Tropical Grasslands-Forrajes Tropicales (2017) Vol. 5(2):6676 66

DOI: 10.17138/TGFT(5)66-76

Research Paper

Dry matter accumulation and crude protein concentration in

Brachiaria spp. cultivars in the humid tropics of Ecuador

Acumulación de materia seca y concentración de proteína cruda en cultivares

de Brachiaria spp. en el trópico húmedo de Ecuador

JONATHAN R. GARAY1, SANTIAGO JOAQUÍN CANCINO1, PEDRO ZÁRATE FORTUNA1, MARTÍN A.

IBARRA HINOJOSA1, JUAN C. MARTÍNEZ GONZÁLEZ1, RICARDO P. GONZÁLEZ DÁVILA2 AND EUGENIA

G. CIENFUEGOS RIVAS1

1 Universidad Autónoma de Tamaulipas, Facultad de Ingeniería y Ciencias, Ciudad Victoria, Tamaulipas, Mexico.

www.uat.edu.mx

2 Universidad Tecnológica Equinoccial, Campus Santo Domingo, Investigación y Transferencia de Tecnología, Santo Domingo de los Tsáchilas, Ecuador. www.ute.edu.ec

Abstract

Climatic conditions throughout the year and age of plants affect both yield and quality of forage grasses. In this research, we evaluated the effects of age of regrowth and seasonal conditions on dry matter accumulation and crude protein concentration in 5 cultivars of Brachiaria spp.: Señal, Xaraés, Marandú, Piatá and Mulato II, harvested at 2, 4, 6, 8 and 10 weeks after a uniformity cut, during the rainy and dry seasons. The variables were: total dry matter (TDM), leaf dry matter (LDM) and stem dry matter (SDM) yields, leaf area index (LAI), specific leaf area (SLA) and crude protein (CP) concentration. For TDM yield, in the rainy season there was no significant difference (P>0.05) among cultivars, with mean DM yield over 10 weeks of 6.34 t/ha; however, during the dry season Xaraés presented a higher (P<0.05) yield over 10 weeks than other cultivars (5.09 vs. 3.14‒3.89 t/ha). Overall, mean DM yield in the dry season was only 62% of that in the wet season. In both periods, Señal tended to have the highest SDM yields, while Xaraés had the greatest (P<0.01) LDM yields in the dry season. Mulato II tended to have the highest CP concentrations throughout, especially in the dry season. This study was conducted in plots with plants only 12 weeks old at commencement. However, it indicated that all cultivars performed well and larger-scale studies of longer duration are warranted to test these cultivars under grazing, especially Mulato II, which showed both high dry matter yield and retention of high protein concentration throughout the study.

Keywords: Brachiaria decumbens, Brachiaria brizantha, Brachiaria hybrid cv. Mulato II, leaf area index, specific leaf area.

Resumen

En la finca experimental “El Oasis” de la Escuela de Ingeniería Agropecuaria, Universidad Tecnológica Equinoccial, Campus Santo Domingo, Ecuador, en un suelo Andosol se evaluaron los efectos de la edad de rebrote y la época del año sobre la acumulación de materia seca y la concentración de proteína bruta en 5 cultivares de Brachiaria: Señal ( B. decumbens), Xaraés ( B. brizantha), Marandú ( B. brizantha), Piatá ( B. brizantha) y Mulato II ( Brachiaria híbrido), cosechados a 2, 4, 6, 8 y 10 semanas después de un corte de uniformidad, durante las estaciones lluviosa y seca.

Las variables evaluadas fueron la materia seca (MS) total, MS de hoja y de tallo, el índice de área foliar, el área foliar específica y la concentración de proteína bruta. El rendimiento de MS total a 10 semanas de rebrote (6.34 t/ha en ___________

Correspondence: E. G. Cienfuegos Rivas, Centro Universitario Adolfo

López Mateos, Edificio de Gestión del Conocimiento 4º piso. Cd.

Victoria, CP 87149, Tamaulipas, Mexico.

Email: ecienfue@docentes.uat.edu.mx

Tropical Grasslands-Forrajes Tropicales (ISSN: 2346-3775)

Brachiaria spp. evaluation in Ecuador 67

promedio) en la estación de lluvias no varió entre cultivares (P>0.05) mientras en la estación seca el cv. Xaraés presentó el mayor rendimiento (P<0.05) en comparación con los demás cultivares (5.09 vs. 3.14‒3.89 t/ha). El rendimiento de MS total en la época seca fue sólo el 62% del obtenido en la época de lluvia. En ambos períodos, el cv. Señal tendió a tener los rendimientos de MS de tallos más altos, mientras que el cv. Xaraés presentó los mayores rendimientos de MS

de hojas (P<0.01) en la estación seca. El cv. Mulato II tendió a tener las mayores concentraciones de proteína bruta en ambas épocas, especialmente en la seca. Este estudio se realizó en parcelas con plantas de sólo 12 semanas de edad al inicio. Sin embargo, indicó que todos los cultivares se comportaron satisfactoriamente y estudios a mayor escala y de mayor duración se justifican para probar estos cultivares bajo pastoreo, especialmente el cv. Mulato II que mostró tanto un alto rendimiento de MS como una retención de alta concentración de proteína a lo largo del estudio.

Palabras clave: Área foliar específica, Brachiaria decumbens, Brachiaria brizantha, Brachiaria híbrido cv. Mulato II, índice de área foliar.

Introduction

seasonal changes (Fagundes et al. 2005) on the pattern of

growth, quality and chemical composition of pasture

Poor grassland productivity is one of the most important

plants, in order to optimize their use and plan an

limitations in the dual-purpose cattle system in the

appropriate

agronomic

management

strategy

for

Ecuadorian tropics (Avellaneda et al. 2008), because most

grasslands (Avellaneda et al. 2008). Based on the above,

grazing areas are sown with forage species such as

this research aimed to evaluate the effects of plant

Brachiaria humidicola, Brachiaria decumbens and

regrowth age and seasonal conditions on the

Brachiaria brizantha (Vera 2004). These grasses have

accumulation of dry matter and crude protein

limitations in productivity, adaptability and persistence in

concentration in 5 cultivars of Brachiaria spp. in the

these environments; in addition, they are susceptible to

humid tropics of Ecuador.

the spittlebug of pastures caused by Aeneolamia spp.

(Cardona et al. 2006) and foliar fungi such as Rhizoctonia

Materials and Methods

solani, which significantly reduce productivity (Álvarez

et al. 2013). However, new cultivars of the genus

The research was conducted under field conditions from

Brachiaria have been released to the market as options to

December 2011 to November 2012, at the experimental

overcome the problems observed in traditional forages,

farm “El Oasis”, property of the Escuela de Ingeniería

thus providing better fodder options (Pizarro et al. 2013).

Agropecuaria

of

the

Universidad

Tecnológica

In Ecuador, several cultivars of Brachiaria spp. are

Equinoccial,

Campus

Santo

Domingo,

Ecuador

available that have potential for increasing the

(00°13'20' S, 79°15'39'' W; 406 masl). The experimental

productivity of existing grass production systems (Faría

site has a predominantly humid tropical climate, with

Mármol 2006; Miles 2006). Jácome and Suquilanda

annual average temperature of 23.4 °C and annual rainfall

(2014) indicated that cultivars Mulato I and Xaraés are

of 2,600‒3,500 mm with 2 seasons, defined as rainy and

well accepted by farmers because of their high nutritional

dry (SENPLADES 2015). The monthly rainfall and mean

value, adaptation to a range of soils and resistance to or

maximum and minimum temperatures during the

tolerance of pests and diseases (Cardona et al. 2006; Argel

evaluation period are reported in Figure 1.

2008).

In this research 5 Brachiaria cultivars were used:

The productive capacity of forage species can be

Señal ( B. decumbens, considered as the control since

modified by changing various factors, such as: using

it is most widely cultivated in the area); Marandú

different genotypes or cultivars; modifying the

( B. brizantha), Piatá ( B. brizantha), Mulato II ( Brachiaria

agronomic management and age of the plants at harvest;

hybrid) and Xaraés ( B. brizantha); and these were

and by weather conditions throughout the year (Zaragoza

harvested at 5 growth stages (2, 4, 6, 8 and 10 weeks after

et al. 2009; Lara et al. 2010). It is therefore important to

an initial harvest) during the rainy (March‒May) and dry

assess the dynamics of herbage accumulation of

(September‒November) seasons to monitor the rate of

different pasture species and to understand the impact of

DM accumulation over time and assess quality aspects.

Tropical Grasslands-Forrajes Tropicales (ISSN: 2346-3775)

68 J.R. Garay, S. Joaquín, P. Zárate, M.A. Ibarra, J.C. Martínez, R.P. González and E.G. Cienfuegos

35

900

800

30

700

)

C)

25

m

600

m(

20

ure

500

on i

400

at

15

perat

pit

300

ci

em 10

re

T

200

P

5

100

0

0

jan feb mar apr may jun

jul aug sep oct nov dec

Precipitation

Temp. Max.

Temp. Min.

Figure 1. Monthly precipitation and mean maximum and minimum temperatures during 2012 at the Climatological Station of the Universidad Tecnológica Equinoccial, Campus Santo Domingo, Ecuador.

A randomized complete block design in an

season, observations commenced 2 weeks after each

arrangement in divided plots was used, where the large

uniformity cut to measure: accumulation of total (TDM),

plot was the cultivar and the subplots the ages of

leaf (LDM) and stem (SDM) dry matter; plant height

regrowth. On 3 December 2011, seeds of each cultivar

(PH); leaf area index (LAI); specific leaf area (SLA);

were placed in bags (3 seeds/bag) of black polyethylene

and crude protein concentration (CP). During each

with a capacity of approximately 2 kg of soil, to ensure

sampling for the relevant cutting treatment we measured

one plant survived per bag and placed in a greenhouse. At

the height of plants from ground level to ligule of the

6 weeks after seedling emergence (21 January 2012; 7

fully developed leaf and harvested the forage at 15 cm

weeks after sowing), the plants were transplanted to large

above ground level from the relevant 1 m2 subplot

plots of 5 x 5 m (25 m2), leaving an intra-row and inter-

(4 plants). The harvested forage (green matter) was

row spacing of 0.50 m. The useful area within each large

weighed on a precision scale (Model PB3002-S, Mettler

plot was 3 x 3 m, divided into 9 quadrants of 1 m2

Toledo) and 2 subsamples were taken, the first

(subplots), of which 5 subplots were randomly selected to

(approximately 200 g) to define the proportions of leaf

represent each age of regrowth (2, 4, 6, 8 and 10 weeks).

(leaf blade + sheath) and stem after separating these

The soil of the experimental site is of volcanic origin

components. Leaves from this subsample were used for

(Andosol) and a soil sample was taken using a 5-point

leaf area estimation as well. A multifunctional printer

method to 20 cm depth to provide the physicochemical

scanner (HP Photosmart D110) was employed using the

analyses shown in Table 1. All plots were fertilized after

methodology reported by Rincón et al. (2012). The

transplanting with 120 kg N/ha (as urea), 60 kg P/ha (as

second subsample (approximately 1 kg) was used to

Daphos), 70 kg K/ha (as KCl), 60 kg Mg/ha (as

determine CP concentration by the Kjeldahl method (N x

magnesium oxide) and 50 kg S/ha (as ammonium sulfate).

6.25) (AOAC 2000); CP% was determined only for 4, 6

In the sixth week after transplantation the plants

and 8 week cuttings. All samples were dried in a forced-

were 80‒100 cm tall with 45‒65 tillers/plant; senescent

air oven at 65 °C for 48 h to estimate the dry matter (DM)

material was obvious in the lower stratum, so we

concentration and calculate DM yields.

applied an initial uniformity cut on 3 March 2012 to 15

The data were analyzed in time within season by using

cm above ground level. The evaluation during the rainy

PROC GLM of SAS (SAS Institute 2010), in a complete

season then followed. At the end of this period (12 May

randomized block design in an arrangement in divided

2012), the plots were allowed to stand for 15 weeks,

plots. Tukey’s Studentized range tests (P≤0.05) were

when a second uniformity cut was applied on 1 September

performed, when treatments were significantly different

2012. The dry season evaluation then followed. In each

in each season.

Tropical Grasslands-Forrajes Tropicales (ISSN: 2346-3775)

Brachiaria spp. evaluation in Ecuador 69

Table 1. Chemical1 characteristics of the soil used in the experiment2.

pH

OM3

NH4

P

S

Fe

Cu

Zn

Mn

B

K

Ca

Mg

(%)

(mg/kg)

(cmol/kg)

5.9

2.2

41.0

6.5

6.3

42.0

5.6

1.9

2.8

0.3

0.3

8.3

2.9

1Obtained in the Chemistry Laboratory, Universidad Tecnológica Equinoccial, Campus Santo Domingo.

2The analysis to determine the phosphorus concentration in the soil was performed by the method of Olsen modified with a solution of sodium bicarbonate and EDTA adjusted to pH 8.5 with 10N NaOH.

3The organic matter (OM) analysis was performed using the method of Walkley Black, suitable for the conditions of Ecuador.

Results

with the remaining cultivars producing 3.14‒3.89 t

DM/ha. Average accumulation of TDM during the dry

Total dry matter accumulation

season was 62% of that produced in the rainy season.

Accumulation of total dry matter (TDM) followed an

Leaf dry matter accumulation

exponential pattern in both rainy and dry seasons (Table

2). There were no significant differences (P>0.05) in

Leaf dry matter (LDM) yield during the rainy season

TDM accumulation among cultivars of Brachiaria during

varied between cultivars at the various cutting ages but at

the rainy season, with yields ranging from 5.83 to 6.87 t

the tenth week, there were no significant differences in

DM/ha (mean 6.34 t DM/ha) over the 10 week period.

leaf yield between cultivars (P>0.05; Table 3). On the

However, in the dry season differences between cultivars

other hand, during the dry season, significant differences

(P<0.01) were observed in each of the regrowth periods.

(P<0.05) in leaf yield between cultivars at all ages of

While there was little consistency in which cultivars were

regrowth were observed. Towards the end of the dry

superior within particular cutting frequencies, cultivar

season (8 and 10 week cuttings) Xaraés produced more

Xaraés accumulated the greatest amount of TDM during

leaf (P<0.05) than most other cultivars. Dry season leaf

the later part of the dry season and produced the highest

yield at the 10 week cutting varied from 4.28 t DM/ha

DM yield during the dry season period (5.09 t DM/ha),

(Xaraés) to 2.50 t DM/ha (Marandú) (P<0.001).

Table 2. Total dry matter yields at different ages of regrowth of 5 Brachiaria cultivars during the rainy (March‒May) and dry (September‒November) seasons in humid tropical conditions of Ecuador.

Cultivar

Total dry matter (t/ha)

Rainy season

Dry season

Age of regrowth (weeks)

Age of regrowth (weeks)

2

4

6

8

10

2

4

6

8

10

Señal

0.24

0.96

2.96

4.02

6.87

0.16

0.39

0.75

1.98

3.69

Marandú

0.24

0.85

2.54

4.00

5.83

0.23

0.42

0.73

1.59

3.14

Mulato II

0.29

1.31

2.66

4.33

6.57

0.13

0.28

0.64

1.95

3.76

Piatá

0.28

1.13

2.36

4.36

6.14

0.13

0.41

0.91

1.93

3.89

Xaraés

0.32

1.28

3.16

4.49

6.27

0.13

0.41

0.87

2.49

5.09

Mean

0.27

1.11

2.73

4.24

6.34

0.16

0.38

0.78

1.99

3.92

LSD1

0.09

0.29

1.05

1.59

2.38

0.04

0.1

0.19

0.54

1.09

LS2

0.058

<0.001

0.177

0.812

0.686

<0.001

0.005

0.004

0.003

0.001

1Least Significant Difference (Tukey P≤0.05).

2Level of significance.

Tropical Grasslands-Forrajes Tropicales (ISSN: 2346-3775)

70 J.R. Garay, S. Joaquín, P. Zárate, M.A. Ibarra, J.C. Martínez, R.P. González and E.G. Cienfuegos Table 3. Leaf dry matter yields at different ages of regrowth of 5 Brachiaria cultivars during the rainy (March‒May) and dry (September‒November) seasons in humid tropical conditions of Ecuador.

Cultivar

Leaf dry matter (t/ha)

Rainy season

Dry season

Age of regrowth (weeks)

Age of regrowth (weeks)

2

4

6

8

10

2

4

6

8

10

Señal

0.21

0.75

1.71

1.85

3.13

0.16

0.35

0.66

1.55

2.76

Marandú

0.24

0.79

1.76

2.49

3.46

0.23

0.42

0.72

1.48

2.50

Mulato II

0.29

1.26

1.90

2.66

3.85

0.13

0.28

0.63

1.88

3.19

Piatá

0.28

0.97

1.60

2.60

3.21

0.13

0.41

0.88

1.66

3.02

Xaraés

0.32

1.20

2.45

3.50

4.18

0.13

0.41

0.87

2.34

4.28

Mean

0.27

1.00

1.88

2.62

3.57

0.16

0.37

0.75

1.78

3.15

LSD1

0.08

0.24

0.76

0.93

1.17

0.04

0.10

0.19

0.52

1.01

LS2

0.017

<0.001

0.029

0.002

0.072

<0.001

0.004

0.002

0.001

0.001

1Least Significant Difference (Tukey P≤0.05).

2Level of significance.

Stem dry matter accumulation

respectively), while Marandú and Mulato II had the least

(0.64 and 0.57 t DM/ha) (P<0.001).

As for leaf production, stem dry matter (SDM)

accumulation during the rainy season varied between

Plant height

cultivars from the second to the sixth week (P<0.01). At

the 8 and 10 week cuts cultivars Señal and Piatá tended to

During the rainy season plant height of the different

have the highest stem yields and Xaraés the lowest but

cultivars varied but by the eighth and tenth weeks Señal

differences were not significant (P>0.05; Table 4). During

was shorter than all other cultivars (P<0.001; Table 5).

the dry season, SDM accumulation followed an expo-

During the dry season Piatá, Señal and Xaraés progres-

nential pattern, with greatest stem production between the

sively showed superiority in height over the other

8 and 10 week cuts. At the 10 week cut, cultivars Señal

cultivars and by the tenth week Piatá and Xaraés were

and Piatá had the highest SDM (0.93 and 0.87 t DM/ha,

taller than all others (P<0.01).

Table 4. Stem dry matter yields at different ages of regrowth of 5 Brachiaria cultivars during the rainy (March‒May) and dry (September‒November) seasons in humid tropical conditions of Ecuador.

Cultivar

Stem dry matter (t/ha)

Rainy season

Dry season

Age of regrowth (weeks)

Age of regrowth (weeks)

2

4

6

8

10

2

4

6

8

10

Señal

0.03

0.20

1.22

1.69

2.81

-

0.04

0.09

0.41

0.93

Marandú

-

0.06

0.78

1.39

1.95

-

-

0.02

0.11

0.64

Mulato II

-

0.05

0.76

1.43

2.09

-

-

0.01

0.08

0.57

Piatá

-

0.16

0.76

1.62

2.61

-

-

0.03

0.27

0.87

Xaraés

-

0.08

0.71

1.00

1.86

-

-

0.01

0.15

0.80

Mean

-

0.11

0.84

1.43

2.26

-

-

0.03

0.21

0.76

LSD1

-

0.07

0.37

0.82

0.95

-

-

0.02

0.10

0.22

LS2

-

<0.001

0.005

0.127

0.068

-

-

<0.001

<0.001

0.001

1Least Significant Difference (Tukey P≤0.05).

2Level of significance.

Tropical Grasslands-Forrajes Tropicales (ISSN: 2346-3775)

Brachiaria spp. evaluation in Ecuador 71

Table 5. Plant height at different ages of regrowth of 5 Brachiaria cultivars during the rainy (March‒May) and dry (September‒

November) seasons in humid tropical conditions of Ecuador.

Cultivar

Plant height (cm)

Rainy season

Dry season

Age of regrowth (weeks)

Age of regrowth (weeks)

2

4

6

8

10

2

4

6

8

10

Señal

43.1

53.5

81.8

95.4

80.2

29.6

42.5

70.4

91.3

77.5

Marandú

40.3

48.8

77.2

117.1

129.8

32.4

30.0

41.5

55.0

67.8

Mulato II

47.7

57.3

83.5

108.3

129.2

32.5

31.2

43.1

57.9

75.8

Piatá

45.8

63.8

98.8

132.1

137.1

32.0

39.2

65.7

95.4

105.5

Xaraés

54.5

71.6

91.2

126.3

137.8

32.3

48.3

71.0

91.3

103.5

Mean

46.3

59.0

86.5

115.8

122.8

31.8

38.2

58.4

78.2

86.0

LSD1

9.4

10.6

14.3

10.1

19.8

10.1

9.1

16.5

16.8

23.8

LS2

0.005

0.001

0.003

<0.001

<0.001

0.879

<0.001

<0.001

<0.001

<0.001

1Least Significant Difference (Tukey P≤0.05).

2Level of significance.

Leaf area index

the lowest SLA throughout the wet season, while Señal,

Mulato II and Marandú presented the highest values. In

During the rainy season, the leaf area index (LAI)

the dry season, cultivar differences again emerged with

followed a linear pattern for all cultivars, increasing from

Mulato II, Marandú and Señal having the greatest SLA

a mean of 0.6 at week 2 to 5.3 at week 10 (Table 6). The

and Xaraés and Piatá the lowest (P<0.01).

cultivars Mulato II and Xaraés reached the highest LAI at

week 10 of 6.6 and 5.8, respectively. Throughout the wet

Crude protein concentration

season Piatá had lower LAI than most other cultivars

(P<0.001). During the dry season, LAI followed an

Data for weeks 4, 6 and 8 on whole plant samples are

exponential pattern with the highest increase from the

presented in Table 8. Differences in crude protein

sixth week. As for the wet season, Piatá showed lower

concentration (P<0.05) between cultivars were observed

LAI than Xaraés and Mulato II by week 10 (P<0.05).

in both rainy and dry seasons, where Mulato II presented

the highest concentration at most observations. In both

Specific leaf area

seasons, mean crude protein concentration decreased as

the regrowth age increased, declining from 14.1% to 9.1%

Specific leaf area (SLA) declined in all cultivars during

in the wet season and from 12.6% to 7.6% in the dry

both rainy and dry seasons (Table 7). Piatá tended to have

season.

Table 6. Leaf area index at different ages of regrowth of 5 Brachiaria cultivars during the rainy (March‒May) and dry (September‒

November) seasons in humid tropical conditions of Ecuador.

Cultivar

Leaf area index

Rainy season

Dry season

Age of regrowth (weeks)

Age of regrowth (weeks)

2

4

6

8

10

2

4

6

8

10

Señal

0.5

2.0

3.2

3.7

5.1

0.4

0.9

1.2

2.7

4.8

Marandú

0.6

1.8

3.1

4.3

5.4

0.7

1.0

1.6

2.5

4.4

Mulato II

0.8

2.6

3.8

4.7

6.6

0.3

0.8

1.4

3.6

5.8

Piatá

0.5

1.3

2.1

3.4

3.8

0.3

0.7

1.4

2.2

3.6

Xaraés

0.7

1.9

3.5

4.9

5.8

0.3

0.9

1.6

3.6

5.9

Mean

0.6

1.9

3.1

4.2

5.3

0.4

0.9

1.4

2.9

4.9

LSD1

0.2

0.4

1.0

1.1

1.2

0.2

0.2

0.3

0.9

2.2

LS2

0.001

<0.001

0.002

0.004

<0.001

<0.001

0.045

0.016

0.001

0.03

1Least Significant Difference (Tukey P≤0.05).

2Level of significance.

Tropical Grasslands-Forrajes Tropicales (ISSN: 2346-3775)

72 J.R. Garay, S. Joaquín, P. Zárate, M.A. Ibarra, J.C. Martínez, R.P. González and E.G. Cienfuegos Table 7. Specific leaf area at different ages of regrowth of 5 Brachiaria cultivars during the rainy (March‒May) and dry (September‒

November) seasons in humid tropical conditions of Ecuador.

Cultivar

Specific leaf area (cm2/g)

Rainy season

Dry season

Age of regrowth (weeks)

Age of regrowth (weeks)

2

4

6

8

10

2

4

6

8

10

Señal

229

246

181

190

151

248

232

173

163

164

Marandú

242

216

168

163

145

273

221

213

158

162

Mulato II

268

190

187

165

162

238

279

210

178

168

Piatá

156

129

125

124

112

221

173

148

125

112

Xaraés

207

148

134

130

132

237

213

170

142

129

Mean

221

186

159

154

140

243

224

183

153

147

LSD1

29

30

41

40

34

85

39

25

28

27

LS2

<0.001

<0.001

0.001

0.001

0.005

0.433

<0.001

<0.001

<0.001

<0.001

1Least Significant Difference (Tukey P≤0.05).

2Level of significance.

Table 8. Crude protein concentration at different ages of regrowth of 5 Brachiaria cultivars during the rainy (March‒May) and dry (September‒November) seasons in humid tropical conditions of Ecuador.

Cultivar

Crude protein concentration (%)

Rainy season

Dry season

Age of regrowth (weeks)

Age of regrowth (weeks)

4

6

8

4

6

8

Señal

13.4

11.2

9.0

11.6

7.3

6.5

Marandú

14.0

12.1

8.8

13.4

8.6

8.1

Mulato II

14.6

12.5

10.1

14.3

10.1

10.2

Piatá

13.8

10.9

8.9

12.2

8.3

6.8

Xaraés

14.6

11.1

8.6

11.6

7.4

6.5

Mean

14.1

11.6

9.1

12.6

8.3

7.6

LSD1

0.9

1.1

0.8

0.9

1.3

3.4

LS2

0.007

0.003

0.001

<0.001

<0.001

0.022

1Least Significant Difference (Tukey P≤0.05).

2Level of significance.

Discussion

The absence of cultivar differences in total DM

production during the rainy season in this study was

This plot study has provided useful information on the

similar to the findings of Rojas-Hernández et al. (2011)

potential of the Brachiaria cultivars for use in pastures of

for B. decumbens, B. brizantha cv. Libertad, B. hybrid

humid tropical Ecuador. While the grasses were planted

cv. Mulato I and cultivars of B. humidicola. Those authors

from seed, were only 12 weeks old at commencement of

also failed to demonstrate any significant differences in

observations and the study was in only a single year, the

total DM production. However, the differences we

excellent growth obtained and CP levels maintained

showed in proportions of leaf and stem in the different

indicate that these pastures can be very productive under

cultivars are important, as we showed higher production

these conditions. In fact, the DM yields obtained over

of leaves and higher CP concentration than Reyes-Purata

only 10 weeks in pastures that were only 12 weeks old at

et al. (2009). The reduction in growth in these cultivars

commencement are quite remarkable. Further studies

during the dry season was not surprising, as the major

under field conditions would clarify how well these

climatic factors that determine forage production are

preliminary findings can be extrapolated to commercial

precipitation and temperature (Gerdes et al. 2000;

situations.

Cuadrado et al. 2004). Rainfall in the rainy season and

Tropical Grasslands-Forrajes Tropicales (ISSN: 2346-3775)

Brachiaria spp. evaluation in Ecuador 73

hence soil moisture was much higher than in the dry

The differences among cultivars for LAI may be

season and minimum temperatures were also lower in the

because some species such as B. decumbens are shorter

dry season. The reduction in TDM yield in the dry season

than B. brizantha, and there is a positive correlation

was somewhat lower than the 50% reduction reported by

between plant height and LAI (Guenni et al. 2005). In

Benítez et al. (2007). This marked reduction in yield

addition, LAI increases as the plant grows (Gómez-

accompanied by a drop in CP concentration indicates the

Carabalí et al. 1999), and this increase is closely related

reduced carrying capacity of these pastures during the dry

to tillering of the crop and soil cover (Rincón et al. 2007;

season.

Ramírez-García et al. 2012). Therefore, as LAI of the

The differences between cultivars in leaf DM yield we

pasture increases, the amount of light that reaches the

found in this study may be due to B. decumbens and

ground will be reduced, which can prevent or retard weed

B. brizantha having different rates of elongation in leaves

growth.

(Dias-Filho and Carvalho 2000). In this regard,

Furthermore, it has been reported that SLA in

elongation rates in leaves of 2.5 and 3.5 cm/d and leaf

Brachiaria cultivars varies according to species; Baruch

percentages in total DM of 54 and 77% in B. decumbens

and Guenni (2007) reported that at 4 weeks of age SLA in

and B. brizantha, respectively, have been reported by

B. decumbens and B. brizantha were 300 and 270 cm2/g,

Guenni et al. (2005). In addition, Hare et al. (2009)

respectively. Also, it has been reported that, as the height

reported that Xaraés and Mulato II have higher foliar

and age of the plant increase, SLA decreases (Gómez et

DM yields than other Brachiaria cultivars, especially

al. 2012), as it did in our study, because there is an

during the dry season. Similarly, Gerdes et al. (2000)

increment in the thickness of the leaves. Grasses with

reported a significant decrease in the accumulation of

greater SLA have thinner leaves and higher concen-

total DM during the period of minimum precipitation,

trations of nitrogen (Pérez et al. 2004) and higher rates of

with as much as 98% of the accumulation corresponding

photosynthesis (Reich et al. 1997), while a high SLA is

to leaf DM.

related to greater palatability and consumption by animals

The pattern of stem DM accumulation during both

(Lloyd et al. 2010; Zheng et al. 2014).

periods would have been determined by environmental

The CP values obtained in this study for Señal

conditions that affected growth of the cultivars. Stem

were higher than those reported by Alvarado et al. (1990)

production during the dry season is reduced (Cab et al.

at 6 and 9 weeks of age (9.4 and 8.8%, respectively),

2008) due to stress caused by soil moisture deficiency,

while the values obtained in Mulato II were similar to

curbing the growth of the plant and reducing stem growth

those found by Castillo et al. (2006) at 3 weeks (15%).

(Cruz et al. 2011a). Cruz et al. (2011b) reported that

The values obtained in Xaraés, Marandú and Piatá were

season has a marked effect on the relative growth of leaf

similar during the rainy season to, and higher in the dry

and stem, and differences between stem elongation rates

season than, those obtained by Pérez et al. (1999) in

of 1.2, 0.8 and 0.6 mm/stem/d in Señal, Xaraés and

B. brizantha at 6 and 9 weeks (11.9 and 8.6%,

Marandú, respectively, have been reported by Paciullo et

respectively). Water deficiency stress can have a negative

al. (2011). The marked reduction in stem growth in the

effect on Brachiaria cultivars, decreasing the con-

dry season results in increases in leaf:stem ratios in this

centration of CP. In this regard, Cuadrado et al. (2004)

season as found in this study. Leaf:stem ratios also

reported that at 24 days of regrowth CP concentration in

declined significantly as plants matured and the

the wet season was higher than in the dry season for

proportion of stem in the forage increased.

Marandú (10.5 vs. 9.3%), Señal (15.4 vs. 9.2%) and

Plant height was affected to a greater extent by

Xaraés (11.5 vs. 8.2%). Increasing age of the plant also

seasonal conditions than by cultivar. Marandú has been

decreases the concentration of CP in forage because of

shown to be quite susceptible to stress caused by soil

accumulation of dry matter (resulting in dilution of

water deficiency, presenting taller plants in the rainy

nutrients), increase in stem DM and decline in the

season (67 cm) and shorter ones in the dry season (36 cm)

proportion of leaf (Juárez-Hernández et al. 2004; Reyes-

at 35 d of regrowth (Gerdes et al. 2000). On the other

Purata et al. 2009), as shown in our study. All cultivars

hand, B. decumbens presented lower height, due to its

had CP concentrations above 7% which is considered the

decumbent growth habit, which is characteristic of this

minimum, below which intake by ruminants could be

species (Pérez et al. 1999). In this study, B. brizantha

suppressed (Lazzarini et al. 2009). The choice of cultivar

cultivars were taller than the Brachiaria hybrid and B.

to use could depend on a range of factors including DM

decumbens, which agrees with what was reported by

yield of leaf, CP concentration, rate of regrowth, response

Gómez et al. (2000).

to fertilizer etc.

Tropical Grasslands-Forrajes Tropicales (ISSN: 2346-3775)

74 J.R. Garay, S. Joaquín, P. Zárate, M.A. Ibarra, J.C. Martínez, R.P. González and E.G. Cienfuegos Conclusions

Avellaneda C; Cabezas JF; Quintana G; Luna R; Montañez O;

Espinoza I; Zambrano S; Romero D; Vanegas J; Pinargote

While the dry season negatively affected the production

E. 2008. Comportamiento agronómico y composición

and nutritional value of forage of all grasses evaluated, the

química de tres variedades de Brachiaria en diferentes

edades de cosecha. Ciencia y Tecnología 1:87‒94.

most outstanding cultivars in terms of leaf yield, leaf area

https://goo.gl/orgr4w

index and specific leaf area were: Mulato II, Marandú and

Baruch Z; Guenni O. 2007. Irradiance and defoliation effects in

Xaraés. Xaraés certainly showed the best tolerance of dry

three species of the forage grass Brachiaria. Tropical

conditions, while performing well in terms of total DM

Grasslands 41:269‒276. https://goo.gl/ttgvtU

and leaf production in the rainy season. However, Mulato

Benítez D; Fernández JL; Ray J; Ramírez A; Torres V; Tandrón

II showed high DM yield and nutritional value over time,

I; Díaz M; Guerra J. 2007. Factores determinantes en la

which supports its further evaluation in grass production

producción de biomasa en tres especies de pastos en

systems in humid tropical conditions, especially as good

sistemas racionales de pastoreo en el Valle del Cauto, Cuba.

CP levels (11‒14%) were maintained for long periods in

Revista Cubana de Ciencia Agrícola 41:231‒235.

both rainy and dry seasons. This was a plot study and

https://goo.gl/dKPEhj

plants were only 12 weeks old when observations

Cab F; Enríquez J; Pérez J; Hernández A; Herrera J; Ortega E;

commenced, but the results do indicate that all cultivars

Quero A. 2008. Potencial productivo de tres especies de

performed well and further evaluation of the more

Brachiaria en monocultivo y asociadas con Arachis pintoi

promising cultivars under field conditions, particularly

en Isla, Veracruz. Técnica Pecuaria en México 46:317–332.

under grazing, is warranted.

Cardona C; Sotelo G; Miles JW. 2006. Resistencia en

Brachiaria a especies de salivazo: Métodos, mecanismos y

avances. Pasturas Tropicales 28(1):30‒35. https://goo.gl/

Acknowledgments

RvWXjU

Castillo MS; Vélez M; Rosas JC; Trabanino R. 2006.

We thank the Consejo Nacional de Ciencia y Tecnología

Producción y composición de los cultivares Mulato I y II de

(CONACYT) for the grant that made possible the stay of

Brachiaria híbrido inoculados con micorriza y Trichoderma

the first author in Ecuador. Our thanks to the Universidad

harzianum. Ceiba 47:25‒32. DOI: 10.5377/ceiba.v47i1-

Tecnológica Equinoccial, Campus Santo Domingo

2.444

(Ecuador) and the Facultad de Ingeniería y Ciencias of the

Cruz PI; Hernández A; Enríquez JF; Mendoza SI; Quero AR;

Universidad Autónoma de Tamaulipas (Mexico) for the

Joaquín BM. 2011a. Desempeño agronómico de genotipos

facilities provided to make this research possible.

de Brachiaria humidicola (Rendle) Schweickt en el trópico

húmedo de México. Revista Fitotecnia Mexicana 34:123‒

131. https://goo.gl/2dVuyp

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Testing functional trait-based mechanisms underpinning

(Received for publication 15 September 2016; accepted 02 May 2017; published 31 May 2017)

© 2017

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Tropical Grasslands-Forrajes Tropicales (ISSN: 2346-3775)