- Email: [email protected]
Behaviour of dairy calves suckling the dam in a barn with automatic milking or being fed milk substitute from an automatic feeder in a group pen
Applied Animal Behaviour Science 117 (2009) 150–158
Contents lists available at ScienceDirect
Applied Animal Behaviour Science journal homepage: www.elsevier.com/locate/applanim
Behaviour of dairy calves suckling the dam in a barn with automatic milking or being fed milk substitute from an automatic feeder in a group pen Sofie Fro¨berg a,*, Lena Lidfors b a b
Department of Animal Nutrition and Management, Swedish University of Agricultural Sciences (SLU), Kungsa¨ngen Research Centre, SE-753 23 Uppsala, Sweden Department of Animal Environment and Health, SLU, P.O. Box 234, SE-532 23 Skara, Sweden
A R T I C L E I N F O
A B S T R A C T
Article history: Accepted 16 December 2008
The aim of this study was to investigate the suckling behaviour of dairy calves allowed to suckle freely in a barn with an automatic milking (AM) system and how their behaviour differed from the behaviour of calves given milk substitute from an automatic feeder. For the first 8 weeks of life, calves of the dairy breed Swedish Red were either suckling freely (FS, n = 16) from the dam in an AM barn or kept in groups with automatic feeding (HM, n = 18) with an allowance of 9 kg of milk substitute per day. All treatments had concentrate and hay ad lib. Focal calves in FS and HM treatments were observed one day in weeks 2, 4 and 8, at 08:00–10:00 and the FS calves’ suckling behaviour was observed for a 24-h period once weekly. The HM calves moved, ate concentrate and ate hay more than the FS calves, whereas the FS calves lay down and ruminated more than the HM calves. Crosssucking and tongue-rolling were displayed in HM but not in FS calves. Of the suckling bouts recorded in FS calves 80% was on the dam. Four calves were recorded suckling solely on their dam, but most of the calves were observed attempting to suckle or suckling on other cows in the group once or twice. The total suckling time and number of suckling bouts during week 2–8 on all cows decreased with increasing calf age. In conclusion, the free-suckling calves ate less solid feed, rested more and exhibited fewer non-nutritive oral and abnormal behaviours compared to the calves fed milk substitute from an automatic feeder, and some calves often suckled on other cows. ß 2009 Elsevier B.V. All rights reserved.
Keywords: Dairy calves Behaviour Free suckling Automatic milk feeding Automatic milking system
1. Introduction Herd sizes in industrialized countries are increasing, e.g. in 1990 the Swedish mean herd size was 22 cows, while in 2007 it was 51 (Swedish Dairy Association, 2008). As an effect of increased herd sizes and technology the number of calves kept in groups with automatic milk feeding is likely to increase. Group pens facilitate movement, play and social contact among calves. However, behavioural problems such as calves sucking on body parts
* Corresponding author. Tel.: +46 18 67 16 79; fax: +46 18 67 29 46. E-mail address: sofi[email protected] (S. Fro¨berg). 0168-1591/$ – see front matter ß 2009 Elsevier B.V. All rights reserved. doi:10.1016/j.applanim.2008.12.015
of other calves (cross-sucking) can be common in calves kept in groups (Stephens, 1982; Lidfors, 1993; Fro¨berg et al., 2008). Cross-sucking may facilitate disease transmission (de Passille´, 2001). Cross-sucking and other nonnutritive oral behaviours such as licking objects are less common in calves allowed to suckle the dam (Krohn et al., 1999; Fro¨berg et al., 2008) or being with the dam without access to suckle (Krohn et al., 1999). Wiepkema et al. (1987) proposed that depriving a calf of sucking a teat might cause stress or conflict that could develop into stereotypies. Other studies (Sambraus, 1985; Sato et al., 1994) also suggest that stereotypies may be related to insufficiently stimulated or suppressed feeding behaviours. Moreover, the presence of the cow may be
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
important for social learning (Krohn et al., 1999). Le Neindre and Sourd (1984) found a higher social activity in heifers that had been reared by their dams the first 10–12 weeks compared to those reared artificially. The function of an attachment between parent and young is to ensure that food, warmth and protection is provided for one’s own young and not to others (Gubernick, 1981). Parental attachment is found in species living in larger groups in which the young are precocial, but, in herds where the females are related a combination of attachment and cooperative care of their young may have evolved (Gubernick, 1981). The suckling behaviour of calves that have continuous access to their dams has been observed in several studies, e.g. on semi-wild Maremma cattle (Bos primigenius taurus, Vitale et al., 1986), pure beef or beef and dairy-beef crosses (Nicol and Sharafeldin, 1975; Lidfors et al., 1994; Waltl et al., 1995; Vı´chova´ and Bartosˇ, 2005) and Zebu cattle and their crosses (Reinhardt and Reinhardt, 1981). There are only few studies on free suckling dairy calves and most of them focus on the first days after parturition with a few exceptions, e.g. Ylipekkala (1990). Behavioural variations between breeds may be expected due to different selection and breeding goals among the breeds. In general beef cattle production systems encourage and benefit from strong maternal behaviour, whereas there may be a selection against it in dairy production, as the calf is most often separated from its dam shortly after birth. It is also likely that a low attachment of the dam to its calf is linked to easy milking in high milk-yielding breeds (Le Neindre, 1989). The calves’ suckling behaviour may also be affected by several factors, i.e. the cow’s milk production, if the cow is milked during the milk feeding period, and whether the calf has free or restricted access to suckle. In a study on free suckling in loose housing with parlour milking it was concluded that the system functioned satisfactorily, but no systematic behavioural studies were conducted (Grøndahl et al., 2007). As far as we know there are no previous scientific publications on raising calves with their dams in loosehousing with an automatic milking (AM) system. The aim of this study was to investigate the suckling behaviour of dairy calves allowed to suckle freely in an AM barn and how their behaviour differed from the behaviour of calves given milk substitute from an automatic feeder. The first hypothesis was that due to the supposed relaxation of the mother–young bond in dairy cattle the calves would suckle on other cows than their dams and when they suckle on other cows their position would not be reverse parallel position. The second hypothesis was that free-suckling calves would be more active but exhibit less non-nutritive behaviour such as cross-sucking than calves fed milk from an automatic feeder. 2. Materials and methods 2.1. Animals and experimental design The present study was carried out at the Swedish University of Agricultural Sciences, Kungsa¨ngen Research Centre, Uppsala. Calves of the Swedish Red dairy breed single-born from August to March were studied from birth
151
to weaning at 8 weeks of age. The cows giving birth to the calves ranged from the first to the fifth lactation. In order of birth calves were allocated to each of the two following treatments; milk substitute from an automatic feeder (HM) and free suckling (FS). When up to 12 calves had been allocated to the first treatment, calves were assigned to the next treatment. Later calves were allocated to a second replicate resulting in a total of 23 (replicate 1; n = 11, replicate 2; n = 12) calves in HM and 18 (replicate 1; n = 8, replicate 2; n = 10) calves in FS treatment. In the FS treatment there were seven primiparous and nine multiparous cows. Four FS dams were found to have been infected by Staphylococcus aureus in the udder; two within the first week (replicate 1 and 2) and another two at 4 and 6 weeks respectively (replicate 1) after parturition. Since cows infected by S. aureus were not admitted to the AM barn these dams and their calves had to be excluded. A fifth FS calf (replicate 1) was excluded after 4 weeks of age because the dam’s let-down at milking was inhibited. One HM calf (replicate 1) was taken out of the study before time of weaning because of excessive cross-sucking, and the risk that this would lead to detrimental effects on the other calves. Data from these calves were included in the study until the calves were taken out of the study, with exception of the two FS calves that were excluded within 1 week after birth. 2.2. Management and feeding All the cows gave birth to their calves in a separate calving pen (3.0 m 3.4 m) where the calves had the opportunity to suckle. The cows and calves in FS were moved to the AM barn (Fig. 1a) after 5–10 days (mainly on day 5), where FS calves continued to have free access to their dam but also had the opportunity to suckle other cows. The dams of the FS calves were permitted to enter the milking unit every 8th hour, which resulted in an average of two and a half milkings per 24 h. There was a calf creep where commercial concentrate (Talang, Lantma¨nnen, Sweden) and hay was provided ad lib. The calves could not access the cows’ feed. Water was available ad lib. in water bowls in the alleys. The cubicles and the calf creep were equipped with rubber mats and covered with chopped straw or peat (peat was used half of the time in the AM barn for replicate 2 in FS) and the walking areas were of concrete. The alleys were automatically scraped every second hour. However, from the moment a new calf and its dam had been introduced to the barn, the scrapers were started manually until the following morning. Calves in HM treatment were separated from their dams within 24 h after birth and moved to a straw-bedded single pen (1.2 m 1.0 m) in a separate calf barn, where they stayed for 3 days, and were fed colostrum (2.5 L) through a nipple bottle twice daily. Thereafter they were moved to a group pen (5.9 m 5.0 m, Fig. 1b) where only animals from HM treatment were kept. The first calves in each replicate were placed in an empty, cleaned pen. The group pen was equipped with an automatic milk feeder (CF300A, DeLaval, Tumba, Sweden) with a rubber teat available at all times. When introduced to the group pen
152
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
Fig. 1. (a) General layout of the AM barn (34 m 16 m). MU = milking unit, S = selective controlling gate (directs cows to the MU or the feeding area), W = water bowl, CF = concentrate feeder for cows. H = hay, C = concentrate, available for calves only. (b) General layout of the calf barn with single pens and two group pens (each 5.9 m 5 m) with transponder-controlled automatic milk (MF) and concentrate (CF) feeder. HAY = hay troughs.
the HM calves were given 5 kg of milk substitute daily at body temperature. Until the end of the second week the allowance in HM was gradually increased to 9.0 kg, which remained constant for the rest of the milk feeding period. The total daily allowance was available in 0.5 l portions evenly distributed over 24 h. Up to three portions could be consumed in one meal. The milk substitute (Kalvisan, AB Kvarnby Foder, Sweden) was prepared using 150 g of powder per litre, and represented the ME content of whole milk with 4% fat content. Hay, concentrate in an automatic feeder (CF300A, DeLaval, Tumba, Sweden) and water were offered ad lib. (Fig. 1b). The milk-feeder was cleaned once daily. The automatic feeders for milk and concentrate were calibrated once weekly. At about 5 weeks of age all calves were dehorned under anaesthetic and the male calves castrated at the same time. All calves were abruptly weaned at 8 weeks of age, when the milk substitute was taken out of the diet in HM and FS calves were moved to the calf barn into a group pen of the same type as HM. Calves were successively removed from the group pen after 10 weeks of age. 2.3. Behaviour of FS and HM calves week 2, 4 and 8 The behaviour of all 16 FS calves introduced to the AM barn and that of the 18 HM calves were observed 1 day in the end of weeks 2, 4 and 8, at 8:00–10:00. The observations were made by four different persons who had practised together before the start of the behavioural studies. Three calves were observed during the same 2-h observation period, in which each calf was observed as a focal animal for 1 min at a time in a rotating schedule. This resulted in each calf being observed for 40 min every observation period. The following behaviours were recorded instantaneously at 3-min intervals; - lie; lying down in different positions, - eat concentrate; taking concentrate into the mouth or chewing on concentrate, - ruminate; regurgitating bolus of rumen content and chewing on it, - sucking teat; sucking on any cow’s teat or the rubber teat in the milk-feeder, - closest neighbour; closest standing or lying calf, dam or other cow when lying down.
The frequency of the following behaviours was recorded within every third minute (if a behaviour stopped and then continued again after 5 s a new event was recorded, with the exception of vocalise, eat hay and eat straw): - move; moving at least one leg so that the body was moving in any direction (including all movements such as run/jump), - run/play; trotting, galloping or jumping, alone or with other calves, - eat hay; every occasion when taking hay into the mouth, - eat straw; every occasion when taking straw into the mouth, - sniff object; muzzle close to an object and inhaling air, - self lick; tongue touching the own body, - social; sniffing (muzzle in contact with or within 5 cm of another calf/cow), rubbing against (movements of the head against the body of another calf/cow), pushing head against head, or mounting another calf, - lick calf, dam or cow; tongue touching another calf, dam or cow, - lick/suck object; tongue or mouth touching an object, - cross-sucking; sucking on any body part of another calf, - tongue rolling; tongue moving in a circulating way. 2.4. Twenty-four hour observations of the suckling behaviour of FS calves The suckling behaviour of FS calves in the AM barn was observed approximately once weekly continuously for 24 h, starting at 21:00, which resulted in 444 h of observations. Up to four observers carried out the observations for 2–8 h each. It was possible to observe all FS calves during the observations. Whenever a calf was suckling the following recordings were made: - suckling attempt; taking a teat into the mouth or suckling for 1 min or less. A new bout was defined after an interruption of 10 min, - suckling bout; a suckling bout lasting longer than 1 min, - suckling position; suckling from inverse parallel, antiparallel, or from behind, - suckled teats; which teats that were suckled, but not the duration per teat,
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
- suckled animal; own dam, other cow with calf or cow without calf, - lick; before, during or after a suckling bout, - initiator of a suckling bout; the calf or dam moving towards the other before the start of a suckling bout, - terminator of a suckling bout; the calf, dam or cow that moved away or dam/cow that kicked during or at the end of a suckling bout. 2.5. Statistical analysis For all statistical analysis SAS version 9.1 (Statistical Analysis System Inc., Cary, USA, 2002–2003) was used. Behavioural recordings of FS and HM calves were Poisson distributed with the exception of ‘lie’ which was normally distributed. Hypotheses were tested with the generalized linear models procedure (PROC GENMOD) for effects of treatment (FS and HM), replicate (1 and 2), calf age (2, 4 and 8), sex, observer and interaction between treatment and calf age as well as treatment and replicate. As no significant effect of sex, replicate, observer or interaction between sex and replicate was found these terms were excluded from the final analysis. Model specifications included repeated measurement models with repeated measurements on individual calves supposed to follow an autoregressive (AR) correlation structure. For the recordings of ‘eat concentrate’ and ‘eat hay’ it was not possible to include the interaction between treatment and calf age due to too few recordings. Observations of the suckling behaviour of the FS calves were normally distributed. Suckling behaviour over 24 h were analysed using the mixed linear models procedure (PROC MIXED) for effects of calf age (week 2, 4 and 8), parturition number of the calf’s dam (primi- or multiparous), replicate (1 and 2), sex, and interaction between calf age and the parturition number of the calf’s dam. As no significant effect of sex or replicate was found these terms were excluded from the final analysis. Model specifications included a random calf effect and a repeated statement concerning weeks where weeks followed an unstructured covariance structure. The analysis was tested with and without the calf that was removed because of the inhibition of milk let-down of the dam and the two calves that were excluded at 4 and 6 weeks of age. No significant difference in the analysis was observed compared to when these calves were included, and data was decided to remain in the analysis. To reduce the risk of misleading interpretations of significant tests due to multiple comparisons in the analysis only P-values <0.01 were regarded as significant in multiple comparisons. Behavioural recordings of FS and HM calves are expressed as least square mean number with the confidence interval calculated from the log scale of recordings per calf and 40-min observation period. Mean values are presented as least squares means as defined in Proc Mixed. Behaviours without treatment effects or analysed by a simpler model are presented as overall arithmetic means and arithmetic means, respectively. A Chi-square test with 1 degree of freedom was used to test for differences between treatments in number of calves that were cross-sucking, number of sucking bouts
153
(on cow or milk feeder from instantaneous recordings) lasting 2 min or more and number of recordings of suckling on front and rear teats and initiator of a suckling bout (dam, calf or both dam and calf). 3. Results 3.1. Behaviour of FS and HM calves week 2, 4 and 8 There was a treatment effect on ‘lie’ (Fig. 2). The HM lay down less than FS calves weeks 2 and 8. There was no effect of calf age or interaction between treatment and calf age on ‘lie’. There was a treatment effect on ‘eat concentrate’ and ‘eat hay’ (Table 1). The HM displayed more ‘eat concentrate’ and ‘eat hay’ than the FS calves. There was a tendency of an effect of calf age on ‘eat concentrate’ (x2 = 4.9, P < 0.1) but not on ‘eat hay’. The calves in HM had more recordings of ‘eat concentrate’ and ‘eat hay’ in weeks 4 and 8 than in week 2. However, ‘eat concentrate’ and ‘eat hay’ could not be tested for interaction between treatment and calf age due to too few recordings. There was no effect of treatment or calf age on ‘eat straw’ (1.39 recordings/ 40 min). There was an effect of treatment (Table 2), calf age (x2 = 16.6, P < 0.001) and a tendency of an interaction between treatment and calf age on the recordings of
Fig. 2. Number of ‘lying’ every third minute between 8:00 and 10:00 of calves either free suckling (FS, n = 16) or fed milk substitute from an automatic feeder (HM, n = 18). Anti-logged least square means (LSM) with confidence intervals (CI) for P = 0.01. LSM with different letters are significantly different (P < 0.01).
Table 1 Arithmetic mean number of behaviours between 08:00 and 10:00 of calves either free suckling (FS, n = 16) or fed milk substitute by an automatic feeder (HM, n = 18). For effect of treatment (T) the x2-value and P are given. Behaviour
Eat concentrate Eat hayb a b
T
a
Week 2
Week 4
Week 8
T
Mean
Mean
Mean
x2
P
FS HM
0.1 0.7
0.1 0.9
0 1.9
11.5
<0.001
FS HM
0.1 1.9
0.7 2.8
0 5.3
11.9
<0.001
Number of recordings every third minute. Number of recordings within every third minute.
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
154
Table 2 Number of behaviours within every third minute between 8:00 and 10:00 of calves either free suckling (FS, n = 16) or fed milk substitute by an automatic feeder (HM, n = 18). Anti-logged least square means (LSM) with confidence intervals (CI) for P = 0.01. Different numbers within and letters between weeks are significantly different (P < 0.01). For effect of treatment (T) the x2-value and P are given. Behaviour
T
Week 2 Mean
*
1a
ab
Mean
T
x2
CI
P
2.7–6.2 2.7–8.0
b
8.4 9.0b
6.1–11.6 6.0–13.5
4.1
<0.05
11.5–24.3 20.2–31.0
14.4 22.7
10.5–19.7 18.7–27.6
10.3
<0.01
Move
FS HM
12.9 22.4
Run/play
FS HM
0.7 2.6
0.3–1.7 1.4–4.6
2.0 3.4
0.8–4.9 1.8–6.3
0.8 1.5
0.3–1.9 0.7–3.3
3.7
<0.1
Social
FS HM
3.5 6.3ab
2.1–6.0 4.6–8.6
4.8 8.1a
3.1–7.4 6.3–10.2
2.6 3.8b
1.7–3.8 2.6–5.4
5.5
<0.05
Sniff object
FS HM
4.4* 10.02
2.7–7.1 8.1–12.4
6.8 7.4
3.6–12.6 6.1–9.2
5.2
<0.05
8.0–20.8 17.7–28.4
4.5–10.5 7.6–13.7
4.1 4.7b
Week 8 CI
FS HM
6.9 10.2
1.9–5.1 0.3–1.9
Mean
Ruminate
*
3.1 0.82a
Week 4 CI
16.7 25.0
Number of recordings every third minute.
Fig. 3. Least square means (LSM SE) of the total number of suckling bouts and suckling attempts and total suckling time on the dam, cows with a calf and cows without a calf from 24 h observations once weekly (2–8) of free suckling calves in an AM barn. LSM marked with a star are significantly different (**P < 0.01, *** P < 0.001).
‘ruminate’ (x2 = 5.3, P < 0.1). The FS calves were more often recorded to ‘ruminate’. Both treatments had more recordings of ‘ruminate’ week 8 than 2, but the FS calves performed more ‘ruminate’ week 2 than the HM calves. There was no effect of treatment or calf age on ‘sucking teat’ (1.29 recordings/40 min). When comparing sucking bouts lasting 2 min or longer the FS had more long bouts than HM calves (P < 0.001). There was a tendency of a treatment effect or a treatment effect on ‘move’, ‘run/play’ ‘social’ and ‘sniff object’ (Table 2), and HM calves had more recordings of all these behaviours than FS calves. There was no effect of calf age or interaction between treatment and calf age on the behaviours ‘move’, ‘run/play’ and ‘sniff object’. There was an effect of calf age on ‘social’ (x2 = 9.5, P < 0.01), and the calves were more social in week 4 compared to week 8 (x2 = 18.6, P < 0.001). There was no effect of treatment or age on ‘lick calf, dam or cow’ (0.72 recordings/40 min). There was no effect of treatment or calf age on ‘lick or suck object’ (3.69 recordings/40 min) and ‘self lick’ (6.89 recordings/40 min). Cross-sucking was never recorded in the FS calves as compared to HM, where 11 out of 18 calves (48 recordings) displayed the behaviour (P < 0.001).
Approximately 50% of the occasions of cross-sucking were recorded within 20 min after sucking from the automatic feeder. Tongue-rolling was observed in two HM calves week 8 (3 recordings), but was never recorded in FS calves. In 63% (27 out of 43 observations) of the 40-min observations the dam of the focal FS calf was recorded as ‘‘closest neighbour’’ for at least 1 min. Of the recorded suckling bouts 19 out of 22 were on the dam, but, on one occasion each, three different calves were observed making a suckling attempt or suckling a cow with calf or a cow without calf.
Table 3 Total number of recordings of different suckling positions of free suckling calves (n = 16) when suckling from the dam, cows with a calf or cows without a calf. Suckled cow
Suckling position Reverse parallel
Anti-parallel
From behind
Dam Cow with a calf Cow without a calf
244 22 0
93 12 13
12 21 7
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
3.2. Twenty-four hour observations of suckling behaviour of FS calves The total numbers of suckling attempts were 32 and the total numbers of suckling bouts were 367, whereby 80% of these were on the dam. Counting only the suckling bouts, 84% were on the dam, 9% on cows with a calf and 7% on cows without a calf. Of the suckling bouts on cows with a calf 74% were performed by two calves and one of these calves, together with another calf, was also responsible for 66% of the suckling bouts on cows without a calf. There was a big individual difference between calves; four calves were recorded suckling on their dam only, but most of the calves were observed making an attempt to suckle or suckling another cow once or twice. Of all of the suckling bouts on cows with a calf, 52% (22 of 42) occurred simultaneously with the cow’s own calf suckling. Of the total number of suckling attempts and suckling bouts on the dam, she was most often the initiator (63%, P < 0.001) followed by the calf (21%), and both dam and calf as initiators (15%). However, an initiator was recorded in only 148 of a total of 318 suckling attempts and suckling bouts on the dam, since it was sometimes difficult to identify the initiator. One cow without a calf was initiator to three suckling bouts with one calf, and this cow was also observed licking the calf during or after all three suckling bouts. When a suckling attempt or suckling bout was on a cow with a calf, the calf not suckling its dam was never licked. The calves were licked during or afterwards in 39% of the suckling attempts or suckling bouts on their dam, and there was no difference between primi- or multiparous dams. The dam was rarely observed ending a suckling attempt or suckling bout, only 18 times, compared to a cow with a calf or a cow without a calf most often acting as the terminator when suckled by other calves. Calf age (F = 2.2, P < 0.1) and parturition number of the dam (F = 11.0, P < 0.01) tended to affect the total suckling time (suckling attempts plus suckling bouts) per 24 h on all cows (dam, cows with a calf and cows without a calf), with the total suckling time tending to decrease with increasing calf age (Fig. 3). The calves suckled longer if born to a multi- compared to primiparous cow (43.9 2.2 min vs. 32.5 2.6 min). However, when suckling on the dam, calf age or parturition number had no effect on the total suckling time per 24 h. There was an effect of calf age (F = 4.5, P < 0.001) and parturition number of the dam (F = 6.4, P < 0.05) and a tendency of an interaction between calf age and parturition number of the dam (F = 2.2, P < 0.1) on the total number of suckling bouts on all cows during 24 h. The number of suckling attempts or suckling bouts decreased with increasing calf age (Fig. 3). There was a greater number of suckling attempts or bouts in calves born to a multi- compared to a primiparous dam (5.5 0.3 vs. 4.1 0.4). Calf age (F = 4.3, P < 0.01) also had an effect on the total number of suckling bouts on the dam over 24 h, with the number of bouts decreasing with increasing calf age. During a suckling attempt or suckling bout on a cow with a calf or a cow without a calf the cow kicked at the calf or walked away in 50 and 40% of these occasions. The calf’s
155
suckling position was different depending on whether it suckled on the dam or on a cow with or without a calf (Table 3). When suckling its dam the reverse parallel position was most common. 77% of the suckling bouts in reverse parallel position on cows with a calf occurred simultaneously as the cows’ own calves suckled. During the suckling bouts the calves had 73% (429) of the teat recordings on front teats compared to 27% (155) on the rear teats (P < 0.001). 4. Discussion 4.1. Behaviour of FS and HM calves The HM calves in this study seemed to be more active due to, e.g. less lying and more movement than FS calves. The greater amount of time spent lying down in FS could be a result of the anti-stress effect of oxytocin, shown, e.g. by calmness (Uvna¨s-Moberg, 1997). The release of oxytocin has been reported to be stimulated by suckling (Lupoli et al., 2000). With increasing calf age there was no decrease in lying in either of the FS and HM treatments contrary to previous studies (Wood-Gush et al., 1984; Ylipekkala, 1990; Fro¨berg et al., 2008). This could be related to the rearing system. In the studies by Ylipekkala (1990) and Wood-Gush et al. (1984) the calves were kept on pasture. Calves kept in group pens have been found to spend as much as 16–19 h per 24 h lying down at 21–70 days of age (Jensen, 2004). In Fro¨berg et al. (2008) the calves were observed in the evening and seemed to be more active at that time than during the warmer day time. The calves in this study might have been more active at other times of the day than the observation times, and in addition, the high amount of milk consumed in both treatments could have contributed to a longer resting time. Although the FS calves were recorded to eat both concentrate and hay at the age of 2 weeks, their consumption of solid feed were mere traces, which can be related to a high milk intake. A high amount of milk is known to diminish the solid feed intake (Roy, 1980). Surprisingly, at 2 weeks of age the FS calves ruminated more than the HM calves. Since the amount of solid feed was considerably lower in FS (Fro¨berg et al., 2008, unpublished data), this could rather have been an effect of diurnal differences related to the management. The suckling time on the dam was longer than from the milk-feeder in this study. Sambraus (1984) found that milk feeding through nipples was followed by 10 min of crosssucking that was about as long as a suckling bout on the cow. Cross-sucking has been found to be frequent after milk ingestion (Dybkjær, 1988; Lidfors, 1993). de Passille´ et al. (1992) found that the motivation to suckle is elicited by the taste of milk. Deprivation of sucking may interfere with digestive processes or satiety (de Passille´, 2001). The HM calves in the present study were fed milk in many small portions. Perhaps the motivation to suckle was stimulated repeatedly but not satisfied due to the short time of milk ingestion. The HM calves were often pushed out of the feeder by other calves, although this was not recorded. In a study on a closed gate to the feeder the time in the feeder and non-nutritive sucking was prolonged
156
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
compared to a feeder without a gate, but cross-sucking without proximity to milk intake was not reduced (Weber and Wechsler, 2001). Eight litres of milk in four instead of eight portions reduced the competition at the feeder but not the number of calves that performed cross-sucking, 51% (Jensen, 2004). In contrast, De Paula Vieira et al. (2008) observed very few events of cross-sucking that they related to a slow milk flow, sufficient non-nutritive sucking, ad lib. feeding of milk, and reduced competition due to a small group size. Taken together with the findings of these studies, the HM calves in the present study may have displayed cross-sucking as an effect of the short milk intake time and the many small portions. Even tongue-rolling was exhibited in a few HM calves in this study. Also Redbo (1992) observed tongue-rolling in the second month of life in nipple or bucket-fed calves, and suggested that the short duration of milk intake could be a causal factor for the development of abnormal oral behaviours. Stereotypies often are expressed in environments that seem barren and less optimal; nevertheless, according to a review of Mason (1991) the connection to animal welfare is yet unclear. 4.2. Suckling behaviour of FS calves The calves in the present study were often licked while suckling on their dams but not on other cows. An indication of strong social bonds is a high incidence of social licking (Sato et al., 1993), which is largely restricted to the dams licking their calves (Reinhardt et al., 1986). That the dams in this study were often observed as their calf’s closest neighbour was a further indication of attachment. In spite of this, several of the calves in the present study were observed to suckle from other cows, which has been observed previously both in dairy cows (Sˇpinka and Illmann, 1992; Illmann and Sˇpinka, 1993) and beef cows as well as dairy-beef crosses (Waltl et al., 1995; Vı´chova´ and Bartosˇ, 2005). Several studies have observed that cows usually refused other calves’ suckling attempts (Beef cattle: Nicol and Sharafeldin, 1975; Price et al., 1981; Lidfors, 1994, Dairy cattle: Lidfors, 1994). In the present study there seemed to be an attachment between cow and calf, but within the herd there were cows without a calf, and these had no bonding to a calf. It is likely that it is the cows that prevent other calves suckling, more than the calves not making a suckling attempt. Other studies have reported that few calves suckled on other cows, but it was more common in dairy than beef breeds (Le Neindre, 1989) and in Zebu Holstein crossbred than Zebu cattle (Das et al., 2000). The Zebu showed also a stronger bond with their calves and more aggression towards other calves compared to the crossbreds (Das et al., 2001). Le Neindre (1989) proposed that lower attachment to the calf in dairy compared to beef breeds is probably linked to an easier milking in the former. In addition, the practise of removing the dairy calves from their dams after birth may have resulted in a relaxation of natural selection (Price, 1984) in strong maternal behaviour. In a study on an old Finnish dairy breed there was only one incidence of a calf suckling on another cow (Woivalin, 1990), which emphasises a high attachment to the calf that may relate to less selection for
milk production. Consequently, in the highly selected breed in the present study the attachment to the calf would be lower. Vı´chova´ and Bartosˇ (2005) found that calves suckling on other cows were likely compensating for some deficiency such as low birth weight or insufficient milk supply from the dam. The dairy cows in the present study were likely to have enough milk for their calves, thus it is more likely that the calves suckled on other cows because they allowed it. Although the calves often received a kick during an attempt to suckle on another cow, the cows were rarely seen to show aggression that could have implied a risk of injury. The calves suckled mostly on other cows from behind or in an anti-parallel position, similar to previous findings (Le Neindre, 1989; Illmann and Sˇpinka, 1993; Waltl et al., 1995). Waltl et al. (1995) proposed that suckling positions other than the reverse parallel did not allow the cow to identify the calf by smelling or anogential licking, and hence calves adopted these positions when suckling on other cows. Suckling bouts on other cows with a calf occurred often simultaneously with the cow’s own calf, which might have been strategic since then the cow did not show aggression. One study found that the method of rearing the dam affected the bond to her calf; mothered Friesian cows licked and nursed their calves longer than cows that had been reared in isolation (Le Neindre, 1989). Also management around calving may effect attachment. Keeping the cow and calf separate from the herd after birth, as is the case in the present study, might facilitate the attachment between dam and calf. Lidfors (1994) observed more closeness between dam and calf and more licking of the calf, when the calf had been born in isolation. In Sˇpinka and Illmann (1992) and Illmann and Sˇpinka (1993) the calves and their dams were kept in a pen together with other animals, which thus could have interfered with attachment. In this study most of the suckling bouts were initiated by the dam. Lidfors et al. (1994) found that dams of beef cattle initiated many of the suckling bouts during the first week of life, whereas the calves were more often the initiator at older ages. In young dairy calves kept in a loosehousing system with parlour milking the dam initiated most of the suckling bouts, however, only six calves were ¨ rtendahl, 1996). In semi-wild Maremma cattle observed (O the cows were rarely seen to move towards their calves (Vitale et al., 1986). The calves in the present study suckled high-yielding cows; hence their motivation to initiate a suckling bout was probably lower compared to calves of breeds with a lower production. In this study the suckling time on the dam was similar until weaning. The total suckling time on all cows decreased with increasing calf age, which was probably related to the high frequency of suckling attempts and bouts on other cows early in life. The number of suckling bouts decreased from 6.3 at 2 weeks of age to 3.8 at 8 weeks of age. Nicol and Sharafeldin (1975) studied beef calves until the age of 120 days and found that the number of suckling bouts per day decreased from 5.6 at 7 days of age to 3.5 at 24 days and then remained at 3.0–3.5. The total suckling time has been reported to decrease with age
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
(Zebu cattle: Reinhardt and Reinhardt, 1981; Maremma cattle: Vitale et al., 1986) or to remain relatively constant with age (Beef cattle: Nicol and Sharafeldin, 1975; KileyWorthington and de la Plain, 1983; Lidfors and Jensen, 1988). However, there may be discrepancies between studies due to different duration of the observation periods, and many have not covered the entire diurnal circle of 24 h. Calves of multi- compared to primiparous cows suckled for a longer time and more frequently in this study. A possible explanation could be that milk removal during milking seemed to be more efficient in multiparous dams, and consequently they were likely to have less milk available for their calves compared to primiparous cows. The calves showed a preference for front teats compared to rear teats which is in line with previous studies on dairy calves (Jung, 1994; Fro¨berg et al., 2008). In high-yielding cows, the front teats are easier to reach than the rear teats and they have enough milk to satisfy the calves (Jung, 1994), which probably was the case in the present study. Jung (1994) proposed that the suckling time on the rear teats is increased when the age of the calf or time since last suckling increases or when the dam’s milk yield seemed to be low. 5. Conclusion This study showed that calves with free access to suckle on their dam in an AM barn rested more, ate less solid feed and exhibited less non-nutritive oral behaviours, than calves that were fed milk substitute from an automatic feeder in group pen. Further, there are indications that free suckling calves and their dams develop a strong attachment to each other, as the free suckling calves suckled mainly on their dam, and in reverse parallel position, and the dams often licked their calves and were in close proximity to them. Acknowledgements This study was financed by the Swedish Animal Welfare ¨ sterberg, Agency. The authors wish to thank Peter O Ingemar Olsson and Kerstin Svennersten-Sjaunja for helping with planning the study and all other help. We also thank the staff at Kungsa¨ngen research centre for help and assistance, Mikaela Patel, Helena Hultborn and Malin Langenfors for help with behavioural recordings, Dietrich von Rosen for statistical advice and Luella Godman and Margaret Knipe for their linguistic revision. References Das, S.M., Redbo, I., Wiktorsson, H., 2000. Effect of age of calf on suckling behaviour and other behavioural activities of Zebu and crossbred calves during restricted suckling periods. Appl. Anim. Behav. Sci. 67, 47–57. Das, S.M., Redbo, I., Wiktorsson, H., 2001. Behaviour of zebu and zebu crossbred cows in restricted suckling groups. Appl. Anim. Behav. Sci. 72, 263–270. de Passille´, A.M., 2001. Sucking motivation and related problems in calves. Appl. Anim. Behav. Sci. 72, 175–187. de Passille´, A.M.B., Metz, J.H.M., Mekking, P., Wiepkema, P.R., 1992. Does drinking milk stimulate sucking in young calves? Appl. Anim. Behav. Sci. 34, 23–26.
157
De Paula Vieira, A., Guesdon, V., de Passille´, A.M., von Keyserlingk, M.A.G., Weary, D.M., 2008. Behavioural indicators of hunger in dairy calves. Appl. Anim. Behav. Sci. 109, 180–189. Dybkjær, L., 1988. Sma˚kalvars adfærd. Dansk Veterinærtidsskrift 71, 113– 121. Fro¨berg, S., Gratte, E., Svennersten-Sjaunja, K., Olsson, I., Berg, C., Orihuela, A., Galina, C.S., Garcı´a, B., Lidfors, L., 2008. Effect of suckling (‘restricted suckling’) on dairy cows’ udder health and milk let-down and their calves’ weight gain, feed intake and behaviour. Appl. Anim. Behav. Sci. 113, 1–14. Grøndahl, A.M., Skancke, E.M., Mejdell, C.M., Jansen, J.H., 2007. Growth rate, health and welfare in a dairy herd with natural suckling until 6–8 weeks of age: a case report. Acta Vet. Scand. 49, http://www. actavetscand.com/content/49/1/16 (accessed 5 March 2008). Gubernick, D.J., 1981. Parent and infant attachment in mammals. In: Gubernick, D.J., Klopfer, P.H. (Eds.), Parental Care in Mammals. Plenum Press, New York, pp. 243–305. Illmann, G., Sˇpinka, M., 1993. Maternal behaviour of dairy heifers and suckling of their newborn calves in group housing. Appl. Anim. Behav. Sci. 36, 91–98. Jensen, M.B., 2004. Computer-controlled milk feeding of dairy calves: the effects of calves per feeder and number of milk portions on use of feeder and social behaviour. J. Dairy Sci. 87, 3428–3438. Jung, J., 1994. Temporal patterning of natural suckling behaviour of dairy calves. Swedish University of Agricultural Sciences, Faculty of Veterinary Medicine, Department of Animal Hygiene, Specialarbete 21, 46 pp. Kiley-Worthington, M., de la Plain, S., 1983. The behaviour of beef suckler cattle (Bos taurus). Tierhaltung 14. Birkha¨user Verlag, Basel, 195 pp. Krohn, C.C., Foldager, J., Mogensen, L., 1999. Long-term effect of colostrum feeding methods on behaviour in female dairy calves. Acta Agric. Scand., Sect. A. Anim. Sci. 49, 57–64. Le Neindre, P., 1989. Influence of cattle-rearing conditions and breed on social relationships of mother and young. Appl. Anim. Behav. Sci. 23, 117–127. Le Neindre, P., Sourd, C., 1984. Influence of rearing conditions on subsequent social behaviour of Friesian and Salers heifers from birth to six months of age. Appl. Anim. Behav. Sci. 12, 43–52. Lidfors, L.M., 1993. Cross-sucking in group-housed dairy calves before and after weaning off milk. Appl. Anim. Behav. Sci. 38, 15–24. Lidfors, L., 1994. Mother-young behaviour in cattle. Parturition, development of cow-calf attachment, suckling and effects of separation. Ph.D. Diss. Swedish University of Agricultural Science, Skara, Sweden, Report 33, 71 pp. Lidfors, L., Jensen, P., 1988. Behaviour of free-ranging beef cows and calves. Appl. Anim. Behav. Sci. 20, 237–247. Lidfors, L.M., Jensen, P., Algers, B., 1994. Suckling in free-ranging beef cattle—temporal patterning of suckling bouts and effects of age and sex. Ethology 98, 321–332. Lupoli, B., Johansson, B., Uvna¨s-Moberg, K., Svennersten-Sjaunja, K., 2000. Effect of suckling on the release of oxytocin, prolactin, cortisol, gastrin, cholecystokinin, somatostatin and insulin in dairy cows and their calves. J. Dairy Res. 68, 175–187. Mason, G.J., 1991. Stereotypies: a critical review. Anim. Behav. 41, 1015– 1037. Nicol, A.M., Sharafeldin, M.A., 1975. Observations on the behaviour of single-suckled calves from birth to 120 days. Proc. N. Z. Soc. Anim. Prod. 35, 221–230. ¨ rtendahl, M., 1996. Milk production combined with suckling calves—a O case study on Ekena¨s estate of behaviour, calf growth, milk production and udder health. Avdelningen fo¨r byggnadsvetenskap, Institutionen fo¨r lantbruksteknik, Institutionen fo¨r husdjurens utfodring och va˚rd, Sveriges Lantbruksuniversitet, Rapport 212, 45 pp. (In Swedish, abstract in English). Price, E.O., 1984. Behavioral aspects of animal domestication. Quart. Rev. Biol. 59, 1–32. Price, E.O., Thos, J., Anderson, G.B., 1981. Maternal responses of confined beef cattle to single versus twin calves. J. Anim. Sci. 53, 934–939. Redbo, I., 1992. Stereotypies in dairy cattle—and their relation to confinement, production-related factors, physiological reactions, and adjoining behaviours. Ph.D. Diss. Swedish University of Agricultural Science, Skara, Report 212, 49 pp. Reinhardt, V., Reinhardt, A., 1981. Natural sucking performance and age of weaning in zebu cattle (Bos indicus). J. Agric. Sci. Camb. 96, 309– 312. Reinhardt, C., Reinhardt, A., Reinhardt, V., 1986. Social behaviour and reproductive performance of semi-wild Scottish highland cattle. Appl. Anim. Behav. Sci. 15, 125–136. Roy, J.H.B., 1980. The Calf. Butterworths, London, 442 pp.
158
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
Sambraus, H.H., 1984. Gegenseitiges besaugen von ka¨lbern bei ku¨nstlicher aufzucht. Berl. Mu¨nch. Tiera¨rztl. Wschr. 97, 119–123 (Abstract in English). Sambraus, H.H., 1985. Mouth-based anomalous syndromes. In: Fraser, A.F. (Ed.), Ethology of Farm Animals. Elsevier, Amsterdam, pp. 391– 422. Sato, S., Tarumizu, K., Hatae, K., 1993. The influence of social factors on allogroming in cows. Appl. Anim. Behav. Sci. 38, 235–244. Sato, S., Nagamine, R., Kubo, T., 1994. Tongue-playing in tethered Japanese Black cattle: diurnal patterns, analysis of variance and behaviour sequences. Appl. Anim. Behav. Sci. 39, 39–47. Sˇpinka, M., Illmann, G., 1992. Suckling behaviour of young dairy calves with their own and alien mothers. Appl. Anim. Behav. Sci. 33, 165– 173. Stephens, D.B., 1982. A review of some behavioural and physiological studies which are relevant to the welfare of young calves. In: Signoret, J.P. (Ed.), Welfare and Husbandry of Calves. Martinus Nijhoff, The Hague, pp. 47–69. Swedish Dairy Association, 2008. Genomsnittligt antal kor per besa¨ttning. http://svenskmjolk.se/ImageVault/Images/id_472/scope_128/ImageVaultHandler.aspx (accessed 12 June 2008) (In Swedish). Uvna¨s-Moberg, K., 1997. Physiological and endocrine effects of social contact; role of oxytocin. Ann Ny Acad Sci 807, 146–163.
Vı´chova´, J., Bartosˇ, L., 2005. Allosuckling in cattle: gain or compensation? Appl. Anim. Behav. Sci. 94, 223–235. Vitale, A.F., Tenucci, M., Papini, M., Lovari, S., 1986. Social behaviour of the calves of semi-wild Maremma cattle, Bos primigenius taurus. Appl. Anim. Behav. Sci. 16, 217–231. Waltl, B., Appleby, M.C., So¨lkner, J., 1995. Effects of relatedness on the suckling behaviour of calves in a herd of beef cattle rearing twins. Appl. Anim. Behav. Sci. 45, 1–9. Weber, R., Wechsler, B., 2001. Reduction in cross-sucking in calves by the use of a modified automatic teat feeder. Appl. Anim. Behav. Sci. 72, 215–223. Wiepkema, P.R., van Hellemond, K.K., Roessingh, P., Romberg, H., 1987. Behavioural and abomasal damage in individual veal calves. Appl. Anim. Behav. Sci. 18, 257–268. Woivalin, A., 1990. Beteende hos friga˚ende mjo¨lkkokalvar—socialt beteende. Veterina¨rmedicinska ho¨gskolan. Institutionen fo¨r husdjurshygien, Helsingfors (Abstract in English). Wood-Gush, D.G.M., Hunt, K., Carson, K., Dennison, S.G.C., 1984. The early behaviour of suckler calves in the field. Biol. Behav. 9, 295–306. Ylipekkala, A., 1990. The Behaviour of Free-Ranging Dairy Calves— Development of Behaviour. Department of Animal Hygiene, University of Veterinary Medicine, Helsinki, Finland, 31 pp. (graduation work).
Contents lists available at ScienceDirect
Applied Animal Behaviour Science journal homepage: www.elsevier.com/locate/applanim
Behaviour of dairy calves suckling the dam in a barn with automatic milking or being fed milk substitute from an automatic feeder in a group pen Sofie Fro¨berg a,*, Lena Lidfors b a b
Department of Animal Nutrition and Management, Swedish University of Agricultural Sciences (SLU), Kungsa¨ngen Research Centre, SE-753 23 Uppsala, Sweden Department of Animal Environment and Health, SLU, P.O. Box 234, SE-532 23 Skara, Sweden
A R T I C L E I N F O
A B S T R A C T
Article history: Accepted 16 December 2008
The aim of this study was to investigate the suckling behaviour of dairy calves allowed to suckle freely in a barn with an automatic milking (AM) system and how their behaviour differed from the behaviour of calves given milk substitute from an automatic feeder. For the first 8 weeks of life, calves of the dairy breed Swedish Red were either suckling freely (FS, n = 16) from the dam in an AM barn or kept in groups with automatic feeding (HM, n = 18) with an allowance of 9 kg of milk substitute per day. All treatments had concentrate and hay ad lib. Focal calves in FS and HM treatments were observed one day in weeks 2, 4 and 8, at 08:00–10:00 and the FS calves’ suckling behaviour was observed for a 24-h period once weekly. The HM calves moved, ate concentrate and ate hay more than the FS calves, whereas the FS calves lay down and ruminated more than the HM calves. Crosssucking and tongue-rolling were displayed in HM but not in FS calves. Of the suckling bouts recorded in FS calves 80% was on the dam. Four calves were recorded suckling solely on their dam, but most of the calves were observed attempting to suckle or suckling on other cows in the group once or twice. The total suckling time and number of suckling bouts during week 2–8 on all cows decreased with increasing calf age. In conclusion, the free-suckling calves ate less solid feed, rested more and exhibited fewer non-nutritive oral and abnormal behaviours compared to the calves fed milk substitute from an automatic feeder, and some calves often suckled on other cows. ß 2009 Elsevier B.V. All rights reserved.
Keywords: Dairy calves Behaviour Free suckling Automatic milk feeding Automatic milking system
1. Introduction Herd sizes in industrialized countries are increasing, e.g. in 1990 the Swedish mean herd size was 22 cows, while in 2007 it was 51 (Swedish Dairy Association, 2008). As an effect of increased herd sizes and technology the number of calves kept in groups with automatic milk feeding is likely to increase. Group pens facilitate movement, play and social contact among calves. However, behavioural problems such as calves sucking on body parts
* Corresponding author. Tel.: +46 18 67 16 79; fax: +46 18 67 29 46. E-mail address: sofi[email protected] (S. Fro¨berg). 0168-1591/$ – see front matter ß 2009 Elsevier B.V. All rights reserved. doi:10.1016/j.applanim.2008.12.015
of other calves (cross-sucking) can be common in calves kept in groups (Stephens, 1982; Lidfors, 1993; Fro¨berg et al., 2008). Cross-sucking may facilitate disease transmission (de Passille´, 2001). Cross-sucking and other nonnutritive oral behaviours such as licking objects are less common in calves allowed to suckle the dam (Krohn et al., 1999; Fro¨berg et al., 2008) or being with the dam without access to suckle (Krohn et al., 1999). Wiepkema et al. (1987) proposed that depriving a calf of sucking a teat might cause stress or conflict that could develop into stereotypies. Other studies (Sambraus, 1985; Sato et al., 1994) also suggest that stereotypies may be related to insufficiently stimulated or suppressed feeding behaviours. Moreover, the presence of the cow may be
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
important for social learning (Krohn et al., 1999). Le Neindre and Sourd (1984) found a higher social activity in heifers that had been reared by their dams the first 10–12 weeks compared to those reared artificially. The function of an attachment between parent and young is to ensure that food, warmth and protection is provided for one’s own young and not to others (Gubernick, 1981). Parental attachment is found in species living in larger groups in which the young are precocial, but, in herds where the females are related a combination of attachment and cooperative care of their young may have evolved (Gubernick, 1981). The suckling behaviour of calves that have continuous access to their dams has been observed in several studies, e.g. on semi-wild Maremma cattle (Bos primigenius taurus, Vitale et al., 1986), pure beef or beef and dairy-beef crosses (Nicol and Sharafeldin, 1975; Lidfors et al., 1994; Waltl et al., 1995; Vı´chova´ and Bartosˇ, 2005) and Zebu cattle and their crosses (Reinhardt and Reinhardt, 1981). There are only few studies on free suckling dairy calves and most of them focus on the first days after parturition with a few exceptions, e.g. Ylipekkala (1990). Behavioural variations between breeds may be expected due to different selection and breeding goals among the breeds. In general beef cattle production systems encourage and benefit from strong maternal behaviour, whereas there may be a selection against it in dairy production, as the calf is most often separated from its dam shortly after birth. It is also likely that a low attachment of the dam to its calf is linked to easy milking in high milk-yielding breeds (Le Neindre, 1989). The calves’ suckling behaviour may also be affected by several factors, i.e. the cow’s milk production, if the cow is milked during the milk feeding period, and whether the calf has free or restricted access to suckle. In a study on free suckling in loose housing with parlour milking it was concluded that the system functioned satisfactorily, but no systematic behavioural studies were conducted (Grøndahl et al., 2007). As far as we know there are no previous scientific publications on raising calves with their dams in loosehousing with an automatic milking (AM) system. The aim of this study was to investigate the suckling behaviour of dairy calves allowed to suckle freely in an AM barn and how their behaviour differed from the behaviour of calves given milk substitute from an automatic feeder. The first hypothesis was that due to the supposed relaxation of the mother–young bond in dairy cattle the calves would suckle on other cows than their dams and when they suckle on other cows their position would not be reverse parallel position. The second hypothesis was that free-suckling calves would be more active but exhibit less non-nutritive behaviour such as cross-sucking than calves fed milk from an automatic feeder. 2. Materials and methods 2.1. Animals and experimental design The present study was carried out at the Swedish University of Agricultural Sciences, Kungsa¨ngen Research Centre, Uppsala. Calves of the Swedish Red dairy breed single-born from August to March were studied from birth
151
to weaning at 8 weeks of age. The cows giving birth to the calves ranged from the first to the fifth lactation. In order of birth calves were allocated to each of the two following treatments; milk substitute from an automatic feeder (HM) and free suckling (FS). When up to 12 calves had been allocated to the first treatment, calves were assigned to the next treatment. Later calves were allocated to a second replicate resulting in a total of 23 (replicate 1; n = 11, replicate 2; n = 12) calves in HM and 18 (replicate 1; n = 8, replicate 2; n = 10) calves in FS treatment. In the FS treatment there were seven primiparous and nine multiparous cows. Four FS dams were found to have been infected by Staphylococcus aureus in the udder; two within the first week (replicate 1 and 2) and another two at 4 and 6 weeks respectively (replicate 1) after parturition. Since cows infected by S. aureus were not admitted to the AM barn these dams and their calves had to be excluded. A fifth FS calf (replicate 1) was excluded after 4 weeks of age because the dam’s let-down at milking was inhibited. One HM calf (replicate 1) was taken out of the study before time of weaning because of excessive cross-sucking, and the risk that this would lead to detrimental effects on the other calves. Data from these calves were included in the study until the calves were taken out of the study, with exception of the two FS calves that were excluded within 1 week after birth. 2.2. Management and feeding All the cows gave birth to their calves in a separate calving pen (3.0 m 3.4 m) where the calves had the opportunity to suckle. The cows and calves in FS were moved to the AM barn (Fig. 1a) after 5–10 days (mainly on day 5), where FS calves continued to have free access to their dam but also had the opportunity to suckle other cows. The dams of the FS calves were permitted to enter the milking unit every 8th hour, which resulted in an average of two and a half milkings per 24 h. There was a calf creep where commercial concentrate (Talang, Lantma¨nnen, Sweden) and hay was provided ad lib. The calves could not access the cows’ feed. Water was available ad lib. in water bowls in the alleys. The cubicles and the calf creep were equipped with rubber mats and covered with chopped straw or peat (peat was used half of the time in the AM barn for replicate 2 in FS) and the walking areas were of concrete. The alleys were automatically scraped every second hour. However, from the moment a new calf and its dam had been introduced to the barn, the scrapers were started manually until the following morning. Calves in HM treatment were separated from their dams within 24 h after birth and moved to a straw-bedded single pen (1.2 m 1.0 m) in a separate calf barn, where they stayed for 3 days, and were fed colostrum (2.5 L) through a nipple bottle twice daily. Thereafter they were moved to a group pen (5.9 m 5.0 m, Fig. 1b) where only animals from HM treatment were kept. The first calves in each replicate were placed in an empty, cleaned pen. The group pen was equipped with an automatic milk feeder (CF300A, DeLaval, Tumba, Sweden) with a rubber teat available at all times. When introduced to the group pen
152
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
Fig. 1. (a) General layout of the AM barn (34 m 16 m). MU = milking unit, S = selective controlling gate (directs cows to the MU or the feeding area), W = water bowl, CF = concentrate feeder for cows. H = hay, C = concentrate, available for calves only. (b) General layout of the calf barn with single pens and two group pens (each 5.9 m 5 m) with transponder-controlled automatic milk (MF) and concentrate (CF) feeder. HAY = hay troughs.
the HM calves were given 5 kg of milk substitute daily at body temperature. Until the end of the second week the allowance in HM was gradually increased to 9.0 kg, which remained constant for the rest of the milk feeding period. The total daily allowance was available in 0.5 l portions evenly distributed over 24 h. Up to three portions could be consumed in one meal. The milk substitute (Kalvisan, AB Kvarnby Foder, Sweden) was prepared using 150 g of powder per litre, and represented the ME content of whole milk with 4% fat content. Hay, concentrate in an automatic feeder (CF300A, DeLaval, Tumba, Sweden) and water were offered ad lib. (Fig. 1b). The milk-feeder was cleaned once daily. The automatic feeders for milk and concentrate were calibrated once weekly. At about 5 weeks of age all calves were dehorned under anaesthetic and the male calves castrated at the same time. All calves were abruptly weaned at 8 weeks of age, when the milk substitute was taken out of the diet in HM and FS calves were moved to the calf barn into a group pen of the same type as HM. Calves were successively removed from the group pen after 10 weeks of age. 2.3. Behaviour of FS and HM calves week 2, 4 and 8 The behaviour of all 16 FS calves introduced to the AM barn and that of the 18 HM calves were observed 1 day in the end of weeks 2, 4 and 8, at 8:00–10:00. The observations were made by four different persons who had practised together before the start of the behavioural studies. Three calves were observed during the same 2-h observation period, in which each calf was observed as a focal animal for 1 min at a time in a rotating schedule. This resulted in each calf being observed for 40 min every observation period. The following behaviours were recorded instantaneously at 3-min intervals; - lie; lying down in different positions, - eat concentrate; taking concentrate into the mouth or chewing on concentrate, - ruminate; regurgitating bolus of rumen content and chewing on it, - sucking teat; sucking on any cow’s teat or the rubber teat in the milk-feeder, - closest neighbour; closest standing or lying calf, dam or other cow when lying down.
The frequency of the following behaviours was recorded within every third minute (if a behaviour stopped and then continued again after 5 s a new event was recorded, with the exception of vocalise, eat hay and eat straw): - move; moving at least one leg so that the body was moving in any direction (including all movements such as run/jump), - run/play; trotting, galloping or jumping, alone or with other calves, - eat hay; every occasion when taking hay into the mouth, - eat straw; every occasion when taking straw into the mouth, - sniff object; muzzle close to an object and inhaling air, - self lick; tongue touching the own body, - social; sniffing (muzzle in contact with or within 5 cm of another calf/cow), rubbing against (movements of the head against the body of another calf/cow), pushing head against head, or mounting another calf, - lick calf, dam or cow; tongue touching another calf, dam or cow, - lick/suck object; tongue or mouth touching an object, - cross-sucking; sucking on any body part of another calf, - tongue rolling; tongue moving in a circulating way. 2.4. Twenty-four hour observations of the suckling behaviour of FS calves The suckling behaviour of FS calves in the AM barn was observed approximately once weekly continuously for 24 h, starting at 21:00, which resulted in 444 h of observations. Up to four observers carried out the observations for 2–8 h each. It was possible to observe all FS calves during the observations. Whenever a calf was suckling the following recordings were made: - suckling attempt; taking a teat into the mouth or suckling for 1 min or less. A new bout was defined after an interruption of 10 min, - suckling bout; a suckling bout lasting longer than 1 min, - suckling position; suckling from inverse parallel, antiparallel, or from behind, - suckled teats; which teats that were suckled, but not the duration per teat,
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
- suckled animal; own dam, other cow with calf or cow without calf, - lick; before, during or after a suckling bout, - initiator of a suckling bout; the calf or dam moving towards the other before the start of a suckling bout, - terminator of a suckling bout; the calf, dam or cow that moved away or dam/cow that kicked during or at the end of a suckling bout. 2.5. Statistical analysis For all statistical analysis SAS version 9.1 (Statistical Analysis System Inc., Cary, USA, 2002–2003) was used. Behavioural recordings of FS and HM calves were Poisson distributed with the exception of ‘lie’ which was normally distributed. Hypotheses were tested with the generalized linear models procedure (PROC GENMOD) for effects of treatment (FS and HM), replicate (1 and 2), calf age (2, 4 and 8), sex, observer and interaction between treatment and calf age as well as treatment and replicate. As no significant effect of sex, replicate, observer or interaction between sex and replicate was found these terms were excluded from the final analysis. Model specifications included repeated measurement models with repeated measurements on individual calves supposed to follow an autoregressive (AR) correlation structure. For the recordings of ‘eat concentrate’ and ‘eat hay’ it was not possible to include the interaction between treatment and calf age due to too few recordings. Observations of the suckling behaviour of the FS calves were normally distributed. Suckling behaviour over 24 h were analysed using the mixed linear models procedure (PROC MIXED) for effects of calf age (week 2, 4 and 8), parturition number of the calf’s dam (primi- or multiparous), replicate (1 and 2), sex, and interaction between calf age and the parturition number of the calf’s dam. As no significant effect of sex or replicate was found these terms were excluded from the final analysis. Model specifications included a random calf effect and a repeated statement concerning weeks where weeks followed an unstructured covariance structure. The analysis was tested with and without the calf that was removed because of the inhibition of milk let-down of the dam and the two calves that were excluded at 4 and 6 weeks of age. No significant difference in the analysis was observed compared to when these calves were included, and data was decided to remain in the analysis. To reduce the risk of misleading interpretations of significant tests due to multiple comparisons in the analysis only P-values <0.01 were regarded as significant in multiple comparisons. Behavioural recordings of FS and HM calves are expressed as least square mean number with the confidence interval calculated from the log scale of recordings per calf and 40-min observation period. Mean values are presented as least squares means as defined in Proc Mixed. Behaviours without treatment effects or analysed by a simpler model are presented as overall arithmetic means and arithmetic means, respectively. A Chi-square test with 1 degree of freedom was used to test for differences between treatments in number of calves that were cross-sucking, number of sucking bouts
153
(on cow or milk feeder from instantaneous recordings) lasting 2 min or more and number of recordings of suckling on front and rear teats and initiator of a suckling bout (dam, calf or both dam and calf). 3. Results 3.1. Behaviour of FS and HM calves week 2, 4 and 8 There was a treatment effect on ‘lie’ (Fig. 2). The HM lay down less than FS calves weeks 2 and 8. There was no effect of calf age or interaction between treatment and calf age on ‘lie’. There was a treatment effect on ‘eat concentrate’ and ‘eat hay’ (Table 1). The HM displayed more ‘eat concentrate’ and ‘eat hay’ than the FS calves. There was a tendency of an effect of calf age on ‘eat concentrate’ (x2 = 4.9, P < 0.1) but not on ‘eat hay’. The calves in HM had more recordings of ‘eat concentrate’ and ‘eat hay’ in weeks 4 and 8 than in week 2. However, ‘eat concentrate’ and ‘eat hay’ could not be tested for interaction between treatment and calf age due to too few recordings. There was no effect of treatment or calf age on ‘eat straw’ (1.39 recordings/ 40 min). There was an effect of treatment (Table 2), calf age (x2 = 16.6, P < 0.001) and a tendency of an interaction between treatment and calf age on the recordings of
Fig. 2. Number of ‘lying’ every third minute between 8:00 and 10:00 of calves either free suckling (FS, n = 16) or fed milk substitute from an automatic feeder (HM, n = 18). Anti-logged least square means (LSM) with confidence intervals (CI) for P = 0.01. LSM with different letters are significantly different (P < 0.01).
Table 1 Arithmetic mean number of behaviours between 08:00 and 10:00 of calves either free suckling (FS, n = 16) or fed milk substitute by an automatic feeder (HM, n = 18). For effect of treatment (T) the x2-value and P are given. Behaviour
Eat concentrate Eat hayb a b
T
a
Week 2
Week 4
Week 8
T
Mean
Mean
Mean
x2
P
FS HM
0.1 0.7
0.1 0.9
0 1.9
11.5
<0.001
FS HM
0.1 1.9
0.7 2.8
0 5.3
11.9
<0.001
Number of recordings every third minute. Number of recordings within every third minute.
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
154
Table 2 Number of behaviours within every third minute between 8:00 and 10:00 of calves either free suckling (FS, n = 16) or fed milk substitute by an automatic feeder (HM, n = 18). Anti-logged least square means (LSM) with confidence intervals (CI) for P = 0.01. Different numbers within and letters between weeks are significantly different (P < 0.01). For effect of treatment (T) the x2-value and P are given. Behaviour
T
Week 2 Mean
*
1a
ab
Mean
T
x2
CI
P
2.7–6.2 2.7–8.0
b
8.4 9.0b
6.1–11.6 6.0–13.5
4.1
<0.05
11.5–24.3 20.2–31.0
14.4 22.7
10.5–19.7 18.7–27.6
10.3
<0.01
Move
FS HM
12.9 22.4
Run/play
FS HM
0.7 2.6
0.3–1.7 1.4–4.6
2.0 3.4
0.8–4.9 1.8–6.3
0.8 1.5
0.3–1.9 0.7–3.3
3.7
<0.1
Social
FS HM
3.5 6.3ab
2.1–6.0 4.6–8.6
4.8 8.1a
3.1–7.4 6.3–10.2
2.6 3.8b
1.7–3.8 2.6–5.4
5.5
<0.05
Sniff object
FS HM
4.4* 10.02
2.7–7.1 8.1–12.4
6.8 7.4
3.6–12.6 6.1–9.2
5.2
<0.05
8.0–20.8 17.7–28.4
4.5–10.5 7.6–13.7
4.1 4.7b
Week 8 CI
FS HM
6.9 10.2
1.9–5.1 0.3–1.9
Mean
Ruminate
*
3.1 0.82a
Week 4 CI
16.7 25.0
Number of recordings every third minute.
Fig. 3. Least square means (LSM SE) of the total number of suckling bouts and suckling attempts and total suckling time on the dam, cows with a calf and cows without a calf from 24 h observations once weekly (2–8) of free suckling calves in an AM barn. LSM marked with a star are significantly different (**P < 0.01, *** P < 0.001).
‘ruminate’ (x2 = 5.3, P < 0.1). The FS calves were more often recorded to ‘ruminate’. Both treatments had more recordings of ‘ruminate’ week 8 than 2, but the FS calves performed more ‘ruminate’ week 2 than the HM calves. There was no effect of treatment or calf age on ‘sucking teat’ (1.29 recordings/40 min). When comparing sucking bouts lasting 2 min or longer the FS had more long bouts than HM calves (P < 0.001). There was a tendency of a treatment effect or a treatment effect on ‘move’, ‘run/play’ ‘social’ and ‘sniff object’ (Table 2), and HM calves had more recordings of all these behaviours than FS calves. There was no effect of calf age or interaction between treatment and calf age on the behaviours ‘move’, ‘run/play’ and ‘sniff object’. There was an effect of calf age on ‘social’ (x2 = 9.5, P < 0.01), and the calves were more social in week 4 compared to week 8 (x2 = 18.6, P < 0.001). There was no effect of treatment or age on ‘lick calf, dam or cow’ (0.72 recordings/40 min). There was no effect of treatment or calf age on ‘lick or suck object’ (3.69 recordings/40 min) and ‘self lick’ (6.89 recordings/40 min). Cross-sucking was never recorded in the FS calves as compared to HM, where 11 out of 18 calves (48 recordings) displayed the behaviour (P < 0.001).
Approximately 50% of the occasions of cross-sucking were recorded within 20 min after sucking from the automatic feeder. Tongue-rolling was observed in two HM calves week 8 (3 recordings), but was never recorded in FS calves. In 63% (27 out of 43 observations) of the 40-min observations the dam of the focal FS calf was recorded as ‘‘closest neighbour’’ for at least 1 min. Of the recorded suckling bouts 19 out of 22 were on the dam, but, on one occasion each, three different calves were observed making a suckling attempt or suckling a cow with calf or a cow without calf.
Table 3 Total number of recordings of different suckling positions of free suckling calves (n = 16) when suckling from the dam, cows with a calf or cows without a calf. Suckled cow
Suckling position Reverse parallel
Anti-parallel
From behind
Dam Cow with a calf Cow without a calf
244 22 0
93 12 13
12 21 7
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
3.2. Twenty-four hour observations of suckling behaviour of FS calves The total numbers of suckling attempts were 32 and the total numbers of suckling bouts were 367, whereby 80% of these were on the dam. Counting only the suckling bouts, 84% were on the dam, 9% on cows with a calf and 7% on cows without a calf. Of the suckling bouts on cows with a calf 74% were performed by two calves and one of these calves, together with another calf, was also responsible for 66% of the suckling bouts on cows without a calf. There was a big individual difference between calves; four calves were recorded suckling on their dam only, but most of the calves were observed making an attempt to suckle or suckling another cow once or twice. Of all of the suckling bouts on cows with a calf, 52% (22 of 42) occurred simultaneously with the cow’s own calf suckling. Of the total number of suckling attempts and suckling bouts on the dam, she was most often the initiator (63%, P < 0.001) followed by the calf (21%), and both dam and calf as initiators (15%). However, an initiator was recorded in only 148 of a total of 318 suckling attempts and suckling bouts on the dam, since it was sometimes difficult to identify the initiator. One cow without a calf was initiator to three suckling bouts with one calf, and this cow was also observed licking the calf during or after all three suckling bouts. When a suckling attempt or suckling bout was on a cow with a calf, the calf not suckling its dam was never licked. The calves were licked during or afterwards in 39% of the suckling attempts or suckling bouts on their dam, and there was no difference between primi- or multiparous dams. The dam was rarely observed ending a suckling attempt or suckling bout, only 18 times, compared to a cow with a calf or a cow without a calf most often acting as the terminator when suckled by other calves. Calf age (F = 2.2, P < 0.1) and parturition number of the dam (F = 11.0, P < 0.01) tended to affect the total suckling time (suckling attempts plus suckling bouts) per 24 h on all cows (dam, cows with a calf and cows without a calf), with the total suckling time tending to decrease with increasing calf age (Fig. 3). The calves suckled longer if born to a multi- compared to primiparous cow (43.9 2.2 min vs. 32.5 2.6 min). However, when suckling on the dam, calf age or parturition number had no effect on the total suckling time per 24 h. There was an effect of calf age (F = 4.5, P < 0.001) and parturition number of the dam (F = 6.4, P < 0.05) and a tendency of an interaction between calf age and parturition number of the dam (F = 2.2, P < 0.1) on the total number of suckling bouts on all cows during 24 h. The number of suckling attempts or suckling bouts decreased with increasing calf age (Fig. 3). There was a greater number of suckling attempts or bouts in calves born to a multi- compared to a primiparous dam (5.5 0.3 vs. 4.1 0.4). Calf age (F = 4.3, P < 0.01) also had an effect on the total number of suckling bouts on the dam over 24 h, with the number of bouts decreasing with increasing calf age. During a suckling attempt or suckling bout on a cow with a calf or a cow without a calf the cow kicked at the calf or walked away in 50 and 40% of these occasions. The calf’s
155
suckling position was different depending on whether it suckled on the dam or on a cow with or without a calf (Table 3). When suckling its dam the reverse parallel position was most common. 77% of the suckling bouts in reverse parallel position on cows with a calf occurred simultaneously as the cows’ own calves suckled. During the suckling bouts the calves had 73% (429) of the teat recordings on front teats compared to 27% (155) on the rear teats (P < 0.001). 4. Discussion 4.1. Behaviour of FS and HM calves The HM calves in this study seemed to be more active due to, e.g. less lying and more movement than FS calves. The greater amount of time spent lying down in FS could be a result of the anti-stress effect of oxytocin, shown, e.g. by calmness (Uvna¨s-Moberg, 1997). The release of oxytocin has been reported to be stimulated by suckling (Lupoli et al., 2000). With increasing calf age there was no decrease in lying in either of the FS and HM treatments contrary to previous studies (Wood-Gush et al., 1984; Ylipekkala, 1990; Fro¨berg et al., 2008). This could be related to the rearing system. In the studies by Ylipekkala (1990) and Wood-Gush et al. (1984) the calves were kept on pasture. Calves kept in group pens have been found to spend as much as 16–19 h per 24 h lying down at 21–70 days of age (Jensen, 2004). In Fro¨berg et al. (2008) the calves were observed in the evening and seemed to be more active at that time than during the warmer day time. The calves in this study might have been more active at other times of the day than the observation times, and in addition, the high amount of milk consumed in both treatments could have contributed to a longer resting time. Although the FS calves were recorded to eat both concentrate and hay at the age of 2 weeks, their consumption of solid feed were mere traces, which can be related to a high milk intake. A high amount of milk is known to diminish the solid feed intake (Roy, 1980). Surprisingly, at 2 weeks of age the FS calves ruminated more than the HM calves. Since the amount of solid feed was considerably lower in FS (Fro¨berg et al., 2008, unpublished data), this could rather have been an effect of diurnal differences related to the management. The suckling time on the dam was longer than from the milk-feeder in this study. Sambraus (1984) found that milk feeding through nipples was followed by 10 min of crosssucking that was about as long as a suckling bout on the cow. Cross-sucking has been found to be frequent after milk ingestion (Dybkjær, 1988; Lidfors, 1993). de Passille´ et al. (1992) found that the motivation to suckle is elicited by the taste of milk. Deprivation of sucking may interfere with digestive processes or satiety (de Passille´, 2001). The HM calves in the present study were fed milk in many small portions. Perhaps the motivation to suckle was stimulated repeatedly but not satisfied due to the short time of milk ingestion. The HM calves were often pushed out of the feeder by other calves, although this was not recorded. In a study on a closed gate to the feeder the time in the feeder and non-nutritive sucking was prolonged
156
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
compared to a feeder without a gate, but cross-sucking without proximity to milk intake was not reduced (Weber and Wechsler, 2001). Eight litres of milk in four instead of eight portions reduced the competition at the feeder but not the number of calves that performed cross-sucking, 51% (Jensen, 2004). In contrast, De Paula Vieira et al. (2008) observed very few events of cross-sucking that they related to a slow milk flow, sufficient non-nutritive sucking, ad lib. feeding of milk, and reduced competition due to a small group size. Taken together with the findings of these studies, the HM calves in the present study may have displayed cross-sucking as an effect of the short milk intake time and the many small portions. Even tongue-rolling was exhibited in a few HM calves in this study. Also Redbo (1992) observed tongue-rolling in the second month of life in nipple or bucket-fed calves, and suggested that the short duration of milk intake could be a causal factor for the development of abnormal oral behaviours. Stereotypies often are expressed in environments that seem barren and less optimal; nevertheless, according to a review of Mason (1991) the connection to animal welfare is yet unclear. 4.2. Suckling behaviour of FS calves The calves in the present study were often licked while suckling on their dams but not on other cows. An indication of strong social bonds is a high incidence of social licking (Sato et al., 1993), which is largely restricted to the dams licking their calves (Reinhardt et al., 1986). That the dams in this study were often observed as their calf’s closest neighbour was a further indication of attachment. In spite of this, several of the calves in the present study were observed to suckle from other cows, which has been observed previously both in dairy cows (Sˇpinka and Illmann, 1992; Illmann and Sˇpinka, 1993) and beef cows as well as dairy-beef crosses (Waltl et al., 1995; Vı´chova´ and Bartosˇ, 2005). Several studies have observed that cows usually refused other calves’ suckling attempts (Beef cattle: Nicol and Sharafeldin, 1975; Price et al., 1981; Lidfors, 1994, Dairy cattle: Lidfors, 1994). In the present study there seemed to be an attachment between cow and calf, but within the herd there were cows without a calf, and these had no bonding to a calf. It is likely that it is the cows that prevent other calves suckling, more than the calves not making a suckling attempt. Other studies have reported that few calves suckled on other cows, but it was more common in dairy than beef breeds (Le Neindre, 1989) and in Zebu Holstein crossbred than Zebu cattle (Das et al., 2000). The Zebu showed also a stronger bond with their calves and more aggression towards other calves compared to the crossbreds (Das et al., 2001). Le Neindre (1989) proposed that lower attachment to the calf in dairy compared to beef breeds is probably linked to an easier milking in the former. In addition, the practise of removing the dairy calves from their dams after birth may have resulted in a relaxation of natural selection (Price, 1984) in strong maternal behaviour. In a study on an old Finnish dairy breed there was only one incidence of a calf suckling on another cow (Woivalin, 1990), which emphasises a high attachment to the calf that may relate to less selection for
milk production. Consequently, in the highly selected breed in the present study the attachment to the calf would be lower. Vı´chova´ and Bartosˇ (2005) found that calves suckling on other cows were likely compensating for some deficiency such as low birth weight or insufficient milk supply from the dam. The dairy cows in the present study were likely to have enough milk for their calves, thus it is more likely that the calves suckled on other cows because they allowed it. Although the calves often received a kick during an attempt to suckle on another cow, the cows were rarely seen to show aggression that could have implied a risk of injury. The calves suckled mostly on other cows from behind or in an anti-parallel position, similar to previous findings (Le Neindre, 1989; Illmann and Sˇpinka, 1993; Waltl et al., 1995). Waltl et al. (1995) proposed that suckling positions other than the reverse parallel did not allow the cow to identify the calf by smelling or anogential licking, and hence calves adopted these positions when suckling on other cows. Suckling bouts on other cows with a calf occurred often simultaneously with the cow’s own calf, which might have been strategic since then the cow did not show aggression. One study found that the method of rearing the dam affected the bond to her calf; mothered Friesian cows licked and nursed their calves longer than cows that had been reared in isolation (Le Neindre, 1989). Also management around calving may effect attachment. Keeping the cow and calf separate from the herd after birth, as is the case in the present study, might facilitate the attachment between dam and calf. Lidfors (1994) observed more closeness between dam and calf and more licking of the calf, when the calf had been born in isolation. In Sˇpinka and Illmann (1992) and Illmann and Sˇpinka (1993) the calves and their dams were kept in a pen together with other animals, which thus could have interfered with attachment. In this study most of the suckling bouts were initiated by the dam. Lidfors et al. (1994) found that dams of beef cattle initiated many of the suckling bouts during the first week of life, whereas the calves were more often the initiator at older ages. In young dairy calves kept in a loosehousing system with parlour milking the dam initiated most of the suckling bouts, however, only six calves were ¨ rtendahl, 1996). In semi-wild Maremma cattle observed (O the cows were rarely seen to move towards their calves (Vitale et al., 1986). The calves in the present study suckled high-yielding cows; hence their motivation to initiate a suckling bout was probably lower compared to calves of breeds with a lower production. In this study the suckling time on the dam was similar until weaning. The total suckling time on all cows decreased with increasing calf age, which was probably related to the high frequency of suckling attempts and bouts on other cows early in life. The number of suckling bouts decreased from 6.3 at 2 weeks of age to 3.8 at 8 weeks of age. Nicol and Sharafeldin (1975) studied beef calves until the age of 120 days and found that the number of suckling bouts per day decreased from 5.6 at 7 days of age to 3.5 at 24 days and then remained at 3.0–3.5. The total suckling time has been reported to decrease with age
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
(Zebu cattle: Reinhardt and Reinhardt, 1981; Maremma cattle: Vitale et al., 1986) or to remain relatively constant with age (Beef cattle: Nicol and Sharafeldin, 1975; KileyWorthington and de la Plain, 1983; Lidfors and Jensen, 1988). However, there may be discrepancies between studies due to different duration of the observation periods, and many have not covered the entire diurnal circle of 24 h. Calves of multi- compared to primiparous cows suckled for a longer time and more frequently in this study. A possible explanation could be that milk removal during milking seemed to be more efficient in multiparous dams, and consequently they were likely to have less milk available for their calves compared to primiparous cows. The calves showed a preference for front teats compared to rear teats which is in line with previous studies on dairy calves (Jung, 1994; Fro¨berg et al., 2008). In high-yielding cows, the front teats are easier to reach than the rear teats and they have enough milk to satisfy the calves (Jung, 1994), which probably was the case in the present study. Jung (1994) proposed that the suckling time on the rear teats is increased when the age of the calf or time since last suckling increases or when the dam’s milk yield seemed to be low. 5. Conclusion This study showed that calves with free access to suckle on their dam in an AM barn rested more, ate less solid feed and exhibited less non-nutritive oral behaviours, than calves that were fed milk substitute from an automatic feeder in group pen. Further, there are indications that free suckling calves and their dams develop a strong attachment to each other, as the free suckling calves suckled mainly on their dam, and in reverse parallel position, and the dams often licked their calves and were in close proximity to them. Acknowledgements This study was financed by the Swedish Animal Welfare ¨ sterberg, Agency. The authors wish to thank Peter O Ingemar Olsson and Kerstin Svennersten-Sjaunja for helping with planning the study and all other help. We also thank the staff at Kungsa¨ngen research centre for help and assistance, Mikaela Patel, Helena Hultborn and Malin Langenfors for help with behavioural recordings, Dietrich von Rosen for statistical advice and Luella Godman and Margaret Knipe for their linguistic revision. References Das, S.M., Redbo, I., Wiktorsson, H., 2000. Effect of age of calf on suckling behaviour and other behavioural activities of Zebu and crossbred calves during restricted suckling periods. Appl. Anim. Behav. Sci. 67, 47–57. Das, S.M., Redbo, I., Wiktorsson, H., 2001. Behaviour of zebu and zebu crossbred cows in restricted suckling groups. Appl. Anim. Behav. Sci. 72, 263–270. de Passille´, A.M., 2001. Sucking motivation and related problems in calves. Appl. Anim. Behav. Sci. 72, 175–187. de Passille´, A.M.B., Metz, J.H.M., Mekking, P., Wiepkema, P.R., 1992. Does drinking milk stimulate sucking in young calves? Appl. Anim. Behav. Sci. 34, 23–26.
157
De Paula Vieira, A., Guesdon, V., de Passille´, A.M., von Keyserlingk, M.A.G., Weary, D.M., 2008. Behavioural indicators of hunger in dairy calves. Appl. Anim. Behav. Sci. 109, 180–189. Dybkjær, L., 1988. Sma˚kalvars adfærd. Dansk Veterinærtidsskrift 71, 113– 121. Fro¨berg, S., Gratte, E., Svennersten-Sjaunja, K., Olsson, I., Berg, C., Orihuela, A., Galina, C.S., Garcı´a, B., Lidfors, L., 2008. Effect of suckling (‘restricted suckling’) on dairy cows’ udder health and milk let-down and their calves’ weight gain, feed intake and behaviour. Appl. Anim. Behav. Sci. 113, 1–14. Grøndahl, A.M., Skancke, E.M., Mejdell, C.M., Jansen, J.H., 2007. Growth rate, health and welfare in a dairy herd with natural suckling until 6–8 weeks of age: a case report. Acta Vet. Scand. 49, http://www. actavetscand.com/content/49/1/16 (accessed 5 March 2008). Gubernick, D.J., 1981. Parent and infant attachment in mammals. In: Gubernick, D.J., Klopfer, P.H. (Eds.), Parental Care in Mammals. Plenum Press, New York, pp. 243–305. Illmann, G., Sˇpinka, M., 1993. Maternal behaviour of dairy heifers and suckling of their newborn calves in group housing. Appl. Anim. Behav. Sci. 36, 91–98. Jensen, M.B., 2004. Computer-controlled milk feeding of dairy calves: the effects of calves per feeder and number of milk portions on use of feeder and social behaviour. J. Dairy Sci. 87, 3428–3438. Jung, J., 1994. Temporal patterning of natural suckling behaviour of dairy calves. Swedish University of Agricultural Sciences, Faculty of Veterinary Medicine, Department of Animal Hygiene, Specialarbete 21, 46 pp. Kiley-Worthington, M., de la Plain, S., 1983. The behaviour of beef suckler cattle (Bos taurus). Tierhaltung 14. Birkha¨user Verlag, Basel, 195 pp. Krohn, C.C., Foldager, J., Mogensen, L., 1999. Long-term effect of colostrum feeding methods on behaviour in female dairy calves. Acta Agric. Scand., Sect. A. Anim. Sci. 49, 57–64. Le Neindre, P., 1989. Influence of cattle-rearing conditions and breed on social relationships of mother and young. Appl. Anim. Behav. Sci. 23, 117–127. Le Neindre, P., Sourd, C., 1984. Influence of rearing conditions on subsequent social behaviour of Friesian and Salers heifers from birth to six months of age. Appl. Anim. Behav. Sci. 12, 43–52. Lidfors, L.M., 1993. Cross-sucking in group-housed dairy calves before and after weaning off milk. Appl. Anim. Behav. Sci. 38, 15–24. Lidfors, L., 1994. Mother-young behaviour in cattle. Parturition, development of cow-calf attachment, suckling and effects of separation. Ph.D. Diss. Swedish University of Agricultural Science, Skara, Sweden, Report 33, 71 pp. Lidfors, L., Jensen, P., 1988. Behaviour of free-ranging beef cows and calves. Appl. Anim. Behav. Sci. 20, 237–247. Lidfors, L.M., Jensen, P., Algers, B., 1994. Suckling in free-ranging beef cattle—temporal patterning of suckling bouts and effects of age and sex. Ethology 98, 321–332. Lupoli, B., Johansson, B., Uvna¨s-Moberg, K., Svennersten-Sjaunja, K., 2000. Effect of suckling on the release of oxytocin, prolactin, cortisol, gastrin, cholecystokinin, somatostatin and insulin in dairy cows and their calves. J. Dairy Res. 68, 175–187. Mason, G.J., 1991. Stereotypies: a critical review. Anim. Behav. 41, 1015– 1037. Nicol, A.M., Sharafeldin, M.A., 1975. Observations on the behaviour of single-suckled calves from birth to 120 days. Proc. N. Z. Soc. Anim. Prod. 35, 221–230. ¨ rtendahl, M., 1996. Milk production combined with suckling calves—a O case study on Ekena¨s estate of behaviour, calf growth, milk production and udder health. Avdelningen fo¨r byggnadsvetenskap, Institutionen fo¨r lantbruksteknik, Institutionen fo¨r husdjurens utfodring och va˚rd, Sveriges Lantbruksuniversitet, Rapport 212, 45 pp. (In Swedish, abstract in English). Price, E.O., 1984. Behavioral aspects of animal domestication. Quart. Rev. Biol. 59, 1–32. Price, E.O., Thos, J., Anderson, G.B., 1981. Maternal responses of confined beef cattle to single versus twin calves. J. Anim. Sci. 53, 934–939. Redbo, I., 1992. Stereotypies in dairy cattle—and their relation to confinement, production-related factors, physiological reactions, and adjoining behaviours. Ph.D. Diss. Swedish University of Agricultural Science, Skara, Report 212, 49 pp. Reinhardt, V., Reinhardt, A., 1981. Natural sucking performance and age of weaning in zebu cattle (Bos indicus). J. Agric. Sci. Camb. 96, 309– 312. Reinhardt, C., Reinhardt, A., Reinhardt, V., 1986. Social behaviour and reproductive performance of semi-wild Scottish highland cattle. Appl. Anim. Behav. Sci. 15, 125–136. Roy, J.H.B., 1980. The Calf. Butterworths, London, 442 pp.
158
S. Fro¨berg, L. Lidfors / Applied Animal Behaviour Science 117 (2009) 150–158
Sambraus, H.H., 1984. Gegenseitiges besaugen von ka¨lbern bei ku¨nstlicher aufzucht. Berl. Mu¨nch. Tiera¨rztl. Wschr. 97, 119–123 (Abstract in English). Sambraus, H.H., 1985. Mouth-based anomalous syndromes. In: Fraser, A.F. (Ed.), Ethology of Farm Animals. Elsevier, Amsterdam, pp. 391– 422. Sato, S., Tarumizu, K., Hatae, K., 1993. The influence of social factors on allogroming in cows. Appl. Anim. Behav. Sci. 38, 235–244. Sato, S., Nagamine, R., Kubo, T., 1994. Tongue-playing in tethered Japanese Black cattle: diurnal patterns, analysis of variance and behaviour sequences. Appl. Anim. Behav. Sci. 39, 39–47. Sˇpinka, M., Illmann, G., 1992. Suckling behaviour of young dairy calves with their own and alien mothers. Appl. Anim. Behav. Sci. 33, 165– 173. Stephens, D.B., 1982. A review of some behavioural and physiological studies which are relevant to the welfare of young calves. In: Signoret, J.P. (Ed.), Welfare and Husbandry of Calves. Martinus Nijhoff, The Hague, pp. 47–69. Swedish Dairy Association, 2008. Genomsnittligt antal kor per besa¨ttning. http://svenskmjolk.se/ImageVault/Images/id_472/scope_128/ImageVaultHandler.aspx (accessed 12 June 2008) (In Swedish). Uvna¨s-Moberg, K., 1997. Physiological and endocrine effects of social contact; role of oxytocin. Ann Ny Acad Sci 807, 146–163.
Vı´chova´, J., Bartosˇ, L., 2005. Allosuckling in cattle: gain or compensation? Appl. Anim. Behav. Sci. 94, 223–235. Vitale, A.F., Tenucci, M., Papini, M., Lovari, S., 1986. Social behaviour of the calves of semi-wild Maremma cattle, Bos primigenius taurus. Appl. Anim. Behav. Sci. 16, 217–231. Waltl, B., Appleby, M.C., So¨lkner, J., 1995. Effects of relatedness on the suckling behaviour of calves in a herd of beef cattle rearing twins. Appl. Anim. Behav. Sci. 45, 1–9. Weber, R., Wechsler, B., 2001. Reduction in cross-sucking in calves by the use of a modified automatic teat feeder. Appl. Anim. Behav. Sci. 72, 215–223. Wiepkema, P.R., van Hellemond, K.K., Roessingh, P., Romberg, H., 1987. Behavioural and abomasal damage in individual veal calves. Appl. Anim. Behav. Sci. 18, 257–268. Woivalin, A., 1990. Beteende hos friga˚ende mjo¨lkkokalvar—socialt beteende. Veterina¨rmedicinska ho¨gskolan. Institutionen fo¨r husdjurshygien, Helsingfors (Abstract in English). Wood-Gush, D.G.M., Hunt, K., Carson, K., Dennison, S.G.C., 1984. The early behaviour of suckler calves in the field. Biol. Behav. 9, 295–306. Ylipekkala, A., 1990. The Behaviour of Free-Ranging Dairy Calves— Development of Behaviour. Department of Animal Hygiene, University of Veterinary Medicine, Helsinki, Finland, 31 pp. (graduation work).