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beef breeding herds have been replaced by yearlings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
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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
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
Zebu
calves.
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(Received for publication 27 October 2016; accepted 30 April 2017; published 31 May 2017)
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