurgente), B. decumbens (Señal), B. humidicola (Chetumal), B. brizantha x B. ruzieziensis (Mulato I); Panicum: P. maximum cv. Mombasa, P. maximum cv. Guinea, P. maximum cv. Tanzania; and, Pennisetum: P. purpureum cv. Taiwan, P. purpureum cv. Cuban king grass, P. purpureum cv. Purple king grass, at 3, 6, 9 and 12 wk of regrowth, in three evaluation cycles. DMY for Pennisetum, Brachiaria and Panicum was: 3,037, 1,689 and 1,872 kg ha-1 (P<0.05). CP concentration decreased from 11.6 to 6.9 % (3 to 12 wk), without differences among genera. Concentration (%) of NDF, ADF and LIG were: 72.1, 43.5 and 8.1 (Brachiaria); 73.0, 45.8 and 9.0 (Panicum); and 68.8, 44.4 and 9.1 % (Pennisetum), values were 68.1, 40.1 and 8.2 (at 3 wk); and 76.2, 49.7 and 9.5 (at 12 wk). The in situ DM degradability was: 70.0 % (Brachiaria), 65.8 % (Panicum) and 72.9 % (Pennisetum) and decreased from 71.7 to 62.9 % (from 3 to 12 wk). Gross energy (GE) pre-incubation, was 3,802, 3,756, and 3,608 kcal kg-1 DM, for Brachiaria, Panicum, and Pennisetum, with no changes due to age of regrowth. In situ ruminal energy degradability (as % of incubated GE) was: 55.6, 51.4, and 57.9 % for Brachiaria, Panicum; and Pennisetum. In conclusion, age of regrowth and genus affected the nutritive value of the evaluated grasses.

Key words: Tropical grasses; Regrowth age; Ruminal degradability; Dry matter

Resumen

El presente trabajo se realizó con el objetivo de determinar el rendimiento de materia seca (RMS), y la degradabilidad ruminal de la materia seca (MS) y la energía, de diez pastos de los géneros Brachiaria: B. brizantha (Insurgente), B. decumbens (Señal), B. humidicola (Chetumal), B. brizantha x B. ruzieziensis (Mulato I); Panicum: P. maximum cv. Mombasa, P. maximum cv. Guinea, P. maximum cv. Tanzania; y Pennisetum: P. purpureum cv. Taiwan, P. purpureum cv. Cuban king grass, P. purpureum cv. Purple king grass, a 3, 6, 9 y 12 semanas de rebrote, en tres ciclos de crecimiento. La degradabilidad ruminal de MS y energía se determinó a 48 h de incubación in situ. El RMS fue 3,037, 1,689 y 1,872 kg ha-1, para Pennisetum, Brachiaria y Panicum (P<0.05). La PC disminuyó de 11.6 a 6.9 % (3 a 12 semanas), sin diferencias entre géneros. Contenidos (%) de FDN, FDA y LIG fueron 72.1, 43.5 y 8.1 (Brachiaria); 73.0, 45.8 y 9.0 (Panicum); y 68.8, 44.4 y 9.1 (Pennisetum); así como 68.1, 40.1 y 8.2 (3 semanas); y 76.2, 49.7 y 9.5 (12 semanas). La degradabilidad in situ de MS fue 70.0 (Brachiaria), 65.8 (Panicum) y 72.9 % (Pennisetum); disminuyendo de 71.7 a 62.9 % de 3 a 12 semanas. La energía pre-incubada promedió: 3,802, 3,756, y 3,608 kcal kg-1 MS, para Brachiaria, Panicum, y Pennisetum, sin cambios por edad de rebrote. La degradabilidad (% de la EB incubada in situ en el rumen) fue 55.6, 51.4 y 57.9, para Brachiaria, Panicum y Pennisetum. En conclusión, la edad de rebrote y género afectaron el valor nutritivo de los pastos evaluados.

Palabras clave: Pastos tropicales; Edad de rebrote; Degradabilidad ruminal; Materia seca; Energía

INTRODUCTION

In the tropical regions, the most extended cattle production is the dual purpose system, based on grazing of native pastures1, whose productivity varies in quality and quantity2. It is thereafter important to effectively select and to introduce grasses with high potential for forage yield and nutritional quality, adapted to the local conditions of clime, soils and management3. Age of regrowth at harvest affects quality and digestibility of grass pastures4,5, due to an increase in the proportion of cell wall, and dead material of the plant as maturity increases. Grasses with low digestibility slow the passage of feed through the animal's digestive system reducing intake levels.

Digestibility of pasture is directly related to its energy and protein content, and influences animal performance6,7,8, being low quality forages generally deficient in digestible protein and energy9,10. Protein supply is limiting in cattle consuming tropical forages with less than 7 % crude protein content11, this affecting intake and digestibility, and animal productivity12. However, surveys dedicated to characterize energy supply and in situ degradability of dry matter of tropical grasses are scarce. In the present study, in situ degradability of dry matter and energy of tropical forages harvested at different ages, and growing under diverse climatic conditions in different seasons of the year, was assessed.

MATERIAL AND METHODS

The experiment was carried out at the Centro de Enseñanza, Investigación y Extensión en Ganadería Tropical of the Facultad de Veterinaria of the Universidad Nacional Autónoma de Mexico (CEIEGT, FMVZ-UNAM), located in Tlapacoyan, Veracruz, Mexico, at an altitude of 114 m. Soils are acidic (pH 4.5 to 5.2), clay-silt Ultisols (durustults), low in available P (3.5 ppm, Bray II) and cation exchange capacity (10.5 meq 100 g-1). Soil has a 25 cm deep hardpan, causing a deficient drainage during rainy and winter seasons. In the dry season, water available to plants is rapidly depleted, because the soil does not hold much moisture. Climate is hot and humid, Af(m)w"(e)13. Yearly average daily temperature is 23.9 ± 0.5 °C, and rainfall is 1931 ± 334 mm. Figure 1 shows rainfall and temperatures registered during the three phases of the experimental period: first, from March 6 to May 29, 2008 (Tmin= 22.2, Tmax= 32.6 °C, 342 mm); second, from August 1 to October 24, 2008 (Tmin= 18.1, Tmax= 29.6 °C, 1,025 mm); and third, from January 30 to April 24, 2009 (Tmin= 18.2, Tmax= 30.2 °C, 217 mm). Rainfall in the first and third cycles was 33 and 20 %, respectively, of the rainfall registered during the second cycle.

Figure 1 Climatic conditions during the experimental period 

Experimental grass plots (5.0 x 2.0 m; 0.5 m distance among furrows) were planted on June 27, 2007, utilizing rooted tillers, without fertilizer. To standardize plant regrowth, grass was cut down on the three following start dates, of each growth cycle: March 6, 2008, August 1, 2008, and January 30, 2009. Forage was harvested with "machete" at cutting heights of 10 cm for decumbent species of Brachiaria: Insurgente, Mulato, Chetumal, Chontalpo; and 20 cm for bunch-type species of P. maximum: Tanzania, Guinea, Mombaza; and erect-type species of Pennisetum: Taiwan, King grass and Purple King grass. In each experimental plot, 1 m2 subplots located within the two central furrows was cut down at 3, 6, 9 y 12 wk of regrowth. Fresh forage was weighed, and dried at 65 °C for 72 h in order to estimate dry matter yield (DMY).

Dried samples were ground on a Wiley #4 mill to pass a 1 mm screen. Duplicates of ground forage were analyzed for crude protein (CP) by the normal (macro) Kjeldahl procedure14, neutral detergent fiber (NDF), acid detergent fiber (ADF) and acid detergent lignin (ADL)15, using an ANKOM2000 Fiber Analyzer (Ankom Technology Corp., Macedon, NY). Concentrations of hemicellulose (NFD - ADF) and cellulose (ADF - Lig)15 were calculated. In situ DM disappearance after 48 h of ruminal incubation (ISDM), was also determined in triplicate on each of three fistulated cows16 after 48 h of ruminal incubation using the formula: {[Incubated sample (g) - Residue (g)]/ Incubated sample (g)}*100.

Gross energy (GE, kcal/kg DM) content of oven-dried forage samples was determined using adiabatic bomb calorimeter (Parr Oxygen Bomb Calorimeter, Parr Instrument Co. Inc., Moline, IL.), using benzoic acid as standard. GE values of samples before and after 48 h of ruminal incubation, were used to estimate ruminal degradable energy content (DegE, expressed in kcal kg-1 DM, using the following equation:

DegE=  [IS x GE of MI- RM x GE of RM]IS

Where: IS, is incubated sample (g DM); RM, is residue of sample after incubation (g DM); y GE= Gross energy (cal/g DM).

Statistical analysis. Results of dry matter yield were analyzed for each cycle utilizing a randomized complete block experimental design, with three blocks (B) as replicates, in order to consider the effect of slope of the plot. Treatments were distributed in a split-plot arrangement, with grass (G) as main plot and age of regrowth (A) as subplot17, according to the following model:

Yijkl = μ + Bj + Gk + (B*G)jk + Al + (G*A)kl + Rijkl

Where, B, G and A are main effects for block, grass (main plot) and regrowth age (subplot), respectively. The interaction B*G is the error in "a", and indicates if the response of each genus is the same for each block; the interaction G*A indicates if the response to regrowth age is the same for each grass. R is the residual variation in "b", and is used to test effects of the subplot A and G*A. Some of the quality variables of the second cycle (CP, NDF, ADF, LIG and ISD) had been previously reported18, however they are presented here in a new approach together with the first and third cycles.

Orthogonal contrasts were also developed to compare genera, and grasses within genus. In the case of in situ degradability (ISD) of the energy, the effect of block was confounded with the effect of the three cows used for the in situ determination. The analysis of variance was performed with PROC GLM of SAS19. Values of DMY (y, kg/ha) were adjusted to the exponential equation y= aebx, where "a" estimates the amount of residual DM left after the standardization cut, "e" is the basis of natural logarithms and "bx" is the relative growth rate expressed as kg DM/kg DM present/day. From this value, average time needed to duplicate the yield was calculated as ln(2)/b, whereas the slope was calculated as b*y20. Data adjustment to the models was made utilizing the GraphPad Prism v5.04 software for Windows (GraphPad Software, San Diego California USA)21. Analysis of variance for other variables was performed using SPSS program22, and mean comparisons were done using Tukey test at P=0.05 level.

RESULTS

Dry matter yield (DMY)

Differences in DMY by genera of grasses is shown in Table 1, with highest (P<0.05) differences between Brachiaria y Penisetum in the first (P=0.0021) and in the second (P=0.0109) cycle. Panicum y Pennisetum were different only in the first cycle (P=0.0331). During the first cycle, orthogonal contrasts for grasses among genera indicated that King grass (5,582 kg ha-1) was different (P<0.05) from Taiwan and Purple King Grass (4,606 y 4,494 kg ha-1). During the second and third cycles, DMY was similar (P>0.05). DMY varied in the second cycle from 1,788 until 2,136, and in the third cycle, from 893 until 1,889 kg ha-1. During the three periods grasses of genus Brachiaria showed the lowest DMY.

Table 1 Least square means of dry matter yield by grass and by genus, corrected to 7.5 weeks of regrowth 

Evaluation cycle: First Second (kg ha-1) Third
Group:
Brachiaria 2334 b 1788 944
Panicum 2848 b 1997 893
Pennisetum 4894 a 2136 1889
SE 203 295 279
P 0.0016 0.6875 0.1040

SE =Standard error; P= Calculated probability.

ab Values of genus with distinct letters are different (P<0.001).

Parameters of the growth exponential model are shown in Table 2. For genus Pennisetum the parameter "a" had the highest variation, with values ranging from 17.5 to 695.9 kg ha-1 (cycles 3 to 1, respectively). The relative growth rates (parameter "b") were less variable, excepting Pennisetum spp (cycle 3) whose value was 2.5 times higher than other values. Variabilities in "a" and "b" suggest an interaction according to the week and the growth cycle. The time to duplicate yield (DMY) varied from 1.5 wk (Pennisetum spp, cycle 3) to 4.6 wk (Brachiaria spp, cycle 2). Growth rates (TC7.5) at 7.5 wk were highly variable: from 127 kg ha-1 wk-1 (18.1 kg ha-1 d-1) for Panicum spp (cycle 3), until 873 kg ha-1 wk-1 (124.7 kg ha-1 d-1) for Pennisetum spp (cycle 1). Determination coefficient R2 varied from 0.46 to 0.85, and F Test was highly significant (P<0.0001) in all the cases.

Table 2 Exponential response type y= aebx, for dry matter yield (y, kg ha-1) according to an increase in regrowth age (x, weeks) in three genera of tropical grasses 

Cycle Genus DF Parameters Derived variables Goodness of fit
a b SDR TC7.5 R2 Sy.x
1 Brachiaria spp 46 426.2 0.200 3.5 383 0.76 898.0
Panicum spp 34 605.5 0.185 3.7 451 0.61 1539.0
Pennisetum spp 34 695.9 0.228 3.0 873 0.71 2489.0
2 Brachiaria spp 46 519.4 0.149 4.6 238 0.70 589.6
Panicum spp 34 316.0 0.213 3.3 331 0.85 548.6
Pennisetum spp 34 231.3 0.251 2.8 383 0.63 1252.0
3 Brachiaria spp 46 188.2 0.188 3.7 145 0.60 454.5
Panicum spp 34 219.8 0.166 4.2 127 0.67 334.8
Pennisetum spp 34 17.52 0.473 1.5 289 0.46 2084.0

DF= Degrees of freedom of the residual. The parameters of the exponential equation are: 'a', Intercept; 'b', relative growth rate kg/kg/wk. The derlvate variables are: 'SDR', weeks to duplicate the yield and TC7.5, slope of the curve or relative growth rate at 7.5 wk of age in kg/ha/wk. The goodness of fit: 'R2', determination coefficient; Sy.x, standard deviation of the nonlinear regression.

Chemical composition

Table 3 shows means for the evaluated quality variables. Purple King grass had the lowest values (P<0.01) of NDF and hemicellulose, although Chetumal and Guinea had the highest values for NDF; and Chetumal had the highest value for hemicellulose. Panicum and Pennisetum showed the highest CP concentrations (>10 %). Even though in situ dry matter degradability was similar among genera, grasses of genus Panicum showed the lowest values, in average 67.1 %.

Table 3 Nutritional composition of ten tropical grasses in a hot and humid climate in Veracruz, Mexico (%) 

Genus grass NDF ADF Lig Hemi Cel CP ISDMD, EDeg, % EDeg, cal
Brachiaria:
Insurgente 71.1 ab 42.8 8.2 28.3 bc 34.6 8.9 c 70.0 56.3 2140
Mulato 69.9 ab 41.4 8.6 28.5 bc 32.8 8.4 c 71.6 55.8 2084
Chetumal 75.4 a 43.1 7.5 32.3 a 35.6 9.1 b 68.7 53.0 2050
Chontalpo 72.1 ab 42.0 8.3 30.1 ab 33.7 8.6 c 71.0 56.5 2168
Panicum:
Tanzania 72.6 ab 44.9 8.5 27.6 bcd 35.5 10.6 ab 68.4 50.3 1879
Guinea 73.7 a 47.1 9.3 26.6 bcde 37.7 10.3 ab 65.3 50.0 1938
Mombasa 72.7 ab 45.3 9.3 27.4bcd 36.1 10.2 ab 67.7 51.9 1933
Pennisetum:
Taiwan 79.5 ab 44.5 8.8 25.0 cde 35.7 11.0 a 72.1 59.9 2198
Cuban KG 70.3 ab 44.5 9.4 24.7 de 36.1 10.2 ab 73.5 59.6 2149
Purple KG 66.7 b 43.2 9.0 23.6 e 34.2 11.3 a 73.9 54.9 1951

NDF= neutral detergent fiber; ADF= add detergent fiber; Llg= llgnln; Heml= hemlcellubse; Cel= cellubse; CP= crude protein; ISDMD= in situ dry matter digestibility; EDeg,%= rumlnal in situ degradabe energy expressed as proportion of the incubated energy. EDeg, cal= expressed as calorie.

abcdeMeans with different letter in a column by each genus, are significant (P<0.0001).

Table 4 shows that Brachiaria had the lowest value for ADF (cycle 1). Among ages of regrowth, this value increased according to the time. Cycles 2 and 3 also showed a genus x age of regrowth interaction (P<0.05). In the first cycle, lignin concentration was similar among genera and among ages of regrowth, but this was different in the other two ones. Cycles 2 and 3 showed a genus x age interaction (P<0.05). Regarding hemicellulose, its content in Brachiaria (cycles 1 and 2) was higher than in Pennisetum (P<0.05), but in cycle 3 there was a genus x age interaction (P<0.05). At 12 wk, the content of hemicellulose was similar for all three genera. The concentration of cellulose was higher for Panicum (cycles 1 and 2) than for the other genera; whereas regarding ages, the concentration of cellulose was higher (P<0.05) at 12 wk in the cycles 1 and 2, but in cycle 3, the average value of cellulose at 6 wk was higher (P<0.05). Content of CP varied among genera from 7.6 to 11.9 %; with significant differences in cycles 2 and 3 (P<0.001). For ages, the concentration of CP was also statistically significant (P<0.001) in all the three cycles, with a reduction in the values from wk 3 to wk 12. Low values of CP were registered in the second cycle, by genera and by ages.

Table 4 Comparison of means among genera and regrowth ages for characteristics of chemical composition (%) of three genera of tropical grasses cultivated in conditions of hot and humid climate in Veracruz, Mexico 

Genus Regrowth ages (weeks) Interaction
Brach Pani Penni SEM P 3 6 9 12 SEM P SEM P
First Cycle
NDF 71.9 ab 73.9 a 69.8 b 0.793 0.005 64.4 c 70.2 b 74.5 a 78.4 a 0.793 0.0001 1.585 0.818
ADF 41.3 b 44.7 a 43.3 ab 0.634 0.001 33.8 c 42.8 b 45.1 b 50.6 a 0.7 0.0001 1.267 0.614
LIG 8.9 a 8.6 a 9.5 a 0.404 0.279 9.3 a 9.7 a 8.1 a 9.0 a 0.447 0.097 0.809 0.086
Hemi 30.7 a 29.2 a 26.5 b 0.51 0.0001 30.6 a 27.4 b 29.4 ab 27.8 b 0.564 0.001 1.020 0.205
Cel 32.4 b 36.0 a 33.8 ab 0.68 0.002 24.5 d 33.1 c 37.1 b 41.5 a 0.752 0.0001 1.360 0.756
CP 9.8 a 11.4 a 11.3 a 0.88 0.337 13.2 a 12.7 a 8.6 b 8.3 b 0.638 0.001 1.158 0.149
Second cycle
NDF 72.9 a 73.8 a 68.2 b 0.776 0.0001 69.8 b 68.8 b 71.6 b 76.3 a 0.858 0.0001 1.552 0.823
ADF 42.8 b 47.5 a 44.0 b 0.431 0.0001 41.3 c 41.0 c 44.6 b 52.1 a 0.476 0.0001 0.862 0.003
LIG 6.7 b 8.4 a 7.7 ab 0.236 0.0001 7.0 b 7.2 b 7.1 b 8.9 a 0.261 0.0001 0.472 0.004
Hemi 30.1 a 26.4 b 24.2 b 0.659 0.0001 28.5 a 27.8 ab 27.0 ab 24.2 b 0.728 0.002 1.317 0.278
Cel 36.1 b 39.1 a 36.4 b 0.378 0.0001 34.3 c 33.8 c 37.4 b 43.2 a 0.418 0.0001 0.756 0.216
CP 7.6 c 8.7 b 9.6 a 0.274 0.003 9.8 a 9.7 a 8.7 b 6.0 c 0.270 0.001 0.508 0.054
Third cycle
NDF 71.5 a 71.3 a 68.5 a 0.945 0.063 69.6 ab 67.3 b 71.0 ab 73.8 a 1.044 0.001 1.889 0.825
ADF 42.9 a 45.3 a 45.9 a 0.727 0.011 45.4 a 39.7 b 46.7 a 47.0 a 0.804 0.0001 1.455 0.032
LIG 8.9 a 10.1 a 10.0 a 0.320 0.009 8.3 b 8.1 b 11.4 a 10.9 a 0.354 0.0001 0.640 0.029
Hemi 28.6 a 26.0 a 22.6 b 0.751 0.0001 24.2 a 27.6 a 24.3 a 26.9 a 0.830 0.012 1.501 0.034
Cel 34.1 a 35.1 a 35.9 a 0.673 0.142 37.1 a 31.6 b 35.3 a 36.1 a 0.744 0.0001 1.346 0.423
CP 8.9 b 11.9 a 11.2 a 0.394 0.001 12.5 a 12.4 a 10.8 b 6.4 c 0.431 0.001 0.787 0.003

Brach- Brachiaria, Pani- Panicum, Penni= Pennisetum; SEM= standard error of the mean; NDF= neutral detergent fiber; ADF= acid detergent fiber; Llg= llgnln; Heml= hemlcellulose; Cel= celluOse; CP= crude proteln; P= probablllty of error P>F.

abc Means ln a row wlth slmllar letter are not dlfferent (P<0.05).

In situ degradation of dry matter and energy

In cycles 1 and 2, the in situ degradability of dry matter of grasses of genus Panicum was less (P<0.0001) than those of grasses Brachiaria and Pennisetum (Table 5). This variable decreased (P<0.0001), in all the three cycles, along with an increase in age at harvest. The degradability of samples harvested at 3 wk was higher (81.0 %; P<0.05) than when grasses were cut at 12 wk (67.0 %). For second and third cycle, there was a significant genus x age interaction.

Table 5 Gross energy (GE, kcal kg-1 DM) before and after incubation, in situ ruminal dry matter degradability (ISDMD, %) and degradable energy (DegE, % and cal) of ten tropical grasses grouped by genus, and regrowth ages 

Genus Regrowth ages (weeks) Interaction
Brach Pani Penni SEM P 3 6 9 12 SEM P SEM P
First cycle
GEpre 3779 a 3753 a 3604 b 31.8 0.001 3742 a 3714 a 3731 a 3661 a 35.1 0.378 63.46 0.670
GEpost 7213 a 6341 a 6675 a 357.0 0.318 7598 a 6548 a 6978 a 6268 a 488.9 0.222 757.4 0.499
ISDMD 74.2 a 63.4 b 76.1 a 1.641 0.0001 81.0 a 73.3 ab 69.5 b 67.0 b 75.14 0.002 2.631 0.339
ResDM 25.8 b 36.6 a 23.9 b 1.641 0.0001 19.0 b 26.7 ab 30.5 a 33.0 a 1.641 0.002 3.222 0.339
DegE,% 51.5 ab 39.6 b 55.4 a 4.213 0.098 61.8 a 53.0 a 43.0 a 44.9 a 5.341 0.109 5.341 0.191
DegE,cal 1953 a 1468 a 2002 a 156.26 0.126 2309 a 1957 a 1607 a 1639 a 156.3 0.094 156.3 0.277
Second cycle
GEpre 3900 a 3836 a 3671 b 28.01 0.0001 3883 a 3733 b 3783 ab 3810 ab 3798 0.016 56.03 0.432
GEpost 3942 a 3914 a 3762 b 24.36 0.0001 3879 a 3786 b 3841 b 3984 ab 26.93 0.0001 48.72 0.019
ISDMD 65.1 b 59.7 c 69.3 a 0.647 0.0001 68.5 a 70.0 a 66.6 a 53.8 b 68.04 0.0001 1.295 0.014
ResDM 34.9 b 40.3 a 31.0 c 0.647 0.0001 31.5 b 30.0 b 33.4 b 46.2 a 0.716 0.0001 1.295 0.014
DegE,% 64.6 b 58.8 c 68.4 a 0.686 0.0001 68.5 a 69.5 a 66.0 a 51.7 b 67.42 0.0001 1.371 0.011
DegE,cal 2519 a 2255 b 2498 a 31.79 0.0001 2658 a 2592 ab 2478 b 1967 c 35.14 0.0001 63.57 0.027
Third cycle
GEpre 3727 a 3681 ab 3550 b 35.54 0.003 3648 a 3587 a 3717 aa 3661 a 39.29 0.162 71.09 0.801
GEpost 6592 a 6767 a 6723 a 262.6 0.86 7193 a 6885 a 6520 a 6179 a 297.6 0.085 606.6 0.115
ISDMD 71.7 a 73.8 a 73.3 a 1.242 0.425 70.0 bc 76.2 ab 77.9 a 67.7 c 1.493 0.0001 3.043 0.006
ResDM 28.3 a 26.2 a 26.7 a 1.242 0.425 30.0 ab 23.8 bc 22.1 c 32.3 a 1.493 0.0001 3.043 0.006
DegE,% 49.2 a 52.1 a 49.3 a 2.912 0.714 40.1 b 54.1 ab 61.0 a 45.5 ab 3.299 0.001 6.724 0.071
DegE,cal 1831 a 1928 a 1759 a 114.13 0.585 1470 b 1939 ab 2276 a 1672 b 129.3 0.001 263.57 0.136

Brach= Brachiaria; Pani= Panicum; Penni= Pennisetum; SEM= standard error of mean; P= significance P>F; GEpre= gross energy before incubation; GEpost= gross energy post incubation, ISDMD= in situ ruminal dry matter degradability (%); ResDM= residual dry matter, DegE,%= degradaba energy expressed in terms of %. DegE,cal= degradaba energy expressed in terms of calories (cal).

abc Means in a row with same letter are not different (P<0.05).

GE content of the samples before ruminal incubation showed significant differences among genera in the three evaluated cycles (P<0.003). Genus Pennisetum had the lowest values, averaging 3,608 kcal kg-1 DM, compared to Brachiaria and Panicum, which averaged 3,802 y 3,757 kcal kg-1 DM, respectively. By ages, only in the second cycle there were significant differences (P<0.016). The GE values of samples after the ruminal incubation was statistically significant only in the second cycle, for genera, ages and its interaction.

Ruminal energy degradability (kcal kg-1 DM) was higher for Panicum in the second cycle, and was different by ages of harvesting during the second and the third cycles (P<0.0001). In the second cycle, grass harvested at 3 wk had a higher content of ruminal degradable energy (2,658 kcal kg-1 DM) than in other ages (P=0.001). In the third cycle, grass harvested at 9 wk had a higher ruminal degradable energy content (2,276 kcal kg-1 DM; P<0.001), than in others harvest ages.

DISCUSSION

Dry matter yield

The seasonality of forage production, a common fact under tropical conditions, leads to fluctuations in the production23 and quality of forage throughout the year. In the present study, grasses of Genus Pennisetum, (erect growth habit), were different compared with species of Brachiaria and Panicum. Similar facts were observed by Valerio et al24. They found that erect-type growing grass species Merkeron grass, Tanzania and Guinea grass, performed better than decumbent ones (B. humidicola, Cynodon spp and Digitaria spp), since erect-type grasses accumulate more forage, as a function of a higher size and leaves area.

In the state of Guerrero, Mexico, Peralta et al25 found significant grass species x age of regrowth interactions in six cultivars of Brachiaria spp (Toledo, Insurgente, Señal, HBA-4062, HBA-2094, Mulato) and two of Panicum (Tanzania and Mombasa), and in regrowth ages of 3 to 12 wk (P<0.01). They mentioned that, cultivars Tanzania and Mombaza had higher yields than those achieved by Brachiaria. In the present study, in the first cycle (dry season), Pennisetum yielded more DM than the other grass.

The exponential growth model described properly the quantitative increase in aerial biomass throughout the time. The positive intersection determined by the model is an estimator of the biomass residue left after the standardization cut (week= 0) at the beginning of each cycle. Parameter "c" estimates the efficiency by which the present dry matter produces more dry matter. This parameter predicts that for every kilo of DM present produced between 149 and 473 g of new biomass per week. This means that between 1.5 and 4.6 wk are needed in order to duplicate the amount of grass harvested. Growth rate (kg ha-1 d-1) is the tangent of the curve in a determined point, which resulted in values varying from 127 to 873 kg ha-1 wk-1, equivalent to 18.1 and 124.7 kg ha-1 d-1 (Table 2). These estimates of growth rates are higher than those measured in the same Experimental Station in several grasses cultivated in small plots, whose values (average of 4 yr) varied from 24.6 to 52.1 kg ha-1 d-1 26, and are clearly higher than those obtained in the same place for native grasses in intensive rotational grazing, with yearly average of 13.7 ± 12.8 kg(27X It is considered that growth differences between plants in small plots and grasslands are due to microenvironments originated by the different managements that are applied to the plants as a consequence of the grazing activity of the animals. In the present work, the genus Pennisetum spp, showed the highest growth during cycle 3, with growth rate at 7.5 wk, of 873 kg ha-1 wk-1, 2.5 times superior than the average of all other genera. The capacity of the species of this genus to produce huge quantities of forage has been verified in studies realized in the tropics of Mexico28 and in the world29.

Considering the dry matter yields for the three evaluation cycles (Table 2), mean annual yields ranged from 13.9 to 29.8 DM t ha-1. Averages by genus were: 15.4 (Brachiaria), 18.7 (Panicum) and 35.5 (Pennisetum) DM t ha-1. In Brazil, Aparecida et al30 reported a range of 43.2 to 66.4 DM t ha-1. For three grasses of genus Brachiaria grasses (B. decumbens, cv Basilisk; B. brizantha, cv Marandu; B. brizantha, cv Xaraes) and two Panicum grasses (P. maximum, cv Mombaza; P. maximum, cv Tanzania), harvested monthly during 10 mo, with application of 45 and 80 kg ha-1 of N and KCl, respectively, after every harvest, and irrigation when needed. In Thailand29 reported significant differences between Pennisetum cultivars, Napier (31.3 DM t ha-1), Merkeron (28.0 DM t ha-1) and Tangashima (30.3 DM t ha-1) on the one side, and Dwarf Napier (26.1 DM t ha-1) and Taiwan A-25 (26.1 DM t ha-1). These findings agree with results of the present study for annual dry matter for Taiwan, but were lower for Cuban King grass and for Purple King grass, with values of 29.8, 41.4 and 35.5 DM t ha-1, respectively.

Chemical composition

Content of NDF for Brachiaria and Panicum (71.5 to 73.9 %), was higher than for Pennisetum (68.2 to 69. 8 %). Hare et al31,32, reported that, NDF of Brachiaria (cultivars Mulato II, Cayman and line BRO2/1794) and Panicum (Mombaza and Tanzania) grasses evaluated at 4, 6, 9 and 12 wk, and varied in ranges from 51.9 to 63.3 % and 62.3 to 68.0 %, respectively. In the present study, content of NDF increased by increasing cutting age from 3 to 12 wk, 21.7 % (first cycle), 9.3 % (second cycle) and 6.0 % (third cycle). This variation is less than that found by previously cited researchers31,32.

The highest ADF content registered in the second and third cycle, were probably due to the fact that stems are building a major proportion of structural tissue, high in fiber content. Hare et al31,32 reported lower values for ADF content on the dates Brachiaria (26.2 to 31.6 %) and Panicum (34.6 to 40.1 %) grasses; and probably differences were due, to fertilizer effect.

The lowest content of lignin was determined during the second cycle, which had the lowest temperatures, (18.1 to 29.6 °C); and 1,025 mm of rainfall. However even these low values, are still higher than those reported by others33, for three Brachiaria brizantha cultivars (Marandu, Piata and Xaraes), (average 3.35 %). Differences in lignin content were found between leaves (3.2 to 3.6 %) and stems (4.6 to 5.9 %) in these grasses. Pastures harvested at early ages, normally showed high quality values, because at this growth stage (vegetative stage), the effect of the climate is not still detrimental to plant quality34. Rodrigues et al35 reported lignin content lower than values reported in the present study (varying from 2.7 to 4.6 %) for three cultivars of Brachiaria (accesions 3,401, 3,413 and 3,451) and one P. maximum (3,616), at 3, 6 and 9 wk of regrowth. In the case of hemicellulose, they found values (27.9 to 33.7 %), higher than results of the present study (24.2 to 30.6 %) at same regrowth ages. Cellulose values of Panicum were increased, as expected, when regrowth age increased, thus, declining the forage quality.

In tropical pastures the crude protein content (CP) falls rapidly by advancing pasture maturity. In this experiment, the high protein concentrations of Pennisetum and Panicum showed a reduction by increasing age of regrowth. In Brazil, Duarte et al36 reported for 24 genotypes of P. maximum, fertilized with 250 and 207 kg/ha/yr of N and K, average contents of CP of 15.7 and 13.5 % for 2 yr. These values were slightly superior to the results of the present study, probably due to fertilizer. In Ethiopia, the effect of defoliation frequency (60, 90 and 120 d) of Napier grass (P. purpureum) was evaluated; these authors reported that CP concentration decreased from 12.1 (60 d) to 8.0 % (120 d)37.

In Florida evaluating the cultivars Mulato II (B. ruziziensis clone 44-6 x B. brizantha cv Marandu) and Cayman (Brachiaria hybrids lines BR02/1794 and BR02/1752) fertilized with 144 kg N, and 112 kg K ha-1, had decreasing CP content, (17.4, 14.2, and 11.8 %) were observed as regrowth interval increased from 2, to 4 and 6 wk, respectively38, estimating that these values higher than those of the present study, could be probably due to fertilization.

In situ dry matter degradation (ISDMD)

In the present study, samples of Pennisetum had always higher ISDMD in each evaluation cycle. It decreased as the cutting age increased, in a proportion of 17.3, 21.5 and 3.3 %, at cycle 1 to 3, respectively. In Indonesia (West Sumatra, 2,289 mm year-1), ISDMD of B. decumbens, P. purpureum and A. compressus, after 72 h of incubation during both, rainy and dry season was 60.5 and 57.4 %, 63.3 and 63.5 %, and 58.3 and 57.6 % respectively39, being these values in general lower than the findings of the present experiment; however, there is not an explanation of this fact.

In Tabasco, México, Jimenez et al40 reported that season and age (21, 28 and 35 d) of regrowth affected significantly the DM degradation of B. humidicola after 48 h of incubation. Highest values were detected during the dry season (59.0 %) followed by rain (57.3 %) and winter (55.7 %) period. By regrowth age, they reported largest degradation at 28 d (59.2 %), followed by 21 (55.5 %) and 35 d (56.8 %). These results were lower than the values recorded for 3 to 12 wk, which varied from 81.0 to 53.8 %. Respect to cycle, report of Jiménez et al40 agrees with the results of the present experiment, when also highest values of ISDMD were recorded during first and third cycles with low rainfall.

Gross energy (pre and post incubation)

In the present experiment, total gross energy content was higher for Brachiaria and Panicum, than for the Pennisetum grasses; however, not a clear trend was observed considering regrowth ages. In north of Veracruz, Mexico, Juárez-Reyes et al41 evaluated four representative tropical pastures, P. maximum (Guinea), P. maximum (var Tanzania), Digitaria decumbens (Pangola) and Cynodon dactylon (Bermuda), at beginning of flowering, and reported a gross energy content of 3,940, 3,930, 4,090 and 4,070 kcal kg-1 DM, respectively.

Ruminal degradable energy

At first and second cycle of evaluation in the present study, Pennisetum, had the highest values of degraded energy, both in terms of percentage and of kcal kg-1 de DM). During the two first cycles, a decreasing value of degraded energy was observed as age increased. In Indonesia, a value of 2.6 Mcal ED kg-1 DM was reported for samples of B. brizantha (vegetative stage), evaluated during the rainy season (1,800 mm rain/yr)42. This figure is close to the value for Brachiaria genus (2.52 Mcal kg-1 DM) reported for the second cycle (rainy season). In India other researchers found that digestibility of gross energy of buffelgrass (Cenchrus ciliaris), declined with maturity, from monsoon (rainy season) to summer (54.6 to 37.1 %)43. This performance is similar to the results of the present study for ruminal degradable energy, where the highest value (64.0 %) was recorded during the rainy season, and lower values were registered during the first (49.9 %) and third (50.1 %) cycles, both with dry conditions.

CONCLUSIONS AND IMPLICATIONS

The exponential growth model that described the dry matter yield of forage in terms of time, identified the genus Pennisetum spp as the most productive one. Pennisetum and Brachiaria had better values of crude protein, varying in a range of 10.2 a 11.3 %, than the other tropical grasses (7 a 9 %). The in situ dry matter degradability is in general satisfactory for the evaluated materials. For an ideal grazing management of these genera, it is recommended that they are harvested between 6 and 9 wk of age. These genera also had better growth periods, when precipitation oscillated between 200 to 350 mm.

ACKNOWLEDGEMENTS

Part of this research was supported by the "Programa de Apoyo a Proyectos de Investigación e Innovación Tecnológica" (PAPIIT) through grant to the Project IN213910 Potencial forrajero de gramíneas introducidas en un clima cálido húmedo del estado de Veracruz, responsibility of the second author. The Programa de Apoyos para la Superación del Personal Académico (PASPA) of the Universidad Nacional Autónoma de México supported a sabbatical stay of the first autor at the Facultad de Agronomía, Universidad Autónoma de Nuevo León, in 2014. MVZ Lucía del Rayo Lozano Caro and MC Nydia Vázquez Aguilar for their technical assistance in laboratory.

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Received: December 10, 2014; Accepted: April 13, 2015

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