Suckling behaviour of Nelore, Gir and Caracu calves and their crosses

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Suckling behaviour of Nelore, Gir and Caracu calves and their crosses Mateus Jose ´ Rodrigues Paranhos da Costa a,b,c, * , Lu ´cia Galva ˜o Albuquerque b,c , Joanir Pereira Eler c,d , Josineudson Augusto II de Vasconcelos Silva e a ETCO-Grupo de Estudos e Pesquisas em Etologia e Ecologia Animal, Brazil b Departamento de Zootecnia, Faculdade de Cie ˆncias Agra ´rias e Veterina ´rias, UNESP, 14884-900 Jaboticabal, SP, Brazil c Pesquisador CNPq, Brazil d Departamento de Cie ˆncias Ba ´sicas, Faculdade de Zootecnia e Engenharia de Alimentos, USP, Pirassununga, SP, Brazil e Estac ¸a ˜o Experimental de Zootecnia de Serta ˜ozinho, IZ-SAA-SP, Serta ˜ozinho, SP, Brazil Accepted 20 February 2006 Available online 18 April 2006 Abstract It is reasonable to assume that the knowledge of suckling behaviour contributes to optimal management and selection of beef cattle. However, there is little information about suckling behaviour of some beef cattle breeds. The aim of this study was to describe the suckling behaviour of two zebu (Bos indicus) and one criollo (Bos taurus) breeds, analysing the potential effects of breed and some environmental factors on suckling frequency and duration. Forty cows, 17 Nelore, 14 Gir (both zebu) and 9 Caracu (criollo) were bred in a diallelic crossing design. The cows and resulting calves were kept on pasture from birth to weaning. Their behaviour was recorded weekly during daylight. Three behavioural traits were considered: number of suckling meals (NSM), duration of each suckling meal (DSM) and total suckling duration (TSD). Allosuckling was not observed. The calves suckled at any time during the daylight and the overall means were: NSM = 2.57 0.05 meals/12 h (from back transformed data), DSM = 9.25 0.11 min/suckling meal and TSD = 23.76 0.47 min/12 h. There was an effect of dam’s breed on NSM and DSM; the calf’s genetic group within breed of cow influenced NSM and TSD when the dams were from www.elsevier.com/locate/applanim Applied Animal Behaviour Science 101 (2006) 276–287 * Corresponding author. Tel.: +55 16 32092678; fax: +55 16 32092684. E-mail address: [email protected] (M.J.R. Paranhos da Costa). 0168-1591/$ – see front matter # 2006 Elsevier B.V. All rights reserved. doi:10.1016/j.applanim.2006.02.006

Transcript of Suckling behaviour of Nelore, Gir and Caracu calves and their crosses

Suckling behaviour of Nelore, Gir and

Caracu calves and their crosses

Mateus Jose Rodrigues Paranhos da Costa a,b,c,*,Lucia Galvao Albuquerque b,c, Joanir Pereira Eler c,d,

Josineudson Augusto II de Vasconcelos Silva e

a ETCO-Grupo de Estudos e Pesquisas em Etologia e Ecologia Animal, Brazilb Departamento de Zootecnia, Faculdade de Ciencias Agrarias e Veterinarias,

UNESP, 14884-900 Jaboticabal, SP, Brazilc Pesquisador CNPq, Brazil

d Departamento de Ciencias Basicas, Faculdade de Zootecnia e Engenharia de Alimentos,

USP, Pirassununga, SP, Brazile Estacao Experimental de Zootecnia de Sertaozinho, IZ-SAA-SP, Sertaozinho, SP, Brazil

Accepted 20 February 2006

Available online 18 April 2006

Abstract

It is reasonable to assume that the knowledge of suckling behaviour contributes to optimal

management and selection of beef cattle. However, there is little information about suckling

behaviour of some beef cattle breeds. The aim of this study was to describe the suckling behaviour

of two zebu (Bos indicus) and one criollo (Bos taurus) breeds, analysing the potential effects of

breed and some environmental factors on suckling frequency and duration. Forty cows, 17 Nelore,

14 Gir (both zebu) and 9 Caracu (criollo) were bred in a diallelic crossing design. The cows and

resulting calves were kept on pasture from birth to weaning. Their behaviour was recorded weekly

during daylight. Three behavioural traits were considered: number of suckling meals (NSM),

duration of each suckling meal (DSM) and total suckling duration (TSD). Allosuckling was not

observed. The calves suckled at any time during the daylight and the overall means were:

NSM = 2.57 � 0.05 meals/12 h (from back transformed data), DSM = 9.25 � 0.11 min/suckling

meal and TSD = 23.76 � 0.47 min/12 h. There was an effect of dam’s breed on NSM and DSM;

the calf’s genetic group within breed of cow influenced NSM and TSD when the dams were from

www.elsevier.com/locate/applanim

Applied Animal Behaviour Science 101 (2006) 276–287

* Corresponding author. Tel.: +55 16 32092678; fax: +55 16 32092684.

E-mail address: [email protected] (M.J.R. Paranhos da Costa).

0168-1591/$ – see front matter # 2006 Elsevier B.V. All rights reserved.

doi:10.1016/j.applanim.2006.02.006

the Nelore breed. The age of calf had significant effects on all traits. Males averaged higher NSM

and TSD (2.60 � 0.03 meals and 25.05 � 1.37 min/12 h, respectively) than females (2.12 � 0.04

meals and 21.51 � 1.55 min/12 h, respectively). The differences in suckling behaviour seem to be

produced by a complex combination of genetic and environmental factors, which result in a

particular behavioural relationship within mother–offspring pairs.

# 2006 Elsevier B.V. All rights reserved.

Keywords: Cattle; Cow–calf relationship; Milk transfer estimation; Breed differences; Suckling

1. Introduction

The importance of maternal influence on offspring’s phenotype has been recognised in

mammals since early animal husbandry. Milk production has been considered the main

cause of maternal effects in cattle (Eler et al., 1989). However, there is a large variation in

the correlation coefficient estimates between milk production of the dam and the

performance of the offspring from birth to weaning, ranging from 0.29 to 0.90 (Neville,

1962; Rutledge et al., 1971; Totusek et al., 1973; Robison et al., 1978; Chenette and Frahm,

1981; Clutter and Nielsen, 1987).

It is also recognised that knowledge of the nursing–suckling relationship has practical

significance, contributing to improve management and selection of cattle (Craig, 1981;

Paranhos da Costa and Cromberg, 1998). Although the obvious function of suckling is to

provide nutrients to the calf, it is also important for providing antibodies and improving

welfare through colostrum and maternal contact, respectively. Therefore, it is expected that

other important factors affect the performance of the calf besides the dam’s milk

production, and that the whole care and protection offered by the dam to offspring during

the suckling period should also be considered part of the maternal influence (Poindron and

Le Neindre, 1979; Toledo et al., 2003).

Important breed differences in suckling behaviour have been reported in several

studies (Nicol and Sharafeldin, 1975; Le Neindre, 1989; Alencar et al., 1995; Das et al.,

2000; Espasandin et al., 2001). Usually, these differences are associated with the

mother’s milk production. However, there is not a consensus on this last statement. The

breed differences in suckling behaviour could also result from non-nutritive suckling (de

Passile and Rushen, 1997), which, as described by Lidfors et al. (1994), represented a

high percentage (between 30.3 and 50.0%) of the total suckling bout duration of beef

cattle.

There are many papers addressing the behaviour of cows and calves from birth

to weaning, but most of them have addressed the issue in just a few breeds of Bos

taurus. The information about suckling behaviour of Bos indicus (zebu) and criollo

breeds is still scarce, especially for those reared in Brazil. The aim of this study was

to describe the suckling behaviour of Nelore, Gir and Caracu cattle and their

crosses, addressing its role in milk transfer estimation and analysing the poten-

tial effects of breed and some environmental factors on suckling frequency and

duration.

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2. Animals and methods

The present study was carried out at the University of Sao Paulo campus of

Pirassununga, State of Sao Paulo, Brazil (218590S 478210W and 560 m of altitude). Forty

cows and their respective calves were observed with a variable frequency (ranging from 11

to 27 days).

The cows were bred in a ‘‘diallel crossing design’’, involving the three breeds mentioned

above. Therefore, all of the cows were pure-breds (being 14 Gir; 9 Caracu and 17 Nelore

breed) and the calves were either, pure (N = 22; being 8 Gir, 5 Caracu and 9 Nelore) or cross-

bred (N = 18; being 2 Gir-Caracu, 4 Gir-Nelore, 3 Caracu-Gir, 4 Caracu-Nelore, 3 Nelore-Gir

and 2 Nelore-Caracu). Nelore and Gir are zebu breeds (Bos indicus) first taken to Brazil in the

19th century and Caracu is a Bos taurus originated in Brazil through crossing of Iberian cattle

(Felius, 1985), which have been introduced into the country since the beginning of the 16th

century. The calving season was from November to March and the observations were carried

out from February to September. Each cow and its calf were identified with the same coloured

rubber collar. All the animals were kept together in pastures and the calves had free access to

suckling during the whole period of observation.

The behavioural data were collected by continuous, direct observations (Martin and

Bateson, 1993). We observed all occurrences of suckling and attempts to suckle, recording

the time of the beginning and ending of each occurrence. An attempt to suckle was

recorded when a calf tried to get a teat into its mouth without success, and suckling when a

calf succeeded in getting a teat into its mouth for any detectable period of time combined

with observable swallowing. The cow-calf pair identification (based on the colour of

collars) and the date of observation were also recorded.

Observations were performed by three people (previously trained) riding a horse with

the aid of binoculars. They were conducted during the daylight period (from 06:00 to

18:00 h) on 28 days. The date of birth of cows and calves, sex and genetic composition of

calves, were obtained from the farm files.

Milk intake was estimated monthly from the 30th to 180th days after calving. The

method of weighing the calf just before and after a suckling meal was used. The method

comprised the following steps: (1) cows and calves were separated at 11:00 h; (2) at

16:00 h the calves were weighed and immediately allowed to suckle their mothers during

approximately 30 min; (3) they were separated again overnight; (4) at 08:00 h the next day

the calves were weighed before suckling; (5) just after weighing they were allowed to

suckle; (6) the calves were reweighed just after suckling. It was assumed that the difference

between the calves’ weight (before and after the second suckling) estimated the normal

milk intake during the 16-h period. The milk production per lactation was estimated by

multiplying the milk intake on sampling days by the number of days between consecutive

milk intake records.

2.1. Data management and statistical analysis

The data file was composed by 774 daily records, characterizing the following variables:

number of suckling meals (NSM), average duration of each suckling meal (DSM) and total

suckling duration (TSD—total time spent suckling in 12 h).

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Repeated measures analysis of variance using the MIXED procedure of the SAS

software package (SAS, 2001) was applied and the model considered the fixed effects of

dam’s breed, calf’s genetic group within the dam’s breed; month of birth, sex of calf, age of

cow at calving and age of calf at the day of observation on NSM, DSM and TSD. Linear

effects of calf’s birth weight and dam’s milk production were also considered, as

covariates. The number of suckling meals was transformed into square root of (NSM + 0.5)

to meet the requirements of normal distribution.

3. Results

Two thousand one hundred ninety-one responses of nursing-suckling interactions from 40

cows and their calves were recorded. Ninety-three percent of them (2038 records) were

effective suckling and the remaining seven percent (153 records) referred to attempts of

suckling in which the calves, for various reasons, did not succeed. The lack of success in

suckling, in spite of the attempts, occurred mainly due to rejection by the cows (62.1%), or

when the calves stopped their attempts before suckling (8.5% of the cases) and when some

other animal disturbed those attempts (1.3%). In 28.1% of the cases the reason for terminating

the attempt could not be determined. All nursing–sucking interactions involved mother–

offspring interactions; allosuckling (a calf suckling from an alien cow) was not observed.

The calves suckled at any time during the daylight, but three peaks of suckling meals

were noted: from 6:00 to 7:00, from 10:00 to 12:00 and from 17:00 to 18:00 h. The overall

means were: NSM = 2.57 � 0.05 suckling meals/12 h (from back transformed data),

DSM = 9.25 � 0.11 min/suckling meal and TSD = 23.76 � 0.47 min/12 h.

There was an effect of dam’s breed on NSM (P < 0.05) and DSM (P < 0.01), but not

TSD (P > 0.05). The NSM means (back-transformed data) of Gir and Nelore dams

(2.56 � 0.04 and 2.53 � 0.03 suckling meals/12 h, respectively) were higher than the

mean of Caracu dams (2.03 � 0.07 suckling meals/12 h). Regarding mean durations of

suckling meals (DSM) the results were the inverse, with Caracu dams presenting higher

mean than Nelore and Gir dams. The least square means of NSM (transformed data), DSM

and TSD, according to the dams breed, are presented in Fig. 1.

The calf’s genetic group within breed of cow influenced NSM and TSD traits only when

the dams were from Nelore breed. The adjusted means according to the calf’s genetic group

are presented in Fig. 2. The highest NSM was presented by pure-bred Nelore–Nelore

calves, which differed significantly (P < 0.05) from the Gir–Nelore and Caracu–Nelore

means. The same order was found for TSD means, and there were significant differences

between the three means (P < 0.05, Fig. 2C).

Age of calf, which corresponds to the stage of lactation, had significant effects

(P < 0.01) on NSM and TSD, but not DSM. NSM decreased continuously with age of calf.

On the other hand, means of TSD presented a light increase until the calf reached 60–90

days of age, decreasing continuously after that (Fig. 3B).

The sex of the calf affected NSM (P < 0.01) and TSD (P < 0.05), but not DSM. Males

presented higher NSM (2.60 � 0.03 suckling meals/12 h; back transformed data) and TSD

(25.05 � 1.37 min/12 h) than females (2.12 � 0.04 suckling meals/12 h and 21.51 � 1.55

min/12 h, respectively).

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Month of birth, age of cow at calving, calf birth weight and dam’s milk production did

not affect (P > 0.05) any of the three traits. Significant coefficients of correlation

(P < 0.05) of milk production with NSM (r = 0.23) and TSD (r = 0.14) were found only for

Nelore breed and with DSM only for Gir (r = 0.25).

4. Discussion

In this study the suckling activities occurred at any time during the daylight, being more

frequent early in the morning, just before noon and late in the afternoon. Similar results

were found in previous researches (Reinhardt and Reinhardt, 1981; Lewandrowski and

Hurnik, 1983; Odde et al., 1985).

According to our results the suckling behaviour of the Nelore, Gir and Caracu cattle

were influenced by various factors, genetic and environmental. In general, these results are

in agreement with those found in earlier studies, describing breed differences on suckling

behaviour (Alencar et al., 1995; Das et al., 2000; Espasandin et al., 2001) and reporting, for

example, that suckling behaviour was influenced by sex of calves and calf’s age (Reinhardt

and Reinhardt, 1981; Lidfors et al., 1994). Specific discussions about these effects are

presented next.

4.1. Descriptive aspects of suckling frequency and duration

The estimated means of NSM, DSM and TSD were slightly lower than those presented

by Nicol and Sharafeldin (1975), who observed 24 mother–offspring pairs of Angus (pure

and cross-bred) cattle from dawn till dusk over the period mid-August to mid-November in

New Zealand. They reported NSM between 3.0 and 3.5 times, DSM between 10 and

11 min and TSD from 30 to 35 min. On the other hand they were slightly higher than those

described by Alencar et al. (1995) for Canchim and Canchim � Nelore cattle

(NSM = 1.6 � 0.0 and 1.9 � 0.0 times/12 h, DSM = 8.2 � 0.2 and 7.0 � 0.0 min/suckling

meal and TSD = 18.3 � 0.7 and 20.6 � 0.7, respectively); Espasandin et al. (2001) for pure

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Fig. 1. Least square means and standard error of square root of number of suckling meal (NSM), duration of each

suckling meal (DSM) and total suckling duration (TSD) according to the breed of cow. Least squares means of the

same trait with common letters do not differ statistically (P > 0.05).

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Fig. 2. Least square means and standard error of square root of number of suckling meals (NSM), duration of each

suckling meal (DSM) and total suckling duration (TSD) for calves mothered by Gir (A), Caracu (B) and Nelore

(C) cows. Least squares means of the same trait with common letters do not differ statistically (P > 0.05).

and cross-bred Nelore cattle (DSM ranging from 6.33 to 7.96 and TSD between 9.28 and

20.05 min/12 h). The mean of DSM from the present study was comparable to others

described in literature for European and Zebu cattle, from 8 to 9 min (Reinhardt and

Reinhardt, 1981; Kiley-Worthington and de La Plain, 1983).

In spite of the main conclusion of Cameron (1998), that ‘‘the assumption that milk

transfer can be measured by time spent suckling has inadequate empirical foundation’’,

we hypothesise that descriptive information of suckling behaviour must have practical

value. In reference to the animal breeding approach, it is common to assess the maternal

effects of the beef cows on calves’ performance by estimating the dam’s milk production.

Usually, this estimation is carried out by weighing the calves, once or twice a day, before

and after suckling (Albuquerque et al., 1993; Alencar et al., 1995; Espasandin et al., 2001),

as described previously in this paper. The precision of this measurement as a milk

production indicator is not completely clear cut, since in this case milk production is

limited by the capacity of udder storage. The milk production of cows presenting udders

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Fig. 3. Least square means and standard error of square root of number of suckling meal (NSM), duration of each

suckling meal (DSM) and total suckling duration (TSD) according to classes of dam’s (A) and calf’s age (B). Least

squares means of the same trait with common letters do not differ statistically (P > 0.05).

with low storage capacity could be underestimated, just by weighing their calves only

once or twice a day.

This statement is based on empirical data. The NSM means from our data, recorded only

during the 12 daylight hours, were always higher than two. Similar results were found by

other authors. Nicol and Sharafeldin (1975), working during the daylight with crossed beef

cattle in New Zealand, reported that the calves presented a suckling frequency ranging

from 3.5 to 5.6 suckling meals/day at 120 and 7 days of age, respectively. Reinhardt and

Reinhardt (1981), studying zebu cattle in Kenya, found 4.8 as the average number of

suckling meals per 24 h; Odde et al. (1985), found an average of 5.0 suckling meals/24 h.

Besides, in our study Nelore and Gir herds presented higher NSM means and lower DSM

means than Caracu. This might explain the breed differences in milk production estimated

by Albuquerque et al. (1993) for Caracu (1077.76 � 134.43 kg), Gir (406.44 � 106.97 kg)

and Nelore breed (631.92 � 90.69 kg), considering 207 days of lactation. Although, some

of the coefficients of correlation of milk production with NSM and DSM were significantly

different from zero, they were always lower than 0.25, characterizing a weak association of

milk production (estimated by the weighing-suckling-weighing method) with suckling

frequency and duration. These results together led us to hypothesise that a calf could get the

same amount of milk from one cow with a low capacity of milk storage, by suckling more

frequently than a calf from a higher udder capacity cow.

We did not observe a single event of allosuckling. Previous studies described the

occurrence of allosuckling in cattle, ranging from 3% (Lewandrowski and Hurnik, 1983

and Das et al., 2000) to 19.02% of the total suckling events (Vıchova and Bartos, 2005).

There are strong evidences that, under restricted milk and space conditions, the probability

of allosuckling increases (Packer et al., 1992; Murphey et al., 1991, 1995). This is, in some

extension, supported by the comparison between the present results and those found by

Lewandrowski and Hurnik (1983), Das et al. (2000) and Vıchova and Bartos (2005). In our

study the animals were kept in free-range conditions, the cows were not milked and the

calves had free access to suckle; in their studies cows and calves had some restriction in

space and milk availability.

4.2. Effect of genetic groups on suckling frequency and duration

In relation to the effects of dams’ breed we have found that the highest NSM mean was

from the breed with the lowest DSM mean, and vice-versa. This could explain the lack of

significance of the dam’s breed effect on TSD, since this variable is a product of NSM and

DSM. This result disagrees with those presented by Das et al. (2000), who found higher

duration and number of suckling meals in zebu than in cross-bred cow–calf pairs, and

significant effects of the breed of cow-calf pairs on the total duration of suckling time.

As pointed out by Nicol and Sharafeldin (1975) breed differences in suckling behaviour

could be attributed to milk production. However, this was not the case in a study with Salers

and Friesian cows–calves pairs carried out by Le Neindre (1989), leading him to conclude

that the total suckling time should result from the need for mother–young physical contact

instead of from the milk production. Both conclusions might be correct, but certainly

incomplete. In our understanding the differences in suckling behaviour among breeds

could result from the combination of similarities and the differences in many traits that are

M.J.R. Paranhos da Costa et al. / Applied Animal Behaviour Science 101 (2006) 276–287 283

involved with physiological and psychological states, and are largely dependent on the

environmental circumstances.

As reported previously, significant differences among the calves’ genetic group were

found depending on the dams’ breed. The calves’ genetic group influenced all variables

when the dams were from Nelore breed. There were no differences among calves mothered

by Gir and Caracu cows. We did not find any logical explanation for such variation among

calves’ genetic group within dams’ breed.

The expectation of a change in suckling behaviour of cross-bred calves to sustain an

increase in nutritional demand promoted by a possible effect of individual heterosis was

not confirmed here, since their suckling frequency and duration were not higher than those

presented by the pure-bred calves. These results disagree with some of those presented

previously. For example, higher DSM means for crossbred than for pure-bred calves were

found by Nicol and Sharafeldin (1975), who proposed that the increase of DSM could

stimulate milk production and consequently proportionate a higher growth rate.

A comparison between pure and crossed Canchim and Canchim–Nelore calves

(Alencar et al., 1995) showed higher NSM for crossed than for pure calves (11.1%), and

lower DSM (14.6%). The difference between TSD means was non significant (18.3 � 0.7

and 20.6 � 0.7 min for pure and crossed calves, respectively). However, Espasandin et al.

(2001) did not find the same tendency when comparing the suckling behaviour of pure and

crossed Nelore calves (Canchim � Nelore, Angus � Nelore and Simental � Nelore) and

their dams’ milk production. The highest mean of DSM was from Simental � Nelore,

which did not differ significantly from the mean of pure Nelore in an intensive system. The

authors did not find significant effects of calves’ genetic group on NSM and TSD.

4.3. Environmental effects on suckling frequency and duration

Several researchers have reported the effects of the age of the calf (or stage of lactation) on

suckling behaviour (Walker, 1962; Nicol and Sharafeldin, 1975; Reinhardt and Reinhardt,

1981; Kiley-Worthington and de La Plain, 1983; Lidfors et al., 1994; Das et al., 2000). In the

present study DSM presented a slight increase between 30 and 60 days of age and remained

relatively constant until 120 days of age, decreasing slightly after this age. The TSD increased

initially, probably due to the increase in DSM, but after 90 days of age it followed the trend

observed for NSM, reducing with the age of calf. Similar results were described by Nicol and

Sharafeldin (1975), reporting an increase in the time spent in each suckling from the 1st to

35th day of life and a decrease in number of sucklings with increasing calf age. However,

other authors found different tendencies in suckling behaviour according to the calf’s age:

Hafez and Lineaver (1968) described an increase in the frequency of suckling from birth to

weaning, Lidfors et al. (1994) observed that the duration of suckling bout decreased with the

calf’s age and the number of suckling bouts increased between 7 and 65 days of age; Das et al.

(2000) reported a continuous decrease in frequency and duration of suckling from birth to 6

months postpartum and Espasandin et al. (2001) did not find any effect of calf’s age on

suckling frequency but the duration of each suckling meal decreased from 60 to 180 days of

age, following the same tendency of milk yield.

The sex of calf significantly affected NSM and TSD, but not DSM. This could be related to

the calves’ weight, since, on average, male calves were heavier than female. These results are

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different from those reported by Nicol and Sharafeldin (1975), Reinhardt and Reinhardt

(1981) and Espasandin et al. (2001). These authors did not find significant effects of sex on

suckling behaviour.

Our results lead us to hypothesise that male calves had a higher demand for milk than

the female calves, resulting in a significant increase in NSM, which led to an increase in

TSD and probably an increase in milk ingestion. This last statement is based upon the

well-known effect of increasing the milking frequency on milk production of dairy cows

(Klei et al., 1997). It is also in agreement with the hypothesis proposed by Trivers (1972),

assuming, for polygynous species like cattle, that natural selection favours parents that

invest more in males, since they present greater variance in reproductive success than

females. We did not find any result addressing the milk production of beef cattle which

could sustain this hypothesis. Instead, some studies with beef cattle (Albuquerque et al.,

1993 and Espasandin et al., 2001) did not find any effect of calf’s sex on milk production.

A similar result was found for water buffalo (Bubalus bubalis), with no difference in

milk production of cows mothering male and female calves (Paranhos da Costa et al.,

2000).

On the other hand, there are some papers showing the higher maternal investment in

male than in female offspring. Alley et al. (1995) studying feral goats (Capra hircus),

showed that single males suckled more frequently than single females, and they also

exhibited a higher growth rate. There are other results supporting the idea of differential

maternal investment based on the sex of offspring: bull calves were heavier at birth and

weaning time (Albuquerque et al., 1993), presented a higher weight gain (Paranhos da

Costa et al., 2000) and were naturally weaned later than heifer calves (Reinhardt and

Reinhardt, 1981).

Considering the results presented here and those reported previously we conclude that

the differences in suckling behaviour cannot be explained by a single isolated underlying

factor. It is more likely that differences in suckling behaviour are produced by a complex

combination of genetic and experiential factors, which would result in a particular

behavioural style for a mother–offspring pair. Nevertheless, under practical conditions, it

could be important to identify specific underlying factors to explain major effects of

suckling behaviour on calf survival or development.

Acknowledgements

We are grateful to Alan Bitu Alves, Andrei K.M. de Castro, Anselmo Bodenmuller,

Marcos V.B. da Silva, Ricardo B. Tonetto and Roberto Carlos de Souza for their help

during data collection. Financial supports were given by CNPq (Conselho Nacional de

Desenvolvimento Cientıfico e Tecnologico) and Fapesp (Fundacao de Amparo a Pesquisa

do Estado de Sao Paulo).

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