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The effect of lying motivation on cow behaviour
Applied Animal Behaviour Science 176 (2016) 1–5
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Applied Animal Behaviour Science journal homepage: www.elsevier.com/locate/applanim
The effect of lying motivation on cow behaviour Marianna Norring ∗ , Anna Valros Research Centre for Animal Welfare, Faculty of Veterinary Medicine, University of Helsinki, FI-00014 Helsinki, Finland
a r t i c l e
i n f o
Article history: Received 26 August 2015 Received in revised form 26 November 2015 Accepted 29 November 2015 Available online 10 December 2015 Keywords: Behaviour Dairy cow Lying Motivation Milk yield
a b s t r a c t Cows in dairy barns spend time standing while waiting for milking, accessing feed and entering the resting area. It has been suggested that high yielding cows may represent a trade off situation regarding eating and lying particularly in systems where there is a long waiting period before milking. We tested the effect on lying motivation of preventing cows of different production levels from lying down. The study included 14 primiparous cows and 14 multiparous cows in tiestalls at their 8th week of lactation. Cows were milked at approximately 06:00 and 18:00 h and milk yield was recorded. The behaviour of the cows was observed over 2 days when the cows were prevented from lying, either from 10:00 to 14:00 or from 14:00 to 18:00 h and during 2 preceding baseline days. Lying, eating, ruminating and lying with neck muscles relaxed (used as a behavioural indicator for sleep) were observed for 4 h before and 4 h after milking. The percentage time allocated to different behaviours per period was analysed using mixed models. In addition, stereotypic leaning behaviour was observed during the deprivation period. We established that the cows lay more after deprivation compared with during the baseline period for that time of day (21% vs. 34% before milking and 49% vs. 55% after milking). The cows spent less time eating after forced standing (23% vs. 21%). Cows used more time lying inactively without ruminating and lying with neck relaxed post treatment (11% vs. 19%). In addition, milk yield and lying time were correlated during the period after forced standing, indicating greater motivation in high yielding cows to rest. In total, 19 of 28 cows housed in tie stalls exhibited leaning behaviour. The duration of leaning was associated with the duration of standing. In conclusion, a 4-h standing period appeared sufficient to increase the motivation for lying and sleeping. Milk yield correlated with shorter lying duration when the cows were motivated to rest. The motivational background of the leaning behaviour in cows requires further investigation. © 2015 Elsevier B.V. All rights reserved.
1. Introduction Many studies have shown the importance of lying behaviour in cows (Munksgaard and Simonsen, 1996; Fisher et al., 2002; Cooper et al., 2008) and under time constraints dairy cows prioritize their lying time even over eating time (Munksgaard et al., 2005). Total daily lying time is higher in low yielding cows than in high yielding cows (Norring et al., 2012). A link between milk yield or anticipation of milking and lying was supported by the observation that time spent lying decreased towards the subsequent milking (Overton et al., 2002) and that cows milked frequently rested more at dawn than cows milked less frequently (Österman and Redbo, 2000). In addition, cows increased their mooing and stood more by the gate when one of the daily milkings was skipped (Pomiès et al., 2007) and a skipped milking event increased milk leakage
∗ Corresponding author. E-mail address: Marianna.Norring@Helsinki.fi (M. Norring). http://dx.doi.org/10.1016/j.applanim.2015.11.022 0168-1591/© 2015 Elsevier B.V. All rights reserved.
and udder firmness, and resulted in decreased total lying duration and shorter lying bouts (O’Driscoll et al., 2011). Changes in lying and standing durations were suggested to be indicative of udder discomfort during mastitis (Medrano-Galarza et al., 2012). Cows subjected to abrupt cessation of milking exhibited behaviours indicative of anticipated milking more than cows that were gradually weaned from milking (Zobel et al., 2013). Moreover, dry-off resulted in shorter duration of lying, indicating udder discomfort and higher motivation to be milked to be linked with reduced lying times (Zobel et al., 2013). The motivational background of increased standing duration of high yielding cows is still unknown. According to Norring et al. (2012) cows were lying down 22% of their time before evening milking and 50% of their time after milking. However, it is not clear why the lying duration prior to milking was lower than after milking. It might be caused by anticipation of milking, or by relaxation after milking. It is also possible that the resting motivation changes during the course of the afternoon due to some other reason. A way to study the underlying motivation, and the link to milk yield, more
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M. Norring, A. Valros / Applied Animal Behaviour Science 176 (2016) 1–5
specifically, while controlling for e.g. diurnal variation, is to manipulate the lying motivation by depriving cows of the opportunity to lie down during a specific period. Lying deprivation induces both physiological and behavioural stress responses in dairy cows (Ruckebusch, 1974; Munksgaard and Simonsen, 1996; Fisher et al., 2002). Even short bouts of 2–4 h standing has an effect (Metz, 1985; Cooper et al., 2007, 2008; Krebs et al., 2011). Pressing or leaning the muzzle on stall constructions is described as leaning behaviour and this behaviour is suggested to be increased during lying deprivation and during tethering (Krohn, 1994; Munksgaard and Simonsen 1996). However, there are not enough studies about leaning behaviour to be able to fully understand how the environment and characteristics of individuals contribute to this behaviour. In order to shed further light on the reasons for the decreased lying observed prior to milking, we investigated the consequences of 4 h of forced standing on behaviour before and after evening milking. We also investigated whether milk yield was connected to the responses to deprivation of lying. In addition, the effect of lying deprivation on leaning behaviour was explored. 2. Material and methods 2.1. Animals and housing The time budget of 28 cows was observed over 4 days. We used Finnish Ayrshire cows in the 8th week of lactation with a mean of 58.3 days (SD 5.1) from parturition on the first day of observation. Fourteen of the cows were primiparous and the remainder had calved 2.5 (SD 1.1) times. The cows weighed a mean of 616 kg (SD 66). The gaits of the cows were visually scored according to Sprecher et al. (1997), and only cows with a normal gait score (1 or 2) and cows that were not being treated for any disease were included in the experiment. The University of Helsinki ethical committee approved the experiment. Water was offered ad libitum from a water bowl. Cows were milked at 06:00 and 18:00 h in their tie-stalls. Before and during the experiment, all cows were housed in the same insulated barn in tie-stalls (170 × 121 cm) with rubber mats (2 cm thick, Delaval, Sweden) bedded with sawdust. The cows were tethered to a neck bar. The cows were fed grass silage ad libitum . Silage was added approximately at 05:00, 12:00, 14:00, 15:00 and 16:00 h according to the working schedule of the caretakers. The leftover silage was removed daily. In addition, all cows were fed a mixture of 2 commercial concentrates (Suomen rehu, Espoo, Finland). Cows were fed concentrate according to milk yield following Finnish feeding recommendations, the daily amount ranging from 12.5 to 20.5 kg per cow (Tuori et al., 2000). The daily total amount of concentrate intake of each cow was divided among 6 equal meals and offered to the animals at approximately at 05:00, 08:00, 11:00, 14:00, 17:00, and 20:00 h. 2.2. Procedures and observations When a cow reached her 8th week of lactation she was placed in one of the experimental stalls. The cows were allowed to adapt to their new location in the barn for 2 days. Subsequently the behaviour of the animals was recorded at 14:00 to 22:00 h during 4 days. If a cow was inseminated filming was postponed for one day and the day of insemination was excluded. Data on baseline behaviour was collected on days 1 and 2. On days 3 and 4 cows were prevented from lying for a 4-h period each day. Lying was prevented using a girth that went around the cow’s trunk and was attached to the ceiling above her tie stall. Lying behaviour was prevented
Table 1 Postures and behaviours observed and the definitions of observations. Lying inactive without ruminating and lying with the neck relaxed were observed to estimate resting and sleeping time. Posture Lying Standing
Rump on the floor Standing on four feet
Behaviour Eating Ruminating Inactive without ruminating Lying with the neck relaxed
Other activities
Taking silage into the mouth and jaw moving, or muzzle in contact with silage and moving Jaw moving, not taking silage into the mouth Jaw not moving, head not moving, head up Jaw not moving, neck relaxed and head in contact with the feeding Table in front of the cow, or reversed along her body All other behaviours of the head not mentioned above, for example head moving actively, drinking, eating concentrate, licking, social behaviour, pressing the muzzle towards equipment
from 10:00 until 14:00 and from 14:00 until the evening milking at approximately 18:00 h. Half of the cows (14) were prevented from lying from 10:00 and the other half beginning from 14:00 h on day 3 and vice versa on day 4. The effect of lying deprivation on the behaviour of the cows was observed both before and after milking, in order to establish the effect of increased lying motivation on behaviour during this specific time period. The milk yield was measured according to milk recording standards at each milking; the accuracy achieved was ±200 g (Tru-test Limited, Pakuranga, New Zealand). Three regular, adjacent tie-stalls, in the middle of the row of tie-stalls, were equipped as experimental stalls. A total of 8 video cameras (Sanyo VCB-3372P, Watford, UK) were used to observe the cows’ behaviour. Time-lapse (Panasonic AG-6730, Secaucus, NJ) recording with 24 h time mode was used, and the pictures from all cameras were fed to a single tape using a multiplexer (Robot, Tyco, Princeton, NJ). 2.3. Behaviours Recordings were made of the lying or standing posture, head and jaw movements indicative of eating, rumination and sleeping behaviours (Table 1). The behaviours and postures were recorded through continuous observation (Observer Noldus, Wageningen, the Netherlands) and were analysed with an accuracy of 1 min while observing the videos. The bout of a single behaviour was considered to end when another behaviour began. Leaning behaviour was observed from 14:00 until 18:00 on 2 baseline days and on one day while the cows were prevented from lying from 14:00 to 18:00. The video analyses of leaning behaviour were made during separate viewings using continuous observation. We were not able to record if the leaning included the weight shift towards the head, thus all pressing or leaning or contact of the muzzle or nostril on stall constructions, including the feeding bowl or front curb, were recorded as leaning behaviour. 2.4. Statistical methods Based on visual inspection of histograms the data was considered to be normally distributed. Percentage time used in different behaviours during the 4-h period before and after evening milking was analysed using the mixed procedure of SPSS. The behavioural responses in time periods 14:00–18:00 and 18:00–22:00 h were analysed separately. Treatment (no forced standing, forced standing 10:00–14:00 or forced standing 14:00–18:00 h), order of treatment (forced standing at 10:00–14:00 followed by forced
M. Norring, A. Valros / Applied Animal Behaviour Science 176 (2016) 1–5 Table 2 Average percentage of time used in different behaviours during 4 h before evening milking (14:00 to 18:00 h) showing the difference between the baseline phase behaviour and post-standing (n = 28). % of Behaviour at 14:00 to 18:00 h
70
a
23 56 20 46
21 45 18 43
14:00-18:00
60
SE P-value
Baseline
50
Baseline Post standing 10:00 to 14:00 Eating Standing, not eatinga Ruminating Other active behaviours
3
1 2 1 2
0.031 0.001 ns 0.052
Post standing 10-14
*
40 % 30
*
Percent of standing time minus percent of eating time.
20 standing 14:00–18:00 h or vice versa), and parity (primiparous or multiparous) were fixed factors. Cow was included as a random factor. Separate mixed models were used to analyse the effect of milk yield on behaviour of lying-motivated cows. The behaviours before (14:00–18:00) and after (18:00–22:00 h) milking during control and treatment times were analysed separately. Cow was included as a random factor. Milk yield was used as a continuous measure and was included in the model as a regression variable. We used yield at the evening milking and average evening milk yield for the baseline behaviour measurement days. Parity was inserted as a fixed factor into the model. Latency from evening milking to lying down was analysed using a linear mixed model. Cow was considered a random factor and treatment (none, after standing 10:00–14:00 or standing 14:00–8:00 h) was inserted into the model as a fixed factor. Parity was inserted as a fixed factor. Milk yield was used as regression variable. The effect of treatment on the total duration of leaning and proportion of duration of leaning during standing was analysed using linear mixed models. Total duration and proportion of leaning from standing time were used in the analysis as an outcome measure. Cow was considered a random factor and day (average of 2 baseline days vs. treatment day) was inserted into the model as a fixed factor. In addition, the baseline behaviour, milk yield and parity of cows that performed leaning were compared with those for cows that did not engage in such behaviour. The characteristics of leaning cows were compared with those of the non-leaning cows using linear mixed models. The lying, eating, ruminating, lying with neck relaxed, inactive without ruminating, other active behaviours, milk yield and parity were analysed. All cows observed leaning at least once were classified as leaning. The model included day as a repeated effect and leaning behaviour as a fixed effect. 3. Results
10 0 70
Lying
18:00-22:00
Lying inacve
*
60
Baseline
50
Post standing 10-14 Post standing 14-18
40 % 30
20 10 0 Lying
Lying inacve
Fig. 1. Percent of time spent on resting behaviours before and after evening milking by different treatments and baseline. Lying inactive consists of behaviour classes lying inactive without ruminating and lying with neck relaxed (*P < 0.05).
The mean evening milk yield of the cows across the study period was 18 ± 0.3 kg. The milk yield decreased with duration of lying (P = 0.028, slope −1.97%/kg) after evening milking on the day when the cows were forced to stand from 10:00 to 14:00; but there was no effect of milk yield on duration of lying when cows were forced to stand from 14:00 to 18:00 h. There was no effect of milk yield on duration of lying before evening milking (P > 0.05). During the baseline period there was no effect of milk yield on duration of lying either before or after evening milking. Milk yield had no effect on other behaviours analysed.
3.1. Behaviour 4-h post treatment 3.2. Lying down latency after milking When the cows were forced to stand from 10:00 to 14:00 h they spent more time lying down and more time lying inactive during the following 4-h period compared with baseline behaviour, (Fig. 1). Animals spent less time eating and less time “standing and not eating” immediately after forced standing (Table 2). Treatment order had an effect on the 2 behaviours during the time period 14:00 to 18:00 h; cows that were assigned to forced standing from 10:00 to 14:00 first spent more time eating (24% vs. 20%, P = 0.042) and were less occupied by “other active behaviours” (40% vs. 49%, P = 0.003) compared with cows tested later. During the time period 18:00–22:00 h treatment order had an effect on eating behaviour; cows first assigned to forced standing from 10:00 to 14:00 h spent longer eating than cows tested in the other order (20% vs. 14%, P = 0.024).
When the cows were forced to stand for the last 4 h before milking, the latency period to lie down after milking tended to be shorter than baseline (16.6 a vs. 15.7 vs. 6.7 b ± 4 min: baseline, forced standing 10:00–14:00, forced standing 14:00–18:00 h, respectively; F2,55 = 2.8, P = 0.07). The milk yield or parity had no effect on latency to lie down after evening milking. 3.3. Leaning behaviour The frequency of leaning increased during the periods with forced standing and duration of leaning tended to be longer during the periods with forced standing compared with baseline periods (Table 4). However, there were no differences in proportion of time
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M. Norring, A. Valros / Applied Animal Behaviour Science 176 (2016) 1–5
Table 3 Average percentage of time used in different behaviours during 4 h after evening milking (18:00 to 22:00 h) showing the difference between the baseline phase behaviour and post standing (n = 28). % of Behaviour at 18:00 to 22:00 h
Eating Standing, not eatinga Ruminating Other active behaviours a
Baseline
Post standing 10:00 to 14:00
Post standing 14:00 to 18:00
17 33b 24 32
18b 28c 24 31
15c 30 27 31
SE
P-value
1 3 2 2
0.025 0.040 ns ns
Percent of standing time minus percent of eating time. Different letters within rows indicate significant difference in post hoc tests.
b,c
Table 4 Mean and SE for leaning during a 4 h period from 14:00 to 18:00 for 28 cows.
Leaning duration min Frequency of leaning bouts Proportion of leaning while standing %
Baseline
Treatment
SE
P-value
7 3.5 3.1
8 5.8 3.5
2 1.1 0.9
0.07 0.008 ns
used for leaning when only the time spent standing was included. Leaning always took place while standing; we did not observe leaning while lying. Leaning behaviour was observed in 19 out of the 28 cows. The cows that leaned also spent more time eating silage during the baseline days than non-leaning cows (227 vs. 194 min/d; P = 0.048). There were no other differences between cows that performed leaning and those that did not. 4. Discussion This study supported earlier results indicating that during normal conditions, cows spend much less time lying down prior to milking than after milking. When cows were deprived of lying during a period between eight and four hours prior to milking, they increased the duration of lying down during the last four hours before milking, indicating an increased motivation for resting. Milk yield did not correlate to lying behaviour prior to milking neither in the baseline situation, nor after the deprivation treatment. We were thus not able to confirm that the amount of milk in the udder before evening milking had an effect on the duration of lying down during this period. In addition, we found that forced standing increased the frequency of leaning. According to our earlier findings (Norring et al., 2012) based on 24 h observation period with these animals, the cows with higher daily milk yield during the baseline phase had shorter lying times. Österman and Redbo (2001) suggested that a full udder prior to milking could hinder lying. O’Driscoll et al. (2011) observed udder tension, milk leakage and a shorter lying after a skipped evening milking. In addition, at the time of an omitted milking, cows stood near the milking parlour gate probably in anticipation of milking (Pomiès et al., 2007; Zobel et al., 2013). In previous studies shorter lying times were recorded around morning milking or during night (Österman and Redbo 2001; O’Driscoll et al., 2011, Norring et al., 2012). Nevertheless, in this study the effect was only noticeable after evening milking and not before it, also suggesting the effect of milk yield on nocturnal lying behaviour. In our study, the behaviour after deprivation was characterized by the cows spending a longer time lying down. In addition, cows engaged more in “lying inactive without ruminating” and performed more neck relaxed lying reportedly indicative of sleep (Ternman et al., 2012). This suggests a build-up of motivation to sleep through a solitary standing posture even though the cows were forced to stand only for 4 h. Earlier studies also demonstrated that short-term lying deprivation builds up the motivation to lie down (Metz, 1985; Cooper et al., 2007 and 2008; Krebs et al., 2011)
even though early studies of cow sleep by Ruckebusch (1974) based on long lying deprivation times, suggested that cows could also sleep while standing. In addition, our results show that the latency to lying down after milking tended to be shorter after the deprivation, again indicating higher motivation to rest as reported by Krebs et al. (2011). Cows also spent a shorter time standing and eating following the deprivation, which was expected because they were able to eat while forced to stand. Even though, this study yields some valuable information, a limitation is the short observation time as well as the effect of deprivation treatment as such. Preventing animals to lie down disturbs them and causes them stress, especially if they have not been allowed to adapt to the procedure. It is, however, likely that stress caused by a disrupted routine will reduce resting time during treatment, as we found increased resting after deprivation compared to baseline situation. Our results indicated that even short bouts of forced standing affected behaviour and increased the motivation to rest, which might have an effect on the overall welfare of cows in intensive dairy units. Loose housing systems impose some waiting time on cows for access to milking and access to feed or stalls (Fregonesi et al., 2007; Gomez and Cook, 2010). In addition, management practices such as collecting cows for waiting for hoof trimming and veterinary check-ups prevent lying and sometimes also feeding. Munksgaard et al. (2005) and Norring et al. (2012) suggested that high yielding dairy cows might already have insufficient time for resting partly, at least, due to their high nutritional demands. Further restrictions on lying behaviour as a result of management, even in the short term, but regularly, might strain their ability to cope and eventually affect their welfare because not getting enough rest does result in stress responses (Munksgaard and Simonsen, 1996; Fisher et al., 2002; Cooper et al., 2008). We recorded a longer duration and higher frequency of leaning behaviour during forced standing compared with during the baseline phase. However, as leaning was never observed while lying, and as lying was not possible during the forced standing period, we further compared leaning as a proportion of standing time. When controlling for standing duration, the above mentioned difference disappeared, thus indicating that leaning was not a sign of frustration due to the forced standing, but was increased merely due to the fact that the cows were standing during this entire period. Leaning has been suggested to be associated with thwarting of lying behaviour (Munksgaard and Simonsen, 1996). According to our results, leaning appeared to be independent of frustration due to a forced standing protocol. Instead, we observed frequent leaning among cows that had free access to lying down even before the forced standing protocol started. Leaning might rather be associated with tethering, which can pose difficulties to lying down movement patterns or lying comfort (Haley et al., 2001) and therefore leaning could be related to frustration due to thwarted lying. However, the predisposing factors and motivational background for formation of leaning behaviour in cows is still poorly understood.
M. Norring, A. Valros / Applied Animal Behaviour Science 176 (2016) 1–5
Numerous cows were recorded leaning during this study. All the cows were allowed to pasture during summers and were tethered during winter. Tethering has been suggested to trigger leaning or oral stereotypic behaviour in cattle compared with loose housing or a pasture environment (Redbo, 1993; Krohn, 1994). In our study, leaning cows differed in their behaviour by spending more time eating silage compared with cows that did not engage in leaning. There are few reports of leaning behaviour, but according to Redbo (1990) most stereotyping individuals used less time for eating. Furthermore, we registered no association between higher milk yield and leaning behaviour as reported by Redbo et al. (1992). Leaning in cows has not been reported from pasture environments, and there is no knowledge about the prevalence of such behaviour in different housing conditions. The present result adds some support to the hypothesis that leaning is related to thwarting lying behaviour. In conclusion, 4 h of lying deprivation increased resting motivation. Milk yield correlated with shorter lying duration when cows were motivated to rest. The lying time before evening milking was relatively short irrespective of the amount of milk in the udder. In this study, leaning behaviour was common among tie-stall cows. Conflict of interest None. Acknowledgement Lene Munksgaard is acknowledged for her help in planning the study, as well as for taking an active part in analysis and revising the manuscript. References Cooper, M.D., Arney, D.R., Phillips, C.J.C., 2007. Two- or four-hour lying deprivation on the behavior or lactating dairy cows. J. Dairy Sci. 90, 1149–1158. Cooper, M.D., Arney, D.R., Phillips, C.J.C., 2008. The effect of temporary deprivation of lying and feeding on the behaviour and production of lactating dairy cows. Animal 2, 275–283. Fisher, A.D., Verkerk, G.A., Morrow, C.J., Matthews, L.R., 2002. The effects of feed restriction and lying deprivation on pituitary-adrenal axis regulation in lactating cows. Livestock Prod. Sci 73, 255–263. Fregonesi, J.A., Tucker, C.B., Weary, D.M., 2007. Overstocking reduces lying time in dairy cows. J. Dairy Sci. 90, 3349–3354. Gomez, A., Cook, N.B., 2010. Time budgets of lactating dairy cattle in commercial freestall herds. J. Dairy Sci. 93, 5772–5781.
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Haley, D.B., de Passillé, A.M., Rushen, J., 2001. Assessing cow comfort: Effects of two floor types and two tie stall designs on the behaviour of lactating dairy cows. Appl. Anim. Behav. Sci. 71, 105–117. Krebs, N., Berry, S.L., Tucker, C.B., 2011. Restless behavior increases over time, but not with compressibility of the flooring surface, during forced standing at the feed bunk. J. Dairy Sci. 94, 97–105. Krohn, C.C., 1994. Behaviour of dairy cows kept in extensive (loose housing/pasture) or intensive (tie stall) environments. III. Grooming, exploration and abnormal behaviour. Appl. Anim. Behav. Sci. 42, 73–86. Medrano-Galarza, C., Gibbons, J., Wagner, S., dePassillé, A.M., Rushen, J., 2012. Behavioral changes in dairy cows with mastitis. J. Dairy Sci. 95, 6994–7002. Metz, J.H.M., 1985. The reaction of cows to a short-term deprivation of lying. Appl. Anim. Behav. Sci. 13, 301–307. Munksgaard, L., Jensen, M.B., Pedersen, L.J., Hansen, S.T., Matthews, L., 2005. Quantifying behavioural priorities—effects of time constraints on behaviour of dairy cows, Bos taurus. Appl. Anim. Behav. Sci. 92, 3–14. Munksgaard, L., Simonsen, H.B., 1996. Behavioral and pituitary adrenal–axis responses of dairy cows to social isolation and deprivation of lying down. J. Anim. Sci. 74, 769–778. Norring, M., Valros, A., Munksgaard, L., 2012. Milk yield affects time budget of dairy cows in tie-stalls. J. Dairy Sci. 95, 102–108. O’Driscoll, K., Gleeson, D., O’Brien, B., Boyle, L., 2011. Does omission of regular milking event affect cow comfort? Livest. Sci. 138, 132–143. Overton, M.V., Sischo, W.M., Temple, G.D., Moore, D.A., 2002. Using time-lapse video photography to assess dairy cattle lying behavior in a free-stall barn. J. Dairy Sci. 85, 2407–2413. Pomiès, D., Martin, B., Chillard, Y., Pradel, P., Rémond, B., 2007. Once-a-day milking of Holstein and Montbèliarde cows for 7 weeks in mid-lactation. Animal 1, 1497–1505. Redbo, I., 1990. Changes in duration and frequency of stereotypies and their adjoining behaviours in heifers, before, during and after grazing period. Appl. Anim. Behav. Sci. 26, 57–67. Redbo, I., 1993. Stereotypies and cortisol secretion in heifers subjected to tethering. Appl. Anim. Behav. Sci. 38, 213–225. Redbo, I., Jacobsson, K.G., van Doorn, C., Pettersson, G., 1992. A note on relations between oral stereotypies in dairy cows and milk production, health and age. Anim. Prod. 54, 166–168. Ruckebusch, Y., 1974. Sleep deprivation in cattle. Brain Res. 78, 495–499. Sprecher, D.J., Hostetler, D.E., Kaneene, J.B., 1997. A lameness scoring system that uses posture and gait to predict dairy cattle reproductive performance. Theriogenology 47, 1179–1187. Ternman, E., Hänninen, L., Pastell, M., Agenäs, S., Nielsen, P.P., 2012. Sleep in dairy cows recorded with a non-invasive EEG technique. Appl. Anim. Behav. Sci. 140, 25–32. Tuori, M., K. Kaustell, J. Valaja. E. Aimonen, E Saarisalo, and P. Huhtanen, 2000 Rehutaulukot ja ruokintasuositukset, Helsingin yliopisto, kasvintuotannon tarkastuskeskus, Maatalouden tutkimuskeskus. Helsinki, Finland. https:// portal.mtt.fi/portal/page/portal/Rehutaulukot/Tietosiilo/Arkisto/vanhat rehutaulukot/6FEB13EA9819F4C4E040A8C0033C5713. Österman, S., Redbo, I., 2001. Effects of milking frequency on lying down and getting up behaviour in dairy cows. Appl. Anim. Behav. Sci. 70, 167–176. Zobel, G., Leslie, K., Weary, D.M., von Keyserlingk, M.A., 2013. Gradual cessation of milking reduces milk leakage and motivation to be milked in dairy cows at dry-off. J. Dairy Sci. 64, 5064–5071.
Contents lists available at ScienceDirect
Applied Animal Behaviour Science journal homepage: www.elsevier.com/locate/applanim
The effect of lying motivation on cow behaviour Marianna Norring ∗ , Anna Valros Research Centre for Animal Welfare, Faculty of Veterinary Medicine, University of Helsinki, FI-00014 Helsinki, Finland
a r t i c l e
i n f o
Article history: Received 26 August 2015 Received in revised form 26 November 2015 Accepted 29 November 2015 Available online 10 December 2015 Keywords: Behaviour Dairy cow Lying Motivation Milk yield
a b s t r a c t Cows in dairy barns spend time standing while waiting for milking, accessing feed and entering the resting area. It has been suggested that high yielding cows may represent a trade off situation regarding eating and lying particularly in systems where there is a long waiting period before milking. We tested the effect on lying motivation of preventing cows of different production levels from lying down. The study included 14 primiparous cows and 14 multiparous cows in tiestalls at their 8th week of lactation. Cows were milked at approximately 06:00 and 18:00 h and milk yield was recorded. The behaviour of the cows was observed over 2 days when the cows were prevented from lying, either from 10:00 to 14:00 or from 14:00 to 18:00 h and during 2 preceding baseline days. Lying, eating, ruminating and lying with neck muscles relaxed (used as a behavioural indicator for sleep) were observed for 4 h before and 4 h after milking. The percentage time allocated to different behaviours per period was analysed using mixed models. In addition, stereotypic leaning behaviour was observed during the deprivation period. We established that the cows lay more after deprivation compared with during the baseline period for that time of day (21% vs. 34% before milking and 49% vs. 55% after milking). The cows spent less time eating after forced standing (23% vs. 21%). Cows used more time lying inactively without ruminating and lying with neck relaxed post treatment (11% vs. 19%). In addition, milk yield and lying time were correlated during the period after forced standing, indicating greater motivation in high yielding cows to rest. In total, 19 of 28 cows housed in tie stalls exhibited leaning behaviour. The duration of leaning was associated with the duration of standing. In conclusion, a 4-h standing period appeared sufficient to increase the motivation for lying and sleeping. Milk yield correlated with shorter lying duration when the cows were motivated to rest. The motivational background of the leaning behaviour in cows requires further investigation. © 2015 Elsevier B.V. All rights reserved.
1. Introduction Many studies have shown the importance of lying behaviour in cows (Munksgaard and Simonsen, 1996; Fisher et al., 2002; Cooper et al., 2008) and under time constraints dairy cows prioritize their lying time even over eating time (Munksgaard et al., 2005). Total daily lying time is higher in low yielding cows than in high yielding cows (Norring et al., 2012). A link between milk yield or anticipation of milking and lying was supported by the observation that time spent lying decreased towards the subsequent milking (Overton et al., 2002) and that cows milked frequently rested more at dawn than cows milked less frequently (Österman and Redbo, 2000). In addition, cows increased their mooing and stood more by the gate when one of the daily milkings was skipped (Pomiès et al., 2007) and a skipped milking event increased milk leakage
∗ Corresponding author. E-mail address: Marianna.Norring@Helsinki.fi (M. Norring). http://dx.doi.org/10.1016/j.applanim.2015.11.022 0168-1591/© 2015 Elsevier B.V. All rights reserved.
and udder firmness, and resulted in decreased total lying duration and shorter lying bouts (O’Driscoll et al., 2011). Changes in lying and standing durations were suggested to be indicative of udder discomfort during mastitis (Medrano-Galarza et al., 2012). Cows subjected to abrupt cessation of milking exhibited behaviours indicative of anticipated milking more than cows that were gradually weaned from milking (Zobel et al., 2013). Moreover, dry-off resulted in shorter duration of lying, indicating udder discomfort and higher motivation to be milked to be linked with reduced lying times (Zobel et al., 2013). The motivational background of increased standing duration of high yielding cows is still unknown. According to Norring et al. (2012) cows were lying down 22% of their time before evening milking and 50% of their time after milking. However, it is not clear why the lying duration prior to milking was lower than after milking. It might be caused by anticipation of milking, or by relaxation after milking. It is also possible that the resting motivation changes during the course of the afternoon due to some other reason. A way to study the underlying motivation, and the link to milk yield, more
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specifically, while controlling for e.g. diurnal variation, is to manipulate the lying motivation by depriving cows of the opportunity to lie down during a specific period. Lying deprivation induces both physiological and behavioural stress responses in dairy cows (Ruckebusch, 1974; Munksgaard and Simonsen, 1996; Fisher et al., 2002). Even short bouts of 2–4 h standing has an effect (Metz, 1985; Cooper et al., 2007, 2008; Krebs et al., 2011). Pressing or leaning the muzzle on stall constructions is described as leaning behaviour and this behaviour is suggested to be increased during lying deprivation and during tethering (Krohn, 1994; Munksgaard and Simonsen 1996). However, there are not enough studies about leaning behaviour to be able to fully understand how the environment and characteristics of individuals contribute to this behaviour. In order to shed further light on the reasons for the decreased lying observed prior to milking, we investigated the consequences of 4 h of forced standing on behaviour before and after evening milking. We also investigated whether milk yield was connected to the responses to deprivation of lying. In addition, the effect of lying deprivation on leaning behaviour was explored. 2. Material and methods 2.1. Animals and housing The time budget of 28 cows was observed over 4 days. We used Finnish Ayrshire cows in the 8th week of lactation with a mean of 58.3 days (SD 5.1) from parturition on the first day of observation. Fourteen of the cows were primiparous and the remainder had calved 2.5 (SD 1.1) times. The cows weighed a mean of 616 kg (SD 66). The gaits of the cows were visually scored according to Sprecher et al. (1997), and only cows with a normal gait score (1 or 2) and cows that were not being treated for any disease were included in the experiment. The University of Helsinki ethical committee approved the experiment. Water was offered ad libitum from a water bowl. Cows were milked at 06:00 and 18:00 h in their tie-stalls. Before and during the experiment, all cows were housed in the same insulated barn in tie-stalls (170 × 121 cm) with rubber mats (2 cm thick, Delaval, Sweden) bedded with sawdust. The cows were tethered to a neck bar. The cows were fed grass silage ad libitum . Silage was added approximately at 05:00, 12:00, 14:00, 15:00 and 16:00 h according to the working schedule of the caretakers. The leftover silage was removed daily. In addition, all cows were fed a mixture of 2 commercial concentrates (Suomen rehu, Espoo, Finland). Cows were fed concentrate according to milk yield following Finnish feeding recommendations, the daily amount ranging from 12.5 to 20.5 kg per cow (Tuori et al., 2000). The daily total amount of concentrate intake of each cow was divided among 6 equal meals and offered to the animals at approximately at 05:00, 08:00, 11:00, 14:00, 17:00, and 20:00 h. 2.2. Procedures and observations When a cow reached her 8th week of lactation she was placed in one of the experimental stalls. The cows were allowed to adapt to their new location in the barn for 2 days. Subsequently the behaviour of the animals was recorded at 14:00 to 22:00 h during 4 days. If a cow was inseminated filming was postponed for one day and the day of insemination was excluded. Data on baseline behaviour was collected on days 1 and 2. On days 3 and 4 cows were prevented from lying for a 4-h period each day. Lying was prevented using a girth that went around the cow’s trunk and was attached to the ceiling above her tie stall. Lying behaviour was prevented
Table 1 Postures and behaviours observed and the definitions of observations. Lying inactive without ruminating and lying with the neck relaxed were observed to estimate resting and sleeping time. Posture Lying Standing
Rump on the floor Standing on four feet
Behaviour Eating Ruminating Inactive without ruminating Lying with the neck relaxed
Other activities
Taking silage into the mouth and jaw moving, or muzzle in contact with silage and moving Jaw moving, not taking silage into the mouth Jaw not moving, head not moving, head up Jaw not moving, neck relaxed and head in contact with the feeding Table in front of the cow, or reversed along her body All other behaviours of the head not mentioned above, for example head moving actively, drinking, eating concentrate, licking, social behaviour, pressing the muzzle towards equipment
from 10:00 until 14:00 and from 14:00 until the evening milking at approximately 18:00 h. Half of the cows (14) were prevented from lying from 10:00 and the other half beginning from 14:00 h on day 3 and vice versa on day 4. The effect of lying deprivation on the behaviour of the cows was observed both before and after milking, in order to establish the effect of increased lying motivation on behaviour during this specific time period. The milk yield was measured according to milk recording standards at each milking; the accuracy achieved was ±200 g (Tru-test Limited, Pakuranga, New Zealand). Three regular, adjacent tie-stalls, in the middle of the row of tie-stalls, were equipped as experimental stalls. A total of 8 video cameras (Sanyo VCB-3372P, Watford, UK) were used to observe the cows’ behaviour. Time-lapse (Panasonic AG-6730, Secaucus, NJ) recording with 24 h time mode was used, and the pictures from all cameras were fed to a single tape using a multiplexer (Robot, Tyco, Princeton, NJ). 2.3. Behaviours Recordings were made of the lying or standing posture, head and jaw movements indicative of eating, rumination and sleeping behaviours (Table 1). The behaviours and postures were recorded through continuous observation (Observer Noldus, Wageningen, the Netherlands) and were analysed with an accuracy of 1 min while observing the videos. The bout of a single behaviour was considered to end when another behaviour began. Leaning behaviour was observed from 14:00 until 18:00 on 2 baseline days and on one day while the cows were prevented from lying from 14:00 to 18:00. The video analyses of leaning behaviour were made during separate viewings using continuous observation. We were not able to record if the leaning included the weight shift towards the head, thus all pressing or leaning or contact of the muzzle or nostril on stall constructions, including the feeding bowl or front curb, were recorded as leaning behaviour. 2.4. Statistical methods Based on visual inspection of histograms the data was considered to be normally distributed. Percentage time used in different behaviours during the 4-h period before and after evening milking was analysed using the mixed procedure of SPSS. The behavioural responses in time periods 14:00–18:00 and 18:00–22:00 h were analysed separately. Treatment (no forced standing, forced standing 10:00–14:00 or forced standing 14:00–18:00 h), order of treatment (forced standing at 10:00–14:00 followed by forced
M. Norring, A. Valros / Applied Animal Behaviour Science 176 (2016) 1–5 Table 2 Average percentage of time used in different behaviours during 4 h before evening milking (14:00 to 18:00 h) showing the difference between the baseline phase behaviour and post-standing (n = 28). % of Behaviour at 14:00 to 18:00 h
70
a
23 56 20 46
21 45 18 43
14:00-18:00
60
SE P-value
Baseline
50
Baseline Post standing 10:00 to 14:00 Eating Standing, not eatinga Ruminating Other active behaviours
3
1 2 1 2
0.031 0.001 ns 0.052
Post standing 10-14
*
40 % 30
*
Percent of standing time minus percent of eating time.
20 standing 14:00–18:00 h or vice versa), and parity (primiparous or multiparous) were fixed factors. Cow was included as a random factor. Separate mixed models were used to analyse the effect of milk yield on behaviour of lying-motivated cows. The behaviours before (14:00–18:00) and after (18:00–22:00 h) milking during control and treatment times were analysed separately. Cow was included as a random factor. Milk yield was used as a continuous measure and was included in the model as a regression variable. We used yield at the evening milking and average evening milk yield for the baseline behaviour measurement days. Parity was inserted as a fixed factor into the model. Latency from evening milking to lying down was analysed using a linear mixed model. Cow was considered a random factor and treatment (none, after standing 10:00–14:00 or standing 14:00–8:00 h) was inserted into the model as a fixed factor. Parity was inserted as a fixed factor. Milk yield was used as regression variable. The effect of treatment on the total duration of leaning and proportion of duration of leaning during standing was analysed using linear mixed models. Total duration and proportion of leaning from standing time were used in the analysis as an outcome measure. Cow was considered a random factor and day (average of 2 baseline days vs. treatment day) was inserted into the model as a fixed factor. In addition, the baseline behaviour, milk yield and parity of cows that performed leaning were compared with those for cows that did not engage in such behaviour. The characteristics of leaning cows were compared with those of the non-leaning cows using linear mixed models. The lying, eating, ruminating, lying with neck relaxed, inactive without ruminating, other active behaviours, milk yield and parity were analysed. All cows observed leaning at least once were classified as leaning. The model included day as a repeated effect and leaning behaviour as a fixed effect. 3. Results
10 0 70
Lying
18:00-22:00
Lying inacve
*
60
Baseline
50
Post standing 10-14 Post standing 14-18
40 % 30
20 10 0 Lying
Lying inacve
Fig. 1. Percent of time spent on resting behaviours before and after evening milking by different treatments and baseline. Lying inactive consists of behaviour classes lying inactive without ruminating and lying with neck relaxed (*P < 0.05).
The mean evening milk yield of the cows across the study period was 18 ± 0.3 kg. The milk yield decreased with duration of lying (P = 0.028, slope −1.97%/kg) after evening milking on the day when the cows were forced to stand from 10:00 to 14:00; but there was no effect of milk yield on duration of lying when cows were forced to stand from 14:00 to 18:00 h. There was no effect of milk yield on duration of lying before evening milking (P > 0.05). During the baseline period there was no effect of milk yield on duration of lying either before or after evening milking. Milk yield had no effect on other behaviours analysed.
3.1. Behaviour 4-h post treatment 3.2. Lying down latency after milking When the cows were forced to stand from 10:00 to 14:00 h they spent more time lying down and more time lying inactive during the following 4-h period compared with baseline behaviour, (Fig. 1). Animals spent less time eating and less time “standing and not eating” immediately after forced standing (Table 2). Treatment order had an effect on the 2 behaviours during the time period 14:00 to 18:00 h; cows that were assigned to forced standing from 10:00 to 14:00 first spent more time eating (24% vs. 20%, P = 0.042) and were less occupied by “other active behaviours” (40% vs. 49%, P = 0.003) compared with cows tested later. During the time period 18:00–22:00 h treatment order had an effect on eating behaviour; cows first assigned to forced standing from 10:00 to 14:00 h spent longer eating than cows tested in the other order (20% vs. 14%, P = 0.024).
When the cows were forced to stand for the last 4 h before milking, the latency period to lie down after milking tended to be shorter than baseline (16.6 a vs. 15.7 vs. 6.7 b ± 4 min: baseline, forced standing 10:00–14:00, forced standing 14:00–18:00 h, respectively; F2,55 = 2.8, P = 0.07). The milk yield or parity had no effect on latency to lie down after evening milking. 3.3. Leaning behaviour The frequency of leaning increased during the periods with forced standing and duration of leaning tended to be longer during the periods with forced standing compared with baseline periods (Table 4). However, there were no differences in proportion of time
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Table 3 Average percentage of time used in different behaviours during 4 h after evening milking (18:00 to 22:00 h) showing the difference between the baseline phase behaviour and post standing (n = 28). % of Behaviour at 18:00 to 22:00 h
Eating Standing, not eatinga Ruminating Other active behaviours a
Baseline
Post standing 10:00 to 14:00
Post standing 14:00 to 18:00
17 33b 24 32
18b 28c 24 31
15c 30 27 31
SE
P-value
1 3 2 2
0.025 0.040 ns ns
Percent of standing time minus percent of eating time. Different letters within rows indicate significant difference in post hoc tests.
b,c
Table 4 Mean and SE for leaning during a 4 h period from 14:00 to 18:00 for 28 cows.
Leaning duration min Frequency of leaning bouts Proportion of leaning while standing %
Baseline
Treatment
SE
P-value
7 3.5 3.1
8 5.8 3.5
2 1.1 0.9
0.07 0.008 ns
used for leaning when only the time spent standing was included. Leaning always took place while standing; we did not observe leaning while lying. Leaning behaviour was observed in 19 out of the 28 cows. The cows that leaned also spent more time eating silage during the baseline days than non-leaning cows (227 vs. 194 min/d; P = 0.048). There were no other differences between cows that performed leaning and those that did not. 4. Discussion This study supported earlier results indicating that during normal conditions, cows spend much less time lying down prior to milking than after milking. When cows were deprived of lying during a period between eight and four hours prior to milking, they increased the duration of lying down during the last four hours before milking, indicating an increased motivation for resting. Milk yield did not correlate to lying behaviour prior to milking neither in the baseline situation, nor after the deprivation treatment. We were thus not able to confirm that the amount of milk in the udder before evening milking had an effect on the duration of lying down during this period. In addition, we found that forced standing increased the frequency of leaning. According to our earlier findings (Norring et al., 2012) based on 24 h observation period with these animals, the cows with higher daily milk yield during the baseline phase had shorter lying times. Österman and Redbo (2001) suggested that a full udder prior to milking could hinder lying. O’Driscoll et al. (2011) observed udder tension, milk leakage and a shorter lying after a skipped evening milking. In addition, at the time of an omitted milking, cows stood near the milking parlour gate probably in anticipation of milking (Pomiès et al., 2007; Zobel et al., 2013). In previous studies shorter lying times were recorded around morning milking or during night (Österman and Redbo 2001; O’Driscoll et al., 2011, Norring et al., 2012). Nevertheless, in this study the effect was only noticeable after evening milking and not before it, also suggesting the effect of milk yield on nocturnal lying behaviour. In our study, the behaviour after deprivation was characterized by the cows spending a longer time lying down. In addition, cows engaged more in “lying inactive without ruminating” and performed more neck relaxed lying reportedly indicative of sleep (Ternman et al., 2012). This suggests a build-up of motivation to sleep through a solitary standing posture even though the cows were forced to stand only for 4 h. Earlier studies also demonstrated that short-term lying deprivation builds up the motivation to lie down (Metz, 1985; Cooper et al., 2007 and 2008; Krebs et al., 2011)
even though early studies of cow sleep by Ruckebusch (1974) based on long lying deprivation times, suggested that cows could also sleep while standing. In addition, our results show that the latency to lying down after milking tended to be shorter after the deprivation, again indicating higher motivation to rest as reported by Krebs et al. (2011). Cows also spent a shorter time standing and eating following the deprivation, which was expected because they were able to eat while forced to stand. Even though, this study yields some valuable information, a limitation is the short observation time as well as the effect of deprivation treatment as such. Preventing animals to lie down disturbs them and causes them stress, especially if they have not been allowed to adapt to the procedure. It is, however, likely that stress caused by a disrupted routine will reduce resting time during treatment, as we found increased resting after deprivation compared to baseline situation. Our results indicated that even short bouts of forced standing affected behaviour and increased the motivation to rest, which might have an effect on the overall welfare of cows in intensive dairy units. Loose housing systems impose some waiting time on cows for access to milking and access to feed or stalls (Fregonesi et al., 2007; Gomez and Cook, 2010). In addition, management practices such as collecting cows for waiting for hoof trimming and veterinary check-ups prevent lying and sometimes also feeding. Munksgaard et al. (2005) and Norring et al. (2012) suggested that high yielding dairy cows might already have insufficient time for resting partly, at least, due to their high nutritional demands. Further restrictions on lying behaviour as a result of management, even in the short term, but regularly, might strain their ability to cope and eventually affect their welfare because not getting enough rest does result in stress responses (Munksgaard and Simonsen, 1996; Fisher et al., 2002; Cooper et al., 2008). We recorded a longer duration and higher frequency of leaning behaviour during forced standing compared with during the baseline phase. However, as leaning was never observed while lying, and as lying was not possible during the forced standing period, we further compared leaning as a proportion of standing time. When controlling for standing duration, the above mentioned difference disappeared, thus indicating that leaning was not a sign of frustration due to the forced standing, but was increased merely due to the fact that the cows were standing during this entire period. Leaning has been suggested to be associated with thwarting of lying behaviour (Munksgaard and Simonsen, 1996). According to our results, leaning appeared to be independent of frustration due to a forced standing protocol. Instead, we observed frequent leaning among cows that had free access to lying down even before the forced standing protocol started. Leaning might rather be associated with tethering, which can pose difficulties to lying down movement patterns or lying comfort (Haley et al., 2001) and therefore leaning could be related to frustration due to thwarted lying. However, the predisposing factors and motivational background for formation of leaning behaviour in cows is still poorly understood.
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Numerous cows were recorded leaning during this study. All the cows were allowed to pasture during summers and were tethered during winter. Tethering has been suggested to trigger leaning or oral stereotypic behaviour in cattle compared with loose housing or a pasture environment (Redbo, 1993; Krohn, 1994). In our study, leaning cows differed in their behaviour by spending more time eating silage compared with cows that did not engage in leaning. There are few reports of leaning behaviour, but according to Redbo (1990) most stereotyping individuals used less time for eating. Furthermore, we registered no association between higher milk yield and leaning behaviour as reported by Redbo et al. (1992). Leaning in cows has not been reported from pasture environments, and there is no knowledge about the prevalence of such behaviour in different housing conditions. The present result adds some support to the hypothesis that leaning is related to thwarting lying behaviour. In conclusion, 4 h of lying deprivation increased resting motivation. Milk yield correlated with shorter lying duration when cows were motivated to rest. The lying time before evening milking was relatively short irrespective of the amount of milk in the udder. In this study, leaning behaviour was common among tie-stall cows. Conflict of interest None. Acknowledgement Lene Munksgaard is acknowledged for her help in planning the study, as well as for taking an active part in analysis and revising the manuscript. References Cooper, M.D., Arney, D.R., Phillips, C.J.C., 2007. Two- or four-hour lying deprivation on the behavior or lactating dairy cows. J. Dairy Sci. 90, 1149–1158. Cooper, M.D., Arney, D.R., Phillips, C.J.C., 2008. The effect of temporary deprivation of lying and feeding on the behaviour and production of lactating dairy cows. Animal 2, 275–283. Fisher, A.D., Verkerk, G.A., Morrow, C.J., Matthews, L.R., 2002. The effects of feed restriction and lying deprivation on pituitary-adrenal axis regulation in lactating cows. Livestock Prod. Sci 73, 255–263. Fregonesi, J.A., Tucker, C.B., Weary, D.M., 2007. Overstocking reduces lying time in dairy cows. J. Dairy Sci. 90, 3349–3354. Gomez, A., Cook, N.B., 2010. Time budgets of lactating dairy cattle in commercial freestall herds. J. Dairy Sci. 93, 5772–5781.
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