Introduction

Andean foothills. Overall, cropped areas represent 4.3‒

11.8%, whereas rangelands cover 72‒89% of the Orinoco

The continuous increase in demand for food requires

basin in Colombia (DANE 2016).

intensification of land use systems in the agricultural

This expansion of crops and plantations has taken

sector and achieving a compromise between increases in

place at the expense of native savannas supporting beef

production and ecological conservation constitutes an

cattle ranching. On areas closest to roads in the Plains,

important challenge (Davies et al. 2010; van Vuuren and

beef breeding herds have been replaced by yearlings

Chilibroste 2013). Additionally, climate change may

brought in from the surrounding area for fattening,

further increase the demands on management of resources

interspersed with crops and plantations (Romero-Ruiz et

and the capacity of livestock farmers and their rural

al. 2012; Huertas-Ramírez and Huertas-Herrera 2015).

communities to adapt (Herrero et al. 2017).

The cattle population of the region is estimated at 4.7 M

The watershed of the Orinoco River covers 35 Mha in

head (DANE 2016), largely supported by native savannas

Colombia (33% of the land area) and includes mountains,

that still constitute the main land use in the rest of the area,

foothills and a variety of seasonally flooded and well

mainly dedicated to extensive beef breeding herds

drained

savannas.

The

well

drained

savannas

(Rausch 2013; Huertas-Ramírez and Huertas-Herrera

(“Altillanura”) extend over 13.5 Mha (CONPES 2014),

2015). The latter are low input-low output systems,

35% of which is plain (“Plains”) and 54% is slopes and

frequently constrained by limited access, physiographic

hills (Rippstein et al. 2001). The latter include small

and water limitations, low soil fertility and often shallow

valleys suitable for cropping, surrounded by pronounced

soils that limit intensification (Seré and Vera 1983). On

slopes that have shallow, stony soils. By 2007, 23% of the

the other hand, these savannas are rich in plant and animal

Plains had been converted to crops, sown pastures, palm

biodiversity (Rippstein et al. 2001; Lasso et al. 2011) and

oil plantations (Romero-Ruiz et al. 2012; Rausch 2013)

have varied landscapes and ecosystem services that attract

and a variety of tree plantations and reforested areas

rural tourism highlighted by educational and cultural

amounting to close to 100,000 ha (MADR 2015). Palm oil

values and traditions (Navas Ríos 1999; Molina and

plantations have experienced a significant and ongoing

Triana 2011; Australian Government 2015).

increase in the savannas, and have reached 112,186 ha

Although the real impact of greenhouse gas emissions

(FEDEPALMA 2015), including large areas in the

from cattle on neotropical savannas is still a challenge for

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

52 R. R. Vera and C.A. Ramírez-Restrepo

scientists, it is likely that plant dynamics may mitigate the

et al. 1988; Schlink et al. 1992; Short et al. 1996; Tyler et

demanding effects of climate variability, reflecting the

al. 2012). In the majority of cases, weaners have either

capacity of these plant-animal evolutionary systems to

been raised in feedlots (Arthington et al. 2005) or supple-

adapt to genetic, environmental and management

mented with concentrates or crop by-products on pasture

stressors (O’Neill et al. 2010; Herrero et al. 2015;

(Vendramini et al. 2007; Vendramini and Arthington

Ramírez-Restrepo and Charmley 2015). In this complex

2008). Consequently, Tyler (2012) indicates that the

scenario, primary producers have to contend with low

effect of tropical pastures on the performance of early

seasonal biomass production and nutritive value

weaners should be prioritized in future pastoral research.

(Paladines and Leal 1979; Rippstein et al. 1996; Durmic

Early weaning has rarely been investigated in the

et al. 2017). Together, these factors negatively impact

neotropics (Moore and da Rocha 1983; Betancourt-López

productivity in terms of slow growth and fertility rates

et al. 2012) and, as elsewhere, the practice has generally

that seldom exceed 50%, yielding no more than 3 or 4

been coupled with regular supplementation of cows and

calves weaned over a cow’s lifetime (Kleinheisterkamp

calves with different combinations of concentrates and/or

and Habich 1985; Plessow 1985; Squires and Vera 1992).

other feedstuffs in different settings. However, to the

These traits contrast with the relatively high per-

authors’ knowledge, raising early weaned beef calves

formance reported when Brahman ( Bos indicus) beef

exclusively on sown pastures in the neotropics has not

cows are grazed year-round on well managed tropical

been investigated.

sown pastures (Vera et al. 2002). Nevertheless, expensive

The objectives of this study were to assess the effects

sown pastures are mostly reserved for fattening yearlings

on cow and calf performance of early weaning of beef

and steers (Vera and Seré 1985), and their year-round use

calves onto sown tropical pastures, while their dams were

by the breeding herd may represent an economically

maintained on savanna, in comparison with cow-calf pairs

suboptimal use of an expensive resource. It is possible

grazing native savanna with weaning at the conventional

that strategic and seasonal grazing of improved pastures

age.

by suckling cows to complement native savanna grazing

may significantly increase reproductive indexes (Vera

Materials and Methods

and Seré 1989).

Reproductive rates may be boosted further by early

Experimental design

weaning of calves, a technology that is 50 years old. In

principle, early weaning can be performed at 45 days of

The study was conducted during the late 1980s over 6

age (Rasby 2007) and it is particularly useful in drought

consecutive years at Carimagua Research Station, located

situations and to contend with the negative effects of

87 km northeast of Puerto Gaitán in the Meta Department

climate change (FAO 2013). Cow-calf research in the

on the eastern plains of Colombia (4°36'44.6" N,

savannas of northern Australia has been amply docu-

74°08'42.2" W). Monthly rainfall, ambient temperature

mented in over 100 references and reports summarized by

and their annual variations were recorded during 13

Tyler (2012) and Tyler et al. (2012). Similarly, a large

consecutive years (Table 1).

amount of research was carried out in the USA

Care of animals and experimental procedures were

(Arthington et al. 2005; Vendramini et al. 2006). Much of

performed by accredited Doctors of Veterinary Medicine

this research has addressed the consequences of early

(DVM), including the second author, following national

weaning on the dams’ reproductive performance (Fordyce

husbandry and animal welfare regulations.

Table 1. Monthly average climatic data over 13 years, and annual rainfall recorded during the 1984‒1987 period at Carimagua Research Station, Meta Department, Colombia.

Jan

Feb

Mar

Apr

May

Jun

Jul

Aug

Sep

Oct

Nov

Dec

Mean rainfall 1979‒1991 (mm)

10

25

76

236

292

368

274

260

292

203

116

50

Mean ambient temp 1979‒1991 (ºC)1 26.9

28.0

28.1

27.1

26.2

25.4

25.2

25.7

26.1

26.5

26.9

26.4

Annual rainfall (mm)

1984

94

47

53

213

125

439

179

246

529

351

222

25

1985

0

0

33

196

544

445

327

250

351

266

180

0

1986

36

32

65

221

570

541

462

367

286

214

147

89

1987

n/a

n/a

n/a

n/a

558

528

445

354

275

204

138

82

n/a - records not available.

1Mean monthly temperature for 1984‒1987 not included as there is less than 1 ºC variation between years.

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

Grass-legume pastures for early weaning in savanna 53

During the first 2 years (Years 1 and 2) exploratory

0.2 cows/ha in large (>200 ha) paddocks managed with

trials were conducted to assess the feasibility of raising

periodic fire following traditional and regional farming

early weaned beef calves on sown pastures alone, and to

practices (Kleinheisterkamp and Habich 1985; Rippstein

monitor the associated health and mortality risks. At the

et al. 2001).

end of the experimental phase of Year 2, the weaners were

In Year 2, the experiment was methodologically

transferred to a savanna paddock and their weight was

similar to Year 1, but 15 calves were weaned at 68 ± 9 kg

monitored until the savanna controls were weaned at the

ILW and 110 ± 8 days of age on 30 May 1985 at the

age of 266 ± 7 days. A further 18 months were required

beginning of the rainy season. They were rotationally

to monitor growth of these weaners on savanna during

grazed on 15 paddocks of AgPp and Ag plus Centrosema

537 days after the early weaning date, and of the

acutifolium cv. Vichada (accession CIAT 5277; AgCa; 6

performance of their dams until the subsequent calving

calves/ha with 7 days grazing, 21 days rest) for 147 days.

event. In the following 2 years (Years 3 and 4), trials

In parallel, 15 similar calves (Control 2) continued to

compared the performance of early weaned calves on a

suckle their dams as part of a large herd grazing native

number of sown tropical pastures with that of normally

savanna, and were weighed at intervals of 21 days during

weaned calves on savanna.

the same period but weaned at 266 ± 7 days of age and

In all cases Brahman and crossbred [Brahman x San

144 ± 20 kg LW. Thereafter, weaned control and early

Martinero (native; B. taurus)] cows and calves were used.

weaned calves were grazed on a native savanna paddock

Calves were born on savanna with a mean live weight

as a single group at 0.25 head/ha for 18 months, and were

(LW) of 22.7 ± 3.4 kg. Body weight of calves during the

weighed about every 120 days.

experimental phase on sown pastures was recorded every

During Year 3, the experiment compared the

7 days. Cows remained on large savanna paddocks as part

performance of groups of 10 weaned calves (68 ± 13 kg

of a large herd and were weighed and rectally palpated at

ILW, 93 ± 4 days of age) rotationally grazing 4 paddocks

~4-monthly intervals to determine if they were pregnant.

(6 calves/ha; 7 days grazing, 21 days rest) each of Ag,

Calving events were recorded daily, but mustering of the

AgPp and AgCa for 123 days during the wet season,

cows was avoided near calving time. Pregnancy rates and

commencing on 25 June 1986 (mid rainy season).

inter-calving intervals were calculated. Approximate date

Year 4 replicated the design of the third year, with the

of conception was back calculated from the calving date.

addition of a Brachiaria humidicola cv. Llanero (syn. B.

Internal and external parasite infestations were controlled

dictyoneura)- Arachis pintoi (BhAp) pasture subjected to

throughout following commercial farming practices.

the same management and sampling practices previously

Mortalities and incidents of ill health were recorded.

described. The experiment began on 23 July 1987. Ten

Cows and calves had free access to fresh water plus a

calves (81 ± 9 kg ILW, 86 ± 5 days of age) were weaned

commercial mineral supplement containing (as-fed):

and placed on each improved pasture, while 10 unweaned

17.5% Na, 26.9% Cl, 8.0% P, 13.7% Ca, 2.0% S, 0.104%

calves grazed on savanna with their dams until weaning

Cu, 0.35% Zn, 0.001% Co and 0.008% I. Supplement

at 319 ± 29 days of age and 155 ± 25 kg live weight.

consumption was recorded every 15 days.

In Year 3 a pilot test of weaning was carried out

Animals, forages and grazing management

starting on 28 July 1986 with one commercial herd at

Carimagua. Forty-six calves born in 2 different savanna

In the first year (Year 1) 10 male calves [112 ± 16 kg

paddocks were weaned at a mean age of 186 ± 62 (range

initial LW (ILW); mean ± standard error of the mean

54‒285) days and LW 131 ± 26 kg, and were transferred

(s.e.m.)] were weaned at 166 ± 10 days old on

to a 1-year-old A. gayanus-Stylosanthes capitata cv.

11 November 1984, coinciding with the end of the rainy

Capica pasture stocked at 3 calves/ha for 99 days until the

season. They were rotationally grazed (6 calves/ha;

end of the rainy season.

7 days grazing, 21 days rest) for 146 days during the dry

season using 4 paddocks of equal size of a 6-year-old

Sample collection and lab analyses

mixture of Andropogon gayanus cv. Carimagua-1 and

commercial Pueraria phaseoloides (AgPp). Fifteen

Pasture. Pre-grazing herbage mass and botanical

cow-calf pairs grazing savanna as part of a larger herd

composition of the introduced pastures were estimated by

served as controls (Control 1) and were monitored at

the BOTANAL method (Tothill 1978; data partially

intervals of ~120 days. Their calves were weaned at

shown). Botanical composition, growth rate and nutritive

280 ± 29 days of age, and the dams were monitored until

value of the savanna have been previously described by

the next calving event. Savanna cows were stocked at

Rivera Sánchez (1988).

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

54 R. R. Vera and C.A. Ramírez-Restrepo

Blood. Jugular blood samples were collected weekly from

shown due to space limitations). Regressions were

each of the calves into two 10 ml BD Vacutainers®

calculated with the GLMSELECT procedure.

(Becton Dickinson, Franklin Lakes, NJ, USA) for

hematology [hematocrit (g/100 ml)], serum enzymes

Results

[aspartate aminotransferase (AST, U/ml) and gamma-

glutamyl transferase (GGT, U/ml)], total protein (g/100

Forages and botanical composition

ml), renal function [urea nitrogen (BUN, mg/100 ml)] and

mineral [P, Ca and Mg (g/100 ml)] analyses. Hematology

Pre-grazing herbage mass in all introduced pastures

and serum biochemistry analyses were performed at the

and across the experimental periods in Years 3 and 4

International Center for Tropical Agriculture, Cali,

always exceeded 2,500 kg DM/ha, except in the Ag

Colombia. Blood enzymes, protein and BUN were

treatment in Year 3 for a brief period during temporary

determined using standard kits (Sigma-Aldrich Corp., St.

flooding of the paddock. Legume percentage in the forage

Louis, MO, USA), N by the micro-Kjeldahl method, P by

on offer was higher for the AgPp (29%) mixture than for

colorimetry and the remaining minerals by atomic

the BhAp (20%) association, with lower amounts

absorption.

(7‒10%) in the AgCa sward. These legume percentages

Feces. Fecal grab samples were individually collected

remained rela-tively stable throughout the experimental

from the rectums of animals at weighing times to

period as shown for Year 3 in Figure 1. The Ag pasture

determine P, Ca, ash and N concentrations, with N

averaged 758 g/kg DM of neutral detergent fiber (NDF)

expressed as percent of fecal organic matter.

and 80 g/kg DM crude protein (CP) during the rainy

season, and 759 and 89 g/kg DM in the dry season,

Statistical analyses

respectively. The AgPp and AgCa pastures had similar

nutritive composition and averaged 756 g NDF and 115 g

Data were analyzed using the Statistical Analysis System,

CP/kg DM and 757 g NDF and 110 g CP/kg DM for the

version 9.4 (SAS Institute, Cary, NC, USA). Results for

rainy and dry seasons, respectively. Similarly, the BhAp

the first 2 years were summarized using descriptive

pasture contained 716 g NDF and 84 g CP/kg DM in the

statistics (means ± s.e.). In Years 3 and 4, calves were

wet season, and 691 g NDF and 21 g CP/kg DM in the dry

balanced for LW and randomly allocated to sown pastures

season.

considering in all cases individual animals as the

experimental unit. Data distribution from all variables

Mortalities

examined (i.e. blood, feces and LW) was reviewed prior

to additional analyses. Repeated-measures of blood, feces

Over the 4 years of the study, 4 deaths out of 95 early

and LW for the same calf were analyzed with the

weaned calves occurred in the AgPp paddocks in Year 3,

GLIMMIX procedure, using a linear mixed model that

which was related to a temporary flooding event. No calf

included the fixed effect of pasture (i.e. sown forages and

mortality of control, suckling calves or their dams was

savanna), and the interaction between pasture and the

recorded in the savanna paddocks.

random effects of year. All interactions were initially

included, and those that were not significant were

Liveweight performance

discarded for the final analysis (Gbur et al. 2012).

Final analyses were preceded by a study of the

Years 1 and 2. Data in Table 2 show the LWs and ages of

covariance structure (Gbur et al. 2012) to adjust the model

calves during Years 1 and 2. Daily LW gains (LWG) of

specification as required. Differences were considered

calves weaned at 166 days of age in Year 1 and grazed on

significant when P≤0.05, and there was tendency to

the AgPp pasture averaged 0.10 ± 0.03 kg/head, whereas

significance if P≤0.10. Denominator degrees of freedom

contemporary suckling calves on savanna gained 0.35 ±

for the test of fixed effects were specified by the

0.19 kg/head. In Year 2, calves weaned at 110 days of age

Kenward-Roger procedure. Multiple comparisons of least

and grazed on AgPp and AgCa pastures gained 0.19 ±

squares means used the Tukey procedure, complemented

0.06 kg/day, and control suckling calves gained 0.60 ±

with graphical interpretation using SAS diffograms (not

0.13 kg/day.

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

Grass-legume pastures for early weaning in savanna 55

8.,000

a)/h ) 6.,000

a

DM /hMkg D (g 4.,000

erf r (k offfe on 2.,000

o

age onr doeFeF

0.,000

0

20

40

60

80

100

120

140

Experimental days

Experimental days

Grass DM AgPp

Grass DM AgCa

Grass DM Ag

Green DM AgPp

Green DM AgCa

Green DM Ag

Leg DM AgPp

Leg DM AgCa

Figure 1. Forage on offer (kg DM/ha) in 3 Andropogon-based pastures and their respective botanical compositions in Year 3. Grass DM stands for total grass DM on offer in the respective pastures; Green DM is green grass DM; Leg DM is total legume DM. Ag =

Andropogon gayanus; Pp = Pueraria phaseoloides; Ca = Centrosema acutifolium.

Live weights of cows for Years 1 and 2 between

(1985‒1986). Early weaned dams in Year 2 calved at

consecutive calving seasons on savanna are shown in

the end of the dry season, whereas control cows did

Figure 2. Year 1 dams of early weaned calves calved in

so at the end of the following rainy season. Calculated

the late rainy season, while control cows (normally

LWs at conception for the 2 years were 303 ± 43 and 321

weaned) calved at the end of the subsequent dry season

± 45 kg, respectively.

Table 2. Mean (± s.e.) live weights and ages of early weaned Brahman and Brahman cross calves during the exploratory observations in Years 1 and 2.

n

Initial

Final

Year 1

Live weight (kg)

10

112 ± 16

126 ± 18

Age (d)

10

166 ± 10

312 ± 10

Gain (kg/d)

0.10

Year 2

Live weight (kg)

15

69 ± 10

97 ± 13

Age (d)

15

109 ± 8

257 ± 8

Gain (kg/d)

0.19

n - number of animals.

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

56 R. R. Vera and C.A. Ramírez-Restrepo

400

380

360

kg))g (tt (kh 340

eighigew w 320

wwoCoC

300

280

260

0

100

200

300

400

500

600

700

Days postpartum

Days postpartum

Early 110

Early 166

Normal 266

Normal 280

Figure 2. Postpartum cows’ live weights in relation to type of weaning. Early and normal refer to weaning treatment, followed by the calves’ ages at weaning. Standard deviations are not shown for clarity, and ranged between 32 and 42 kg, and 24 and 55 kg for early and normal weaning, respectively. Vertical arrows indicate weaning times and ellipses indicate time and spread of calvings.

Years 3 and 4. Consistent with the preliminary observa-

(P<0.0001) in calves on improved forages than in their

tions, calves were weaned at average ages of 93 and 86

suckling counterparts on savanna (Table 3). Differences

days in Years 3 and 4, respectively (Table 3). At the end

in LW productivity amongst sown forages were small

of the 123 days of experimentation, LW was lower

(P>0.05), with the exception of Ag in Year 3 (Table 3).

Table 3. Least squares means (± s.e.) of ages and live weights (LW) of early weaned calves at weaning and their final LW (FLW) after 123 days of experimental grazing on improved pastures (early weaned) and of normally weaned calves off savanna.

Pasture

n

Year 3

Year 4

Age (d)

LW (kg) FLW (kg)

Age (d)

LW (kg) FLW (kg)

Andropogon gayanus

10

92 ± 3

63 ± 15

82a1 ± 4

85 ± 4

77 ± 9

84a ± 4

A. gayanus + Centrosema acutifolium

10

93 ± 5

69 ± 9

91ab ± 3

89 ± 5

77 ± 6

88a ± 4

A. gayanus + Pueraria phaseoloides

10

92 ± 4

72 ± 13

95b ± 4

83 ± 5

89 ± 10

92a ± 4

Brachiaria humidicola + Arachis pintoi

10

n.a.

n.a.

n.a.

86 ± 3

80 ± 7

85a ± 5

Savanna2

10

91 ± 5

74 ± 6

117c ± 3

96 ± 10

89 ± 14

129b ± 5

P

NS

NS

<0.0001

NS

NS

<0.0001

n - number of animals.

n.a. - not applicable.

NS - not significant.

1Within columns values followed by different letters differ significantly (P<0.05).

2Weights and ages of savanna calves were obtained on the same date (± 3 days) as those of the early weaners.

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

Grass-legume pastures for early weaning in savanna 57

Data on daily LWG (DLWG, g/head) of calves for

pastures (P<0.0001), associated with a temporary increase

each individual grazing period and for each of the 3 Ag-

in the percent legume. Over the same period, relative to

based pastures in 1986 were pooled and regressed on the

calves grazing Ag as monoculture, hematocrit concen-

amount of green grass leaf on offer (GGL; kg DM/ha).

tration was higher (P<0.001) in AgCa and AgPp mixtures

The resulting linear regression equation was: DLWG =

by 31 and 19%, respectively. However, although within

0.702 ± 0.156 GGL – 166.667 ± 98.700; r2 = 0.63, P<0.01.

normal physiological values, a larger hematocrit differ-

ence (58%; P<0.001) was found with the unweaned

Pilot test

calves on savanna (Table 4).

Concentrations of minerals in serum and feces (Table

Despite the very low LWs of some of the early weaned

5) varied considerably between years (P<0.0001). How-

calves, no deaths occurred. Weight gains during the sown

ever, there were small and mostly non-significant

pasture phase averaged 0.18 ± 0.10 kg/day, while the

differences between pastures. There were no effects of

correlation between weaning weight and subsequent

nutritional treatment upon ash or fecal N, on either a DM

weight gain on the sown pasture was not significant (r2 =

or an organic matter basis (Table 5).

0.24, P>0.05).

Cow live weights and reproductive performance

Blood and feces profiles

Early weaned cows were significantly heavier at weaning

Blood data showed that the concentrations of total

than those weaned at normal times (Table 6; P<0.05).

protein, urea nitrogen and the AST and GGT enzymes in

Inter-calving intervals increased significantly (P<0.05)

weaned calves differed significantly (P<0.0001) between

with increasing calf weaning age, but were inversely

years, but there were much smaller differences in

related to cow weight at weaning (P<0.001), with the

concentrations of protein and enzymes amongst the

positive effect of cow weaning weight in reducing inter-

pastures (Table 4). In Year 3, blood urea nitrogen levels

calving interval being greater with older than with

on the AgPp pasture were 8‒10 times those on the other

younger weaning ages (regression equation in Table 6).

Table 4. Least squares means for total blood protein (TBP), hematocrit (HCT), urea nitrogen (BUN) and the body fluid enzymes aspartate aminotransferase (AST) and gamma-glutamyl transferase (GGT) of early weaned calves grazing sown pastures and unweaned calves grazing savanna in Years 3 and 4.

Pasture

TBP

HCT

BUN

AST

GGT

(g/100 ml)

(g/100 ml)

(g/100 ml)

(IU/ml)

(IU/ml)

Reference values1

6.3–8.9

24–46

6–22

39–79

14–40

Year 3

A. gayanus

5.9 ± 0.14

26.1a ± 1.56 7.4a ± 0.69

76.0 ± 6.00

7.2 ± 7.00

A. gayanus + C. acutifolium

6.2 ± 0.12

34.4b ± 1.38 5.8a ± 0.31

75.0 ± 5.00

7.7 ± 7.30

A. gayanus + P. phaseoloides

6.3 ± 0.13

31.1b ± 1.46 63.5b ± 41

69.0 ± 5.00

3.3 ± 7.40

Savanna

6.8 ± 0.29

41.3c ± 1.47

6.0a ± 0.35 83.0 ± 14.00

42.9± 8.90

Year 4

A. gayanus

6.7 ± 0.53

41.3 ± 1.47 6.6a ± 0.40

80.0 ± 5.00

16.4 ± 8.30

A. gayanus + C. acutifolium

7.6 ± 0.52

n/a

5.9a ± 0.31

82.0 ± 5.00

5.2 ± 8.10

A. gayanus + P. phaseoloides

8.8 ± 0.61

n/a

8.9b ± 0.82

71.0 ± 6.00

1.9 ± 9.35

B. humidicola + A. pintoi

8.0 ± 0.52

n/a

8.8b ± 0.69

84.0 ± 3.00

6.6 ± 8.00

Savanna

6.8 ± 0.20

n/a

7.5b ± 0.81

83.0 ± 8.00

15.0 ± 7.80

Year, Probability

<0.0001

<0.0001

<0.0001

<0.0001

Pasture (year), Probability

= 0.09

<0.001

<0.0001

= 0.09

= 0.25

n/a - not available.

1Aiello and Moses (2016).

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

58 R. R. Vera and C.A. Ramírez-Restrepo

Table 5. Least squares means for blood serum phosphorus and calcium (mg/dl), fecal phosphorus, calcium and ash (% fecal DM, FDM) and fecal nitrogen (% fecal organic matter, FOM) concentrations (means ± s.e.) of early weaned calves grazing sown pastures and un-weaned calves grazing savanna in Years 3 and 4.

Pasture

Serum P

Serum Ca

Fecal P

Fecal Ca

Fecal ash

Fecal N

Year 3

A. gayanus

4.28c1 ± 0.51 8.96 ± 0.23 0.99a ± 0.25 1.24a ± 0.46 13.57a ± 0.93 1.92 ± 0.07

A.gayanus + C. acutifolium

4.17c ± 0.43 8.83 ± 0.21 0.76a ± 0.13 1.03a ± 0.29 12.29a ± 0.85 1.69 ± 0.07

A. gayanus + P. phaseoloides 5.89b ± 0.47 8.75 ± 0.22 0.61a ± 0.08 2.38a ± 1.54 13.08a ± 0.85 1.67 ± 0.07

Savanna

9.78a ± 0.39 10.65 ± 0.24

n/a

n/a

n/a

n/a

Year 4

A. gayanus

4.88a ± 0.44 9.04 ± 0.23 0.64a ± 0.45 0.77a ± 0.84 9.43a ± 1.05 1.79 ± 0.06

A. gayanus + C. acutifolium

5.46a ± 0.42 9.52 ± 0.22 0.61a ± 0.39 0.75a ±0.77 10.58a ± 1.06 1.75 ± 0.05

A. gayanus + P. phaseoloides 5.39a ± 0.49 9.13 ± 0.27 0.32b ± 0.14 0.76a ± 1.07 7.33a ± 1.67 1.87 ± 0.06

B. humidicola + A. pintoi

5.19a ± 0.42 9.71 ± 0.22 0.46b ± 0.25 2.74b ± 5.0 9.20a ± 1.17 1.80 ± 0.06

Savanna

5.93a ± 0.75 10.25 ± 0.30

n/a

n/a

n/a

n/a

Year, Probability

<0.0001

<0.0001

<0.0001

<0.0001

<0.0001

<0.0001

Pasture (year), Probability

<0.0001

NS

<0.05

<0.0001

<0.05*

NS

n/a - not analyzed.

NS - not significant.

1Letters compare pasture values within years. Differences were found between pastures in different years, but not within the same year.

Table 6. Reproductive performance and live weight (LW) of cows (n = 45) in relation to age and weight of calves at weaning.

Weaning age (d)

Inter-calving interval (d)1,2

Cow weaning LW (kg)

Calf weaning LW (kg)

110

472d3 (454–491)

339a ± 8

68a ± 4

166

514c (504–557)

308b ± 11

112b ± 5

198

625a (594–656)

304b ± 11

114b ± 5

266

642a (616–669)

306b ± 9

143c ± 4

1Regression equation:

Inter-calving interval (d) = 394 (± 38) + 2.76 (± 0.59) * calf weaning age – 0.00588 (± 0.0019) * calf weaning age * cow weaning LW; adj R2 = 0.45; P<0.001.

2Confidence interval in parentheses.

3Within columns values followed by different letters differ significantly (P<0.05).

Calf compensatory body growth

Table 7. Final live weights (LW) of contemporary calves (n)

weaned early and grazed on various sown pastures for 123

The possible carry-over effects of low calf weaning LW

days and normally weaned calves, following 414 days

were examined by monitoring subsequent performance of

grazing savanna as a single group. Data are least squares means

early weaned calves compared with normally weaned

± s.e.m.

calves, for a total of 414 days after the end of the early

Weaning pasture

n

LW (kg)

weaning experimental period (Table 7). At that time, there

A. gayanus

10

187 ± 19

were no significant (P>0.05) differences in final LW

A. gayanus plus C. acutifolium

10

187 ± 16

between the early weaned and normally weaned calves

A. gayanus plus P. phaseoloides

10

192 ± 13

(Table 7). Over the 414 days of common grazing in

Traditional weaning on savanna

10

205 ± 16

savanna, daily weight gains were inversely related to

Probability

NS

weaning weight, weight gains to weaning, and weight at

end of the experimental phase (r2 = 0.98, P<0.05 in all

n - number of animals.

cases).

NS - not significant.

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

Grass-legume pastures for early weaning in savanna 59

Discussion

performance of early weaners on limpograss ( Hemarthria

altissima).

The primary objective of this study was to assess the

Casual visual observations showed highly selective

growth of early weaned calves on sown tropical pastures,

grazing on the Ag-based pastures. In earlier studies

while keeping the breeding cows on savanna. A

(Böhnert et al. 1985; 1986), young steers grazing AgPp

secondary objective was to document the effect of early

mixtures selected diets much higher in N than when

weaning (3‒4 months) on inter-calving interval in

grazing Ag as a monoculture but differences in in vitro

breeders compared with traditional rangeland cow-calf

DM digestibility of the diets selected were not significant.

breeding, where calves are weaned at about 9‒10 months

Although some caution is required in extrapolating

old. The low overall mortality rate of calves of 1% over 4

selective grazing behaviors between weaners and 1‒2

consecutive years was a significant finding when

year old steers, the high BUN concentration on the AgPp

compared with commercial, extensive tropical herds that

pasture in Year 3 was likely due to temporary low

normally exhibit death rates of 7‒9% (Kleinheisterkamp

availability of green leaf on Ag and high levels of Pp on

and Habich 1985; Rivera Sánchez 1988). Reduced calf

offer, as Ag leaves were selectively grazed at the start of

mortality would then complement the improved

grazing in each paddock. However, as the availability of

reproductive performance of the breeding cows in terms

Ag leaves rapidly declined, calves were forced to

of reduced inter-calving intervals associated with early

consume the legume. The preliminary, positive relation-

weaning.

ship between green grass on offer on all Ag-based

There has been limited research on management of

pastures and daily LWG points in the same general

early weaners on supplemented tropical pastures

direction.

(Vendramini et al. 2008; Vendramini and Arthington

Overall, this study also showed that none of the early

2008), and even less if unsupplemented. Post-weaning

weaned calves demonstrated a deficiency of total protein,

LW gains by early weaned calves were low and similar

hematocrit, BUN and the sensitive enzyme marker of liver

across the 4 years of experimentation (Tables 2 and 3)

damage, AST (Table 4). However, compared with blood

allowing for the differences between years in age of

reference values, results from all pastures in Years 3 and

calves at weaning. Schottler and Williams (1975)

4 indicated low GGT activity, and high variability

compared the performance of Brahman-Shorthorn

between animals (Table 4). Although the potential

crossbred calves weaned at 4, 5, 6 or 7 months of age on

physiological effects of these values need to be clarified,

a Para grass ( Brachiaria mutica)-Siratro ( Macroptilium

it is reasonable to assume, as demonstrated by Stojević et

atropurpureum) sward for 2 months, followed by a

al. (2005) with healthy dairy cattle, that the observed GGT

buffel

grass

( Cenchrus

ciliaris)-Siratro

pasture.

concentrations reflected a temporary acute situation (i.e.

Regardless of weaning age, LWG ranged between 0.20

circadian changes) associated with age of the calves,

and 0.32 kg/day. Holroyd et al. (1990) weaned calves in

rather than long-term detrimental metabolic effects.

northern Australia at 5 and 8 months of age, and after a

Furthermore, it is unfortunately impossible to define

10-day period of supplementation with good quality

whether the low enzyme concentrations suggested

pasture hay they were transferred to a savanna paddock.

adverse metabolic effects of secondary compounds in the

The LW difference at 8 months between early and late

legumes. Nevertheless, it is worth noting that recent

weaners of 54 kg for males was reduced to 13 kg at the

studies (Ramírez-Restrepo et al. 2016) demonstrated

age of 3.5 years due to compensatory growth in younger

that supplementation of animals with plant-derived

weaned calves, and the research indicated that pasture

compounds increases GGT blood serum values in

quality was probably the limiting factor for better animal

Brahman cattle, which is contrary to the present results.

performance.

Despite differences between pastures and years in

This hypothesis is supported by the observation that

mineral concentrations in serum and feces (Table 4), all

the LW of early weaners (100 days) placed on a higher

values fell within normal ranges (Doornenbal et al. 1988;

quality annual ryegrass ( Lolium rigidum) temperate

Aiello and Moses 2016). The fairly large between-year

pasture did not differ from those of late weaners at 365

differences, and the between-animal variation indicated

days of age (Potter et al. 2004). The close correlation

by the relative magnitude of the standard error terms,

between green leaf and LWG found in the present

question the reliability of single samplings within years

research supports the hypothesis that performance of

and pastures, and by inference, between farming systems

early weaners on pastures may be limited by forage

as frequently carried out in survey studies. Several

quality. In this context, Aguiar et al. (2015) advocated

mineral deficiencies in unsupplemented adult beef cattle

limited creep feeding with soybean meal to improve the

grazing savanna, accompanied by low breeding cow

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

60 R. R. Vera and C.A. Ramírez-Restrepo

LWs, have been reported (Lebdosoekojo et al. 1980).

supported by the literature, would suggest that stress in

Subsequent and detailed analyses by Rivera Sánchez

early weaned beef calves would have been short-term

(1988) in controlled savanna experiments over several

only. Furthermore, and despite the condition of

years showed adequate serum, liver and fecal

generalized under-nutrition, there was no evidence of

concentrations of all minerals in cows, when complete

negative carry-over effects. In fact, by the end of the

mineral mixes were provided. Furthermore, Rivera

observation period, there were no significant differences

Sánchez (1988) found evidence of an interaction of

in the final LWs (Table 7) despite the early advantage of

mineral supplementation with management strategies that

animals allowed to suckle for the normal time, a finding

allowed access to improved grass-dominated pastures by

confirmed by the longer-term on-farm study of Mejía et

the breeding herd.

al. (2009), who weaned female and male calves at

Although fecal N as an indicator of nutrition has

4 months of age on a rotation of B. decumbens,

sometimes been questioned (Hobbs 1987), it is generally

B. humidicola and savanna pastures plus a medium

regarded as appropriate for free-ranging herbivores in the

quality concentrate until reaching 8 months of age.

absence of better, simple indices (Leslie Jr. et al. 2008).

Thereafter these and normally weaned calves (8 months

Similarly, N in feces has been found to be broadly and

old) were raised on the same pastures as above without

linearly related to N intake with a variety of forage diets

supplementation. No significant differences were found

(Nunez-Hernandez et al. 1992). Allden and Jennings

in age at first conception (35 vs. 33.4 months for early and

(1969) proposed that fecal N levels of 1.4‒1.6% in sheep

traditional weaning, respectively). The inter-calving

are indicative of N-limited diets. If these values are

interval of early weaned dams was 141 days shorter than

applicable to calves, data in Table 5 would suggest that

those weaned late. The corresponding males reached

dietary N would not have been the limiting nutritional

slaughter weight (450 kg) with a non-significant

variable. This view accords with that expressed by

difference of 3.2 months and average monthly LWGs of

Lascano (1991), who showed that digestible energy intake

11.5 vs. 10.6 kg for late and early weaners, respectively.

is the most limiting nutrient for yearlings and adult cattle

Under more severe pasture conditions, Holroyd et al.

in neotropical savannas.

(1990) noted that the early weaners (5 months old at

The low calf LWs at the end of the experimental

weaning) grazing savanna in northern Australia were still

periods, coupled with absence of indicators of specific

13 kg lighter than late weaners (8 months old at weaning)

nutritional deficiencies shown by the blood and fecal

at 3.5 years of age.

analyses, are indicative of a general condition of under-

In view of the compensatory growth experienced by

nutrition, a hypothesis supported by the in-depth analyses

the early weaners in the present study, the low LWG

of metabolic profiles of early weaned calves in tropical

recorded on pasture would be acceptable if, as is generally

northeast Argentina. There, Coppo (2003; 2007a; 2007b)

the case, weaners are not sold immediately, but are kept

assessed the stress produced by early weaning at 60 days

as young steers for an additional 12‒18 months. This

of age in crossbred Zebu cattle supplemented with

trade-off between calf LWG and cow reproductive

concentrates, but could not relate it to a large set of blood

performance is an important consideration for farm

parameters and suggested that there was no evidence of

managers who need to balance different forage resources

specific metabolic stresses. Arthington et al. (2005)

and the nutritional needs of stock and prioritize their use.

studied the dynamics of acute-phase proteins in beef

The new ‘crop’ of improved, high-yielding and leafier

calves weaned at 89 days of age onto pasture and fed a

Brachiaria cultivars (Pizarro et al. 2013), together with

supplement, and found them to rise in the first few days

higher quality grasses such as Panicum maximum, may

following separation from the dams, decreasing

help resolve the above issues to some extent, if the

subsequently to normal values. While early weaned

Brachiaria cultivars do not lead to subclinical and clinical

calves were lighter than contemporary suckling calves at

photosensitization (Lima et al. 2012).

120 days of age, and had lower concentrations of BUN,

Notwithstanding few to no weight differences in later

total proteins, triglycerides, P, Mg, Fe and Cu, indicators

life, possible negative effects of low early growth rate on

of stress such as cortisol, aldosterone and AST did not

lifetime beef production of early weaned females through

differ between the 2 groups of calves. The authors

epigenetic effects affecting them and their progeny cannot

concluded that early weaning does not produce clinical

be ignored (Martin et al. 2007; Funston et al. 2012;Wathes

stress in crossbred Zebu calves, despite a general

et al. 2014), although this would probably be a minor

condition of under-nutrition.

concern in extensive systems.

One measure of the sustainability of beef herds is

In our study, early weaned male calves attained

absence of animal stress, and the present results,

weights at 18 months of age similar to those of control

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

Grass-legume pastures for early weaning in savanna 61

animals of equivalent ages. Vera (1991) and Vera et al.

they stopped cycling, but they needed to reach weights in

(1993) showed that, despite long periods of sustained

excess of 320 kg to start cycling again. As shown in Table

under-nutrition, heifers and cows could achieve mature

6, only cows, whose calves were weaned at 110 days,

body sizes and inter-calving intervals similar to those of

showed LWs above the purported lower critical weight of

better-fed animals, when given the opportunity to make

320 kg. Inter-calving intervals increased significantly

moderate compensatory gains. In the present study,

with increasing weaning age and were negatively related

weaning calves early removed lactation stress on cows,

to cows’ weights at weaning (P<0.001; footnote of Table

and allowed them to attain plateau weights of 350‒400 kg,

6), whereby the positive effect of cow weaning weight on

confounded with pregnancy, even during the dry season.

inter-calving interval was greater with larger than with

While LWs of this magnitude are seldom observed in

lower weaning ages (Table 5).

traditional savanna-based grazing systems (Rivera

Lastly, early weaning of beef calves and the

Sánchez 1988), weights of Zebu cows above 320 kg do

consequent changes in reproductive performance will

not appear to limit re-conception (Mukasa-Mugerwa

likely lead to significant changes in herd dynamics

1989; Vera et al. 1993). In fact, cows weaned at 166 days

(Turner et al. 2013), whose production, environmental

in the present research conceived after weaning with an

and economic effects remain to be studied. These changes

average weight of 303 ± 43 kg.

would significantly impact management decisions

Early weaning is known to enhance reproductive

(Sullivan et al. 1997). The savannas of the Orinoco

performance of underfed beef cows (Moore and da Rocha

watershed are under transition as represented by: (i)

1983; Holroyd et al. 1990; Schlink et al. 1994; Coppo et

increase in crop and tree plantation areas; (ii) oil

al. 2002; Arthington et al. 2004), an effect shown also in

exploration and extraction; (iii) mining; (iv) growing

Table 6 that demonstrates the trade-offs between weaning

recognition of indigenous rights and lands; (v) and

age and LWs of calves, LWs of their dams and inter-

increasing appreciation of the savanna’s relevance in

calving intervals. Moore and da Rocha (1983)

terms of biodiversity and contribution to greenhouse gas

investigated the effects of 2 levels of nutrition and 5

emissions (Rausch 2013; CONPES 2014). This transition

weaning ages in Zebu Gyr breed cows fed hay of

has been also noted in the Australian savannas (Holmes

B. decumbens in the Brazilian Cerrados and found that,

2010). This implies that extensive beef cattle farming,

irrespective of the cows’ nutritional level, early weaning

even if it continues to represent an important land use

of calves improved cow weights at different stages

system, will need to adapt and intensify to the extent

throughout the reproductive cycle and subsequent

possible. Further, farmers and their communities will

reproductive performance. Weights of cows at weaning

need to modify their decision-making to take into account

were 313 kg and 325 kg on low and high supplementary

the multifunctional traits of these lands. The strategic use

energy treatments, respectively, and the authors suggested

of areas of sown pasture could play an increasingly

that Zebu cows rarely conceive if suckling cows weigh

important role in management of breeding herds.

less than 300 kg. Weights of cows at weaning decreased

linearly from 352 kg, when calves were weaned at 1

Conclusions

month of age, to 294 kg if calves were weaned at 6 months

of age, with LW losses during lactation increasing from

Results from this study suggest that there is considerable

21 to 102 kg for the respective weaning ages (Moore and

scope and flexibility in strategic use of small areas of

da Rocha 1983).

sown pastures for weaners in combination with extensive

Mukasa-Mugerwa (1989) reviewed the literature

savannas to improve productivity of beef breeding herds,

regarding reproduction of Zebu cattle in the tropics, and

if calves are given the opportunity to realize

noted that tropical cattle dependent on natural pastures

compensatory growth. Several authors have commented

most often calve in alternate years, but animals with

on the flexibility, adaptability and sustainability of

access to good quality sown pastures have improved

extensive systems (Davies et al. 2010; Astigarraga and

reproductive performance. For example, Rivera Sánchez

Ingrand 2011), while some of the environmental aspects

(1988) reported inter-calving intervals of 618 days on

have been also described (Ramírez-Restrepo and

well managed savannas over 4 years, whereas Vera et al.

Charmley 2015) or are under scrutiny (Ramírez-Restrepo

(2002) found an average interval of 445 days on well

et al. unpublished data 2017). This study has demon-

managed B. decumbens. Equally large differences have

strated over 4 consecutive years, that early weaning of

been reported by other authors (Arthington et al. 2004).

calves onto a variety of sown tropical pastures is

Hale (cited by Mukasa-Mugerwa 1989) found that, when

technically feasible, resulting in improved LWGs and

the LW of suckling Zebu cows fell from 390 to 320 kg,

reproductive rates in their dams.

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

62 R. R. Vera and C.A. Ramírez-Restrepo

Adoption of such early weaning strategies in extensive

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Acknowledgments

chemical composition of the diet selected by fistulated steers

under grazing on improved grass-legume pastures in the

This research was financially supported by the

tropical savannas of Colombia. I. Botanical composition of

International Center for Tropical Agriculture (CIAT)’s

forage available and selected. Journal of Animal Breeding

core funding in Colombia. The authors express our

and Genetics 102:385–394. DOI: 10.1111/j.1439-0388.

appreciation to Hernando Ayala, DVM (deceased), who

1985.tb00707.x

supervised the first 2 years of the experiment. We further

Böhnert E; Lascano C; Weniger JH. 1986. Botanical and

chemical composition of the diet selected by fistulated steers

acknowledge Obed García Durán, DVM, who confirmed

under grazing on improved grass-legume pastures in the

some of the views and opinions expressed here, and

tropical savannas of Colombia. II. Chemical composition of

facilitated a recent trip by the senior author to the region

forage samples of forage available and selected. Journal of

to verify some of the assertions made in the paper. We

Animal Breeding and Genetics 103:69–79. DOI:

thank staff at CIAT and Carimagua Research Station for

10.1111/j.1439-0388.1986.tb00068.x

their help and technical support with all aspects of this

CONPES. 2014. Política para el desarrollo integral de la

work. Finally, special thanks are extended to the

Orinoquia: Altillanura - Fase 1. Documento 3797. Consejo

Commonwealth Scientific and Industrial Research

Nacional de Política Económica y Social (CONPES),

Organisation (CSIRO) for allowing the time to the second

Departamento Nacional de Planeación (DNP), República de

author to co-write the manuscript.

Colombia, Bogotá, Colombia. https://goo.gl/5IXl2S

Coppo JA. 2003. Early weaning as cause of malnutrition

in

half-bred

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calves.

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(Received for publication 27 October 2016; accepted 30 April 2017; published 31 May 2017)

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