Comparison of time budget of behaviors ... - Wiley Online Library

Animal Science Journal (2008) 79, 518–525

doi: 10.1111/j.1740-0929.2008.00558.x

ORIGINAL ARTICLE Comparison of time budget of behaviors between penned and ranged young cattle focused on general and oral behaviors Toshie ISHIWATA,1 Katsuji UETAKE,1 Robert J. KILGOUR,2 Yusuke EGUCHI1 and Toshio TANAKA1 1

School of Veterinary Medicine, Azabu University, Sagamihara, Japan; and 2Agricultural Research Center, NSW Department of Primary Industries, Trangie, New South Wales, Australia

ABSTRACT To consider cattle welfare, time budget of behaviors of young cattle in pen and pasture conditions were compared. Behavioral observations of 103 steers in seven pens and 1136 steers in six ranges were performed during daylight over 3 days in each rearing condition. The pen condition had Japanese Black ¥ Holstein cross, and the pasture conditions had assorted breeds of Angus, Murray Grey, Shorthorn, Hereford, Santa Gertrudis and their crosses aged 5–15 months. The ranges varied in vegetation from extensive native pasture to intensive improved pasture. The daytime proportion of behaviors and activity patterns were compared between rearing conditions. The proportion of walking was much lower in the pen condition (1.0 ⫾ 0.2%) compared to all pasture conditions (at least 9.4 ⫾ 4.0%; all P < 0.05), but grooming (5.9 ⫾ 1.2%), investigating (2.5 ⫾ 0.2%) and tongue playing (1.1 ⫾ 0.2%) covered the loss. Fluctuation patterns of the proportions of eating (P < 0.001) and resting (P < 0.001) were different between pen and pasture conditions, whereas the pattern of the proportion of walking was not different. The proportion of eating in the pen condition (32.4 ⫾ 0.3%) was not different to that in one on the farm (42.2 ⫾ 12.6%), which had a thick improved pasture. However, the proportion of eating in the pen condition (32.4 ⫾ 0.3%) was lower than that in two farms (58.8 ⫾ 5.0% and 61.6 ⫾ 16.5%) with low pasture availability (both P < 0.05). The proportions of oral behaviors such as self-grooming (3.9 ⫾ 1.0%), allogrooming (2.0 ⫾ 0.2%), licking objects (1.3 ⫾ 0.4%) and tongue-playing (1.1 ⫾ 0.2%) compensated for the lack of eating. These results indicate that we can learn a well-balanced proportion of behaviors for cattle by making comparisons of time budget of behaviors and activity pattern between various rearing conditions. In addition, it is indicated that cattle under pen conditions under a restricted feeding period might compensate for a lower time spent feeding by performing other oral behaviors.

Key words: beef cattle, behavior, rearing condition, welfare.

INTRODUCTION The welfare of farmed animals is related to the extent to which they can adapt, without suffering, to environments designed by humans (Animal Welfare Institute 2005). In an attempt to ensure that a farmed animal is raised in an acceptable rearing environment, the ‘Five Freedoms’ firstly defined by the UK’s Farm Animal Welfare Council (RSPCA 2000) are used as a global standard. These five freedoms are: (1) freedom from hunger and thirst; (2) freedom from discomfort;

© 2008 The Authors Journal compilation © 2008 Japanese Society of Animal Science

(3) freedom from pain, injury and disease; (4) freedom to express normal behavior; (5) freedom from fear and distress. The inhibition of freedoms (1) (2) (3) and (5) would usually result in physical as well as psychological wellbeing problems. The freedom to express normal behavior has proved more difficult to justify as Correspondence: Katsuji Uetake, School of Veterinary Medicine, Azabu University, Sagamihara 229-8501, Japan. (Email: [email protected]) Received 15 February 2007; accepted for publication 1 October 2007.

BEHAVIORS OF PEN AND RANGE CATTLE 519

an animal welfare indicator since it often does not result in physically measurable improvements in animal welfare (Young 2003). However, it is not argued that expression of normal behavior is unimportant to animal wellbeing. Broom (1999) has described a variety of normal behaviors expressed as one measure in a list of good animal welfare indicators. One of the means of allowing an animal to express normal behavior is to provide environmental enrichment. Environmental enrichment is defined as the provision of stimuli that promote the expression of species-appropriate behavioral and mental activities (Reinhardt & Reinhardt 2002). It is a dynamic process in which changes to structures and husbandry practices are made with the goal of increasing behavioral choices to animals and drawing out their species appropriate behaviors and abilities, thus enhancing animal welfare. The study and implementation of environmental enrichment has been dominated by two approaches since its inception, these approaches being the naturalistic approach and behavioral engineering. The former relies upon creating the wild environment in captivity to provide stimulation for captive animals, while the later relies upon providing devices and machines that animals operate to receive some form of reward, usually food (Young 2003). Scientists who favor one or other of the two approaches have often been critical of the other. However, the important thing is not to argue that one approach is better than the other but to develop the methodology so that animals can express their species appropriate behaviors in captivity. To achieve this ultimate goal, an in-depth understanding of the time budget of behaviors and diurnal pattern of activity in the species’ natural environment is an essential prerequisite. Once this has been completed their species-appropriate behaviors that animals have a high motivation to perform can be determined along with the consequences of preventing the performance of these behaviors. With the above in mind, the first objective of this study was to determine the time budget of general behavior and diurnal activity pattern of cattle reared either in a conventional pen and of cattle reared in extensive and intensive pastures, which varied in vegetation. One functional category of behavior that we investigated in depth as the second objective of this study was oral behavior, since it has been shown to be a behavior that cattle are strongly motivated to perform (Phillips 2002a).


MATERIALS AND METHODS Pen condition A total of 103 steers aged 7–11 months were observed at one farm in Sano (36°N, 138.5°E), Tochigi prefecture, Japan. The steers were Japanese Black ¥ Holstein steers kept in seven pens (width 6.0 m ¥ length 9.5 m each) with between 12 and 16 steers (3.6–4.8 m2/head) in each pen. At one end of the pen, the steers were permitted access to a feeding alley (length 6.0 m) for grain feed, while at the other end of the pen, the steers had access to a wooden trough (width 0.7 m ¥ length 1.8 m) for hay and a self-filling concrete water bowl (diameter 0.5 m). The pens used in this study were in a part of two open-sided barns and each pen was divided by a metal fence 1.4 m in height. The steers were observed by scan sampling every 10 min during three mornings (from dawn till 11.50 hours) and three afternoons (from 12.00 hours till dusk) during March 2004. The duration of all morning observations was 6 h 10 min, while the minimum and maximum lengths of the afternoon observations were 5 h 50 min and 6 h, respectively. The observations were made by two experienced persons with the unaided eye while walking up and down the feeding alley. The minimum and maximum temperatures during the six observations ranged from -1.2 - +5.7 (+3.4 ⫾ 2.7)°C to +10.3 - +24.4 (+15.1 ⫾ 5.9)°C, respectively. Stormy weather characterized by high winds and/or precipitation was not encountered during any observations. The steers were provided a commercial concentrate diet based on the average body weight in each pen in expectation of DG 1.0–1.2 kg, twice daily between 08.30 and 08.40 hours and between 15.40 and 16.00 hours at the feeding alley. The steers were also allowed free access to a trough containing Italian ryegrass hay. The hay was added to the trough at the same time as the concentrate diet was provided and also between 11.00 and 11.30 hours. The diet contained 57% of grain (corn, wheat flour and soy flour), 18% of bran (corn gluten feed and wheat bran), 14% of plant-origin oil meal (soybean oil meal and rapeseed oil meal) and 11% of the other additives (alfalfa meal, molasses, calcium carbonate and common salt). The steers were allowed free access to the water bowl.

Pasture conditions A total of 1136 steers kept on six commercial beef cattle farms near Dubbo (33°S, 148.5°E), New South Wales, Australia were observed. The details of the farms are shown in Table 1. Observations were conducted on one farm in both 2003 and 2004 (farms C03 and C04, respectively), but on different animals each year. The majority of animals were British breeds and crosses, although there were some Santa Gertrudis and Santa Gertrudis cross animals. Since the growth rate of British breeds is normally in the low to moderate range (Wallis 1994), there is not a big difference from that of Japanese Black ¥ Holstein steers, which are in the moderate range. The subject steers were 5–15 months of age. The steers were continually at pasture with no housing and were able to graze at times of their own choosing. One important difference between the farms was the amount of pasture available. A qualitative assessment of the pasture


Plant height & density¶

†Farm C03 and C04 were observed at the same range for 2 years in 2003 and 2004, respectively. ‡This artificial pasture was used for grazing after mowing. §This alfalfa pasture was used for grazing before mowing. ¶Comparative assessment by experimenters.

High & High Middle & Middle

High & High

Wild grasses Wild grasses Vegetation

Wild grasses & ryegrass Low & Low

Low & Low

Native pasture 220, 100 Pasture type & area (ha)

Native pasture 250

Native pasture 250 Artificial pasture‡ 50 Ryegrass

Native pasture 60–150

Rotational grazing (mainly in artificial pasture) Native pasture 420 Artificial pasture§ 40 Alfalfa Rotational grazing Set grazing Pasture Grazing system

Set grazing

Rotational grazing (2–3 days interval)

140 125.1 5–14 Angus, Angus X Herefords, Angus X Shorthorns 150 143.4 6–12 Shorthorns, Shorthorns X Santa Gertrusdis 500, 170 371.4, 166.6 6–12 Angus, Shorthorns, Murray Grey, Herefords, Brahmans, Santa Gertrudis, & various crosses 120 103.1 6–12 Angus 56 47.5 12–15 Murray Grey Subject ainmal Number of steers Number of steers observed Age (month) Breed

Farm D Farm C03, C04† Farm B Farm A Variable

Table 1

Information about subject animal and pasture conditions observed

Farm E

520 T. ISHIWATA et al.


indicated that pasture quantity on two of the farms (A and B) was low as indicated by the low pasture height and density. On another two of the farms (D and E), pasture height and density were high, whereas on the fifth farm (C), the one that was studied in 2003 and 2004, pasture availability was intermediate. The steers were observed by scan sampling every 15 min over three mornings (from dawn till 11.45 hours) and three afternoons (from 12.00 hours till dusk). Both morning and afternoon observations were conducted at intervals of three or four days during August and September in 2003 and 2004. The minimum and maximum lengths of the morning observations were 5 h 30 min and 5 h 45 min, respectively, while the minimum and maximum length of afternoon observations were 5 h 30 min and 6 h, respectively. The observations were made by three experienced persons through binoculars from such a distance that the animals were unaware of their presence. In cases where the steers broke up into small groups in a pasture, the observers separated taking care not to observe the same steer group. The minimum and maximum temperatures during the six observations at each farm in 2003 ranged from +0.3 - +11.5 (+4.1 ⫾ 3.2)°C to +12.8 - +21.1 (+16.0 ⫾ 2.2)°C and from +0.2 - +14.1 (+5.0 ⫾ 3.3)°C to +13.4 - +27.2 (+19.5 ⫾ 4.2)°C, respectively, in 2004. During periods of inclement weather, observations were not carried out since such conditions caused the steers to curtail their activity and simply stand in the pasture until the weather passed.

Behavioral observation At any one scan, the behavioral state of every steer visible was recorded. In cases where the observers were forced to separate due to the animals forming subgroups, two observers remained together, one acting as observer, the other as recorder while the third observer recorded the behavioral state data on tape from which the behavioral states were later transcribed. The behavioral states recorded were eating (including grazing and browsing under pasture conditions), drinking water, excreting, walking, investigating (including sniffing and licking objects), grooming (including self- and allogrooming), rubbing (i.e. grooming with objects) playing (including mock fighting and mounting), tongue-playing, and resting. Because of the distances involved and because the animals tended to form large groups when resting in the shade, it was difficult to determine whether they were ruminating or simply resting. For this reason, ruminating was recorded as resting. Standing and lying down were also recorded with the behavioral states. In addition, all the behaviors observed in this study were reclassified into oral behaviors and the other behaviors. Oral behaviors consisted of eating, drinking water, self-grooming, allogrooming, licking objects and tongue-playing.

Statistical analyzes Because the scans were carried out every 10 min in Japan and every 15 min in Australia, the proportion of steers engaged in each behavioral state was analyzed for each 30 min observation in order to maintain synchrony. The mean proportion of animals classified in a given behavioral state during daylight hours was compared between rearing



conditions by Tukey’s honestly significant difference adjustment (Tukey’s HSD test) for multiple comparisons (d.f. = 12, n = 3 for each rearing condition). Difference in diurnal activity pattern was tested by statistical significance of a crossed factor of rearing condition ¥ time (d.f. = 161, 368) in twoway repeated measures ANOVA.

Overall comparison of time budget of behaviors between pen and pasture conditions The mean proportion of steers engaged in walking during daylight hours was significantly lower in the ¯ ⫾ SD = 1.0 ⫾ 0.2%) compared to in all farms pen (X (9.4 ⫾ 4.0% at lowest on farm B; all P < 0.05). The proportions of steers investigating (2.5 ⫾ 0.2%), grooming (5.9 ⫾ 1.2%) and tongue-playing (1.1 ⫾ 0.2%) covered a loss in the proportion of walking (all P < 0.05 in comparison to all farms) (Fig. 1). The proportion of steers engaged in eating in the pen (32.4 ⫾ 0.3%) was not significantly different to that in farm E (42.2 ⫾ 12.6%), one of the farms with the highest pasture quantity. However, the proportion of steers engaged in eating was significantly higher in farms A (58.8 ⫾ 5.0%) and B (61.6 ⫾ 16.5%; both P < 0.05), the two farms with the lowest pasture availability. Although the differences between the farms were not statistically significant, the proportion of steers engaged in eating during daylight hours tended to decrease as their pasture availability increased (Fig. 1). The proportion of steers that performed resting in farm E (41.3 ⫾ 9.1%) was also not different to that in the pen (53.2 ⫾ 1.7%). Especially the proportion of steers engaged in lying in this farm (24.2 ⫾ 2.3%) was highest of all farms and showed similar level of that in the pen condition (32.2 ⫾ 1.1%) (Fig. 2). The proportion of steers engaged in resting was not different between pasture conditions, but it tended to increase as their pasture availability increased (Fig. 1). Time diurnal patterns in the proportions of steers eating (P < 0.01; Fig. 3a) and steers resting (P < 0.001; Fig. 3b) were significantly different between rearing conditions. This was mainly due to the fact that there were three distinct peaks of eating in the pen and these coincided with the delivery of concentrate and hay (Fig. 3a). At pasture, however, there was no evidence of a consistent peak of grazing through the day. There were no significant differences between the pen and the pasture in the pattern of steers walking (Fig. 3c).


80

Proportion of steers (%)

RESULTS

100

Resting Tongue-playing Playing

60

Rubbing Grooming Investigating

40

Walking Excreting Drinking water Eating

20

0 A

B

C03 C04

D

E

Pen

Farm

Rearing condition

Figure 1 Mean proportion of steers engaged in each behavior during daylight hours in each rearing condition. Tukey’s honestly significant difference adjustment (all P < 0.05): resting: pen > farms A, B, C03, C04, D in pasture conditions; tongue-playing grooming and investigating: pen > all farms in pasture conditions (A, B, C03, C04, D, E); walking: pen < all farms in pasture conditions (A, B, C03, C04, D, E); eating: pen < farms A, B in pasture conditions.

The mean proportion of pen-housed steers observed to be walking was very low as already mentioned previously.

Comparison of oral behaviors between pen and pasture conditions The mean proportion of steers engaged in all oral behaviors (including eating, drinking water, selfgrooming, allogrooming, licking objects and tongueplaying) during daylight hours was not significantly different between rearing conditions. In the pasture conditions on the farms, the proportion of steers eating on Farms A, B, C03, C04, D and E accounted for 96.1%, 98.2%, 96.5%, 95.4%, 96.9% and 95.1% of all oral behaviors, respectively. On the other hand, the proportion of steers eating in the pen condition accounted for 77.4% of all oral behaviors. There were, however, significant differences between rearing conditions for the mean proportion of steers engaged in self-grooming, allogrooming, licking objects and tongue-playing (all P < 0.001). The proportions of



100


80 Standing

60

Lateral lying Lying

40

20

0 A

B

C03 C04

D

E

Pen

Farm

Rearing condition Figure 2 Mean proportion of steers engaged in each posture during daylight hours in each rearing condition. Tukey’s honestly significant difference adjustment (all P < 0.05): lying: pen > farms A, B, C03, C04 in pasture conditions.

steers engaged in self-grooming (3.9 ⫾ 1.0), allogrooming (2.0 ⫾ 0.2), licking objects (1.3 ⫾ 0.4) and tongue-playing (1.1 ⫾ 0.2) in the pen condition were higher than those in all pasture conditions (all P < 0.05 in comparison to all farms) (Table 2). Time diurnal patterns in the proportions of steers observed performing oral behaviors other than eating and drinking water were significantly different between rearing conditions (P < 0.01; Fig. 3d). The greatest proportion of steers observed performing oral behaviors other than eating and drinking water was observed in the pen condition, with two major periods in the morning and one major period in the afternoon being identified. The peaks occurred before feeding in the morning and after eating in the morning and afternoon. At pasture, the proportion of steers performing oral behaviors other than grazing and drinking water was overall much lower than that in pen environments, with no obvious peaks.

DISCUSSION The mean proportion of steers engaged in walking during daylight hours was extremely low (only 1%)


in pen conditions compared with pasture conditions (approximately 10% at lowest), but time serial change of the proportion of steers walking in there showed similar fluctuation patterns to those in pasture conditions. Motivation for locomotion is known to increase with the duration and severity of resource restriction, especially of space (Dellmeier et al. 1990 as cited by Phillips 2002b). For example, Hart et al. (1993) have reported that cows on a vast pasture (207 ha) traveled farther (6.1 km/day) than cows on a small pasture (24 ha; 3.2 km/day). Of course the motivation for locomotion is influenced by many other factors, both genotypic and environmental (Zeeb et al. 1983 as cited by Phillips 2002b). Breed or genotype difference in behavioral traits (Gare 2004) would be a possible confounding factor in the result of this study. Additionally, grazing system as an environmental factor can affect time spent traveling and the distance walked. They generally increase as the frequency of rotation of pastures increase (Walker 1989). However, differences in walking proportion among ranges that had different grazing systems and kept different breeds and/or crosses in this study were not statistically significant. Considering all these factors together, in some way or other the proportion of steers engage in walking during daylight hours would be encouraged to promote up to 10% even in a pen condition. As pasture quantity got higher, the daytime proportion of steers eating and that of steers resting in a pasture tended to decrease and increase, respectively. They leveled off at approximately 40% in the pasture with highest quantity (farm E), and became not to differ from those in a pen condition. Furthermore, the daytime proportion of steers lying accounted for nearly 20% of all postures both in the farm E and in the pen condition. Grazing time per unit area might possibly differ among pasture types because of variation in preference (Gammon & Roberts 1980). But Walker (1989) has reported that total grazing and sleeping time do not differ among grazing treatments. Not only do they not differ between rotational grazing and continuous grazing, but they do not differ between different stocking densities. Breed or genotype difference in the motivation to eat might have the potential to influence the outcome of this study. Thus it can be recommended that the mean proportions of steers perform eating and resting during daylight hours should occupy 40%, respectively. At lowest lying steers should account for 20% of all steers in the daytime.



(a) Eating


Feed delivery (only to the pen)

Feed delivery ( only to the pen)

100

A B C03 C04 D E Pen

80 60 40 20 0 6:30

8:00

9:30

11:00

12:30

14:00

15:30

17:00

6:30

8:00

9:30

11:00

12:30

14:00

15:30

17:00

8:00

9:30

11:00

12:30

14:00

15:30

17:00

15:30

17:00

( b) Resting 100 80 60 40 20 0

( c) Walking

50 40 30 20 10 0 6:30

( d) Oral behaviors other than eating and drinking water 40 30 20 10 0 6:30

8:00

9:30

11:00

12:30 Hour

14:00

Figure 3 Time serial changes in the proportion of steers engaged in eating (a), resting (b), walking (c) and oral behaviors other than eating and drinking water (d) in each rearing condition.




Table 2

Mean (⫾ SD) proportion of steers engaged in each oral behavior during daylight hours in each rearing condition

Behaviors

Rearing condition Farm A

Self-grooming Allogrooming Licking objects Tongue-playing

B

1.0 ⫾ 0.1 0.6 ⫾ 0.5b 0.0 ⫾ 0.0b 0.00 ⫾ 0.00b b

C03

0.5 ⫾ 0.0 0.3 ⫾ 0.2b 0.0 ⫾ 0.0b 0.01 ⫾ 0.01b b

Pen

C04

1.0 ⫾ 0.3 0.3 ⫾ 0.2b 0.0 ⫾ 0.0b 0.05 ⫾ 0.03b b

D

1.6 ⫾ 0.3 0.4 ⫾ 0.1b 0.0 ⫾ 0.0b 0.15 ⫾ 0.13b b

E

0.9 ⫾ 0.3 0.2 ⫾ 0.1b 0.0 ⫾ 0.0b 0.00 ⫾ 0.00b b

1.4 ⫾ 0.5b 0.3 ⫾ 0.0b 0.0 ⫾ 0.0b 0.01 ⫾ 0.01b

3.9 ⫾ 1.0a 2.0 ⫾ 0.2a 1.3 ⫾ 0.4a 1.11 ⫾ 0.24a

Different letters indicate significant differences (P < 0.05).

In a pen condition, specific categories of behaviors such as investigating, grooming and tongue-playing appeared to act as alternative behaviors for cattle to cover a loss in activity. In a small space where cattle are fed concentrate diet and hay, there is no need for walking around to seek grassy areas. Cattle seemed to engage in these behaviors to spend the spare time, instead of engaging in walking necessarily. In conflicting and/or frustrating circumstances, self-grooming and tongue-playing have been described as ‘disturbed behaviors’ (Sato 1999). Ruminating is also known to increase because of long-term frustration (Sato 1997). It is unlikely that the behaviors such as investigating, grooming and tongue-playing shown more in pen condition occurred out of frustration. More likely it is because the steers in pen conditions avoided the need to walk around for food and so they would not have a high motivation to walk. So it might be appropriate that these behaviors should be called ‘alternative behavior’ for the spare time or activity. Stereotyped behaviors shown in cattle relate to oral behaviors, such as bar-biting and tongue-rolling (Phillips 2002a). Cattle should have a high motivation to move their mouth and tongue as a species-specific behavior. In this study, the steers reared in intensive pen environments and fed a concentrate ration spent less time eating than steers reared in sparsely vegetated pasture. However, total proportion of steers performing all oral behaviors was not significantly different between rearing conditions. This result shows that cattle under any environmental conditions engaged in some sort of oral behaviors for a certain proportion of the daytime. Especially in an intensive pen environment, cattle might perform more oral behaviors other than eating to compensate for the lack of occurrence of feeding behavior. The lack of oral behaviors caused by the loss of eating might be compensated by oral behaviors except for eating as ‘compensatory behaviors’. In this study, tongue-playing


was observed in the pasture conditions as well. So tongue-playing is unlikely to be a typical behavior in intensive pen environments. Additionally, the proportion of tongue-playing was not excessive in either environment. Therefore, tongue-playing observed in this study is regarded as a normal behavior just like investigating, self-grooming and allogrooming. Self-grooming, allogrooming, licking objects and tongue-playing were observed more before and after eating. In artificially reared calves, the frequency of calf-directed oral behavior such as cross-sucking was the greatest in the 10 min following milk ingestion (de Passillé et al. 1992; Lidfors 1993; Bokkers & Koene 2001; Margerison et al. 2003). de Passillé et al. (1992) and Lidfors (1993) have suggested that the ingestion of milk stimulated sucking in calves and increased the motivation to perform sucking behavior. Tongueplaying observed in tethered beef cattle followed feeding and was followed by the other tonguemovement behaviors (Sato et al. 1994). In cattle in an intensive environment, the ingestion of the concentrate diet should stimulate oral behaviors and increase the motivation to perform oral behaviors. Although the steers were allowed free access to hay, eating hay was observed after eating the concentrate diet. In this study, however, the oral behaviors other than eating were also observed before morning-feeding. There is a possibility that the designated feeding time stimulated prefeeding oral behaviors. To conclude, comparisons of time budget of behaviors and activity patterns between pen and various pasture conditions indicated that penned cattle evidently decreased the expression of walking, although its time serial fluctuation pattern was maintained, but they spend their spare time performing other behaviors such as investigating, grooming and tongueplaying. Furthermore, the comparisons indicated that cattle in pen conditions under a restricted feeding period might compensate for the oral behaviors caused



by a lower time spent feeding by performing the other oral behavior, such as self-grooming, allogrooming, licking objects and tongue-playing. As compared to range with the high pasture availability, the implication of the results in this study might be tentative criteria of a well-balanced time budget of behaviors for young cattle: (i) the mean proportion of steers that engage in walking during daylight hours had better promote up to 10% even in a small pen condition; and (ii) the mean proportions of steers perform eating and resting in daylight should maintain 40%, respectively. Lying steers should account for at least 20% of all steers during daylight hours. The level of expressing behaviors is determined by many factors, both genotypic and environmental. So of course we need further studies to come up with the ultimate criteria of welfare-satisfied time budget of behaviors for cattle.

ACKNOWLEDGMENTS We gratefully acknowledge Mr Yoshio Shinohara and his staff at beef farm ‘Kotora’ for providing animals and facilities. We are also grateful to the beef cattle farmers and family at ‘Bowen Park’, ‘Glenfield’, ‘Inverary’, ‘Waitara’, ‘Weemabah’ for their contribution to our study.

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