Enhancing movement opportunity and fulfilling behavioural needs for movement- restricted cattle by providing different conditions of access to exercise

Enhancing movement opportunity and fulfilling behavioural needs for movement- restricted cattle by providing different conditions of access to exercise | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article Enhancing movement opportunity and fulfilling behavioural needs for movement- restricted cattle by providing different conditions of access to exercise Marjorie Cellier, Elise Shepley, Nadège Aigueperse, Marianne Villettaz Robichaud, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4953052/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 18 Feb, 2025 Read the published version in Scientific Reports → Version 1 posted 10 You are reading this latest preprint version Abstract Intensification in animal productions has led to increased indoor confinement for animals, thereby limiting their opportunities to fulfill some behavioural needs such as exploration and locomotion. This poses a crucial question as to how these restrictions can be alleviated by providing additional space designed with that specific purpose in mind. Working with cows housed in a movement-restricted environment, our study aimed to: 1) quantify how providing an exercise area outside the normal housing environment affects locomotor behaviour; 2) evaluate how exercise access conditions can contribute to optimize benefits for animals; 3) investigate the cows’ time budgets during exercise access. Six trials involving different exercise access conditions (indoor vs outdoor, outing duration, exercise area size) enrolled 141 tie-stall-housed lactating Holsteins. A meta-analysis compared daily steps for exercise vs non-exercise treatments, while generalized linear mixed models determined exercise conditions’ impact on daily steps. Providing 1-hour exercise access increased daily steps by 53% (304 more steps), influenced by type of access (167 more steps outdoor vs indoor), larger space (146 more steps) and longer outings (84 more steps). Cows spent 50–85% of exercise time idle, exploring (5–20%) and socializing (5%). Our results highlight the significant impact of 1h daily exercise on movement-restricted cattle. Biological sciences/Zoology/Animal behaviour Biological sciences/Zoology behavioural needs locomotor behaviour movement opportunity steps Figures Figure 1 Figure 2 Figure 3 Introduction In recent decades, the animal production, including dairy industry, has undergone an intensification characterized by changes in animal housing, notably an increase in indoor housing [ 1 ] with limited to non-existent outdoor access. Arguments put forward for the use of this type of system are, among others, individualized care, feed rationing, and reduced risk of agonistic behaviours [ 2 ]. This change has been met with public criticism by a society increasingly concerned about farm animal welfare [ 3 ]. What is often missing from these debates, however, is the perspective of the cow. As we cannot ask her directly, other means must be utilized in order to shed light on this important stakeholder within this debate. How does she use the space provided to her and what utility does it have? Can other elements be introduced to mitigate the negative effects of certain housing systems, or bring additional positive ones? If so, how does she perceive and make use of these elements? We often rely on what is known as environmental enrichment to add to the captive environment. This is a broad term that covers any change or addition to the living environment or routine of an individual with the goal of providing more opportunities for choice, agency, and natural behaviours. The effectiveness and utility of any enrichment can then only be evaluated through the perception of the individual and will thus depend on how the cow perceives and uses what is provided to her [ 4 ]. Moreover, evidence suggests that the greatest positive benefit to animals comes from additions that fulfil existing behavioural needs [ 5 ], further emphasizing our need to understand the cow’s existing needs and perception of her environment. One such need impacted by the increase of indoor housing is the need for movement and particularly the need for locomotion. The provision of exercise area access by increasing movement opportunity, is a potential solution to meet a cow’s needs for movement opportunity, as well as a variety of other natural behaviours not available to her in her home restricted environment [ 6 ]. For example, cows provided with regular access to an exercise area get more opportunities to engage in social interactions, to explore their environment, and to groom hard-to-reach body parts such as their hindquarters [ 7 ]. Movement opportunity can be defined as the level and ease of movement provided to an animal in its housing environment [ 8 ]. This includes the number of steps achieved by the cow [ 9 ], freedom of movement, and locomotor behaviours [ 8 ]. The average number of daily steps taken by cows, differs across housing systems: it ranges from 700 steps/day in tie-stall housing to 4000 steps/day on pasture, with free-stall housing sitting in-between at 1500 steps/day (reviewed by [ 9 ]). These quantitative data support the idea that systems offering more opportunity of movement can positively influence the level of locomotor behaviour performed by a cow (i.e. the cow will make use of the additional space provided). However, new scientific knowledge is necessary relating to the benefits of providing exercise access and the conditions of exercise access, like outdoor or indoor access, required to enhance such benefits. In particular, we need to know if and how cows actually use the exercise area as a movement opportunity to meet their locomotion needs by increasing their locomotor activities. Moreover, cow motivation to engage in locomotor behaviours such as walking, trotting, or running has been studied both in individuals kept in tie-stalls given access to an exercise area [ 10 ]and in those usually housed on pasture (summer) or free-stalls (winter) kept tied for the duration of the experiment [ 11 ]. Though being tied reduced locomotor behaviour for these cows, when additionally provided with daily access to an exercise area (e.g., indoor exercise area, outdoor paddock, or pasture), they showed an increase in locomotor behaviours [ 10 , 11 ]. However, it is not known to what extent access to an exercise area for cows continuously living in a movement-restrictive environment (i.e., cows that are not used to moving) affects their level of locomotor behaviour, most notably in terms of number of steps. It is further unknown how the conditions of access to exercise, such as the duration of access or the size of the exercise area, can influence these outcomes. Our study aimed to 1) quantify how providing movement opportunity outside of the stall impacts the locomotor behaviour of cows housed in a movement-restricted environment; 2) evaluate which conditions of access to exercise optimize locomotor behaviour: indoor vs outdoor, duration of outing, size of the exercise area; and 3) investigate which other behaviours cows perform when granted access to an exercise area. We hypothesized that regardless of the specific conditions, providing access to exercise would lead cows to take more steps than when confined to their stalls. We also hypothesized that cows would take a similar number of steps regardless of the conditions of access to the exercise area and that it is the overall experience provided to the cow outdoors (companionship, etc.) rather than the exercise area per se that would have a greater influence on the increase in locomotor behaviour in cows provided with regular exercise access. Finally, we hypothesized that cows would engage in a variety of behaviours (as opposed to solely engaging in locomotor behaviours) when provided with opportunity for movement outside of their stalls. Results Dataset 1. Exercise vs non-Exercise treatments: Meta-analysis Descriptive statistics for each trial are shown in Table 3. The forest plot of the meta-analysis is presented in Figure 1. The meta-analysis of all comparisons between Exercise and non-Exercise treatments resulted in a between-study heterogeneity variance estimated at τ² = 2686 (95% Confident Interval ( CI ): 0-26859), with an I² value of 9.5% (95% CI: 0% – 68.1%), meaning that less than 10% of the variation is due to between-study heterogeneity. The nonsignificant Q score suggests that the studies represent samples from a common population and that the coalescing of their data was justified (Q = 8.84, degrees of freedom ( df ) = 8, P = 0.356). The prediction interval ranged from g = 151 to 457. The number of daily steps was significantly higher when cows had access to an exercise area, with an average increase of 304 steps per day (95% CI: 215-393; P < 0.001), equivalent to 53% more steps (923 ± 65 daily steps) compared to the 578 (± 32 daily steps) mean daily steps taken in the stall. Dataset 2. Type of exercise area The number of daily steps differed depending on whether cows had access to an indoor or outdoor exercise area. Cows took 167, or around 20%, more daily steps in an outdoor exercise area (987 ± 50.3 daily steps) than in an indoor one (820 ± 42.4 daily steps; χ² = 44.980, df = 1, P < 0.001). Dataset 3. Duration of outing and size of the exercise area The number of daily steps was influenced by the duration of the outing (χ² = 10.090, df = 1, P = 0.002) and by the size of the exercise area (χ² = 18.317, df = 3, P < 0.001). Cows took 84, or around 9%, more daily steps when staying in the exercise area for 2 hours (1009 ± 22.8 daily steps) compared with 1 hour (925 ± 21.3 daily steps; Figure 2A). They also took more daily steps when provided access to a larger exercise area compared to smaller areas. More precisely, they took 16% more daily steps in the large area compared to the smaller (20 m²: 902 ± 28.4 daily steps; 40 m²: 929 ± 27.8 daily steps; 60 m²: 992 ± 27.6 daily steps; 80 m²: 1048 ± 31.0 daily steps; Figure 2B). Dataset 4. Other Behaviours When granted access to an exercise area, cows spent more than 2/3 of their time idle (Figure 3). In the trials where rumination behaviour was recorded (W19, F19 and S19), the cows were observed spending between 25 and 70% of their idle time ruminating. This behaviour was observed 10 to 50% of the time when a second hour of outing was provided. Cows dedicated between 5 and 30% of their time to locomotor behaviours and between 5 and 20% to exploration behaviours. When they were with other individuals in the same exercise area, the cows spent about 5% of their time engaging in social behaviours. Cows also spent about 15% of their time lying down in the indoor area (S20), while there were little (3% in S21) to no records of lying down in the outdoor exercise areas. Discussion While cows spent only 4% of their daily time in an exercise area, their locomotor behaviour increased by 53%. The use of exercise areas, such as the ones used in our study for one hour, led cows to take an extra 300 steps per day compared to when they were kept in tie-stalls for the whole day (700 daily steps, review by [ 8 ]). Animals perform locomotor behaviours when granted with the opportunity, indicating that it might represent an important behavioural need [ 11 , 12 ]. Therefore, providing an exercise area with sufficient space for locomotion, regardless of the other conditions of access, allows cows to increase their level of locomotor activity. When in the exercise area, cows were observed spending one third of their time engaging in locomotor behaviours, a result which corroborates the extra number of steps recorded by the accelerometers. In addition to walking, however, providing access to the exercise areas allowed cows to engage in a number of additional behaviours unavailable (or less available) to them in their home pens. Exploration, for one, is a behaviour for which animals generally show a strong motivation, indicating another possible need [ 5 , 10 , 13 ]. Similarly, cows exercised in dyads or triads and allowed to have visual and vocal contact with cows in other paddocks were observed performing social behaviours. Grazing (when grass was available in summer trials), grooming, and other maintenance behaviours were also observed, albeit to a lesser extent. Access to exercise areas therefore allowed cows to improve the quality of expression of a number of natural behaviours [ 7 ], beyond those tied directly to locomotion, indicating that the outdoor area offered more than just the fulfilment of the need of locomotion. Moreover, some of their behaviours also involve a degree of locomotion. Exploration, for example, is often carried out while walking, to better scan the environment. Certain social behaviours, such as following one another, also increase the proportion of animal’s locomotion. As such, these behaviours also contribute to increasing the animals' overall locomotion. With this in mind, it is important to note that our study likely under-estimated non-locomotive behaviours. In fact, some behaviours occurred only rarely, and/or lasted for a short duration of time; the scan sampling methodology employed to observe the cows in our study was thus unlikely to capture every occurrence of such behaviours (exploration for example). It is clear, however, that the cows outdoors engaged in a number of behaviours unavailable to them in stall. Moreover, the presence of such behaviours, despite a method that was not made to capture them, indicates a strong motivation to engage, and thus a potential behavioural need being met. Nevertheless, in exercise areas, we observed cows spent two-thirds of their time idle, despite the noted increase in both locomotory and other behaviours. This does not imply, however, that the cows did not benefit from or did not need exercise area access. The noted increase in locomotor activity, regardless of the conditions of access, indicates an important improvement in locomotor activity. The high level of inactivity may show that moving for the sake of moving is not necessarily what the cow wants. Indeed, in our study, the exercise areas were empty, i.e. without food, water, shelter or any additional objects. Only access to the exercise area itself was offered to the cows. This made it possible to observe the cows' behaviour in relation to access to the exercise area only, without any other interference. Having the opportunity to perform other behaviours could therefore further increase locomotor activity. Other studies had shown that when additional resources, such as food or shade, are provided in an outdoor area, cows will make use of them [ 12 , 14 , 15 , 16 ]. However, such studies do not distinguish between the benefits provided from the outing itself and from additional items added to the outing environment. Cow behaviour outdoors is moderated based on their needs for the resource in question. When provided with feed at a manger, cows will spend more time at the manger [ 12 ]. When provided with shade, cows will modify behaviour while outside to access it [ 16 ]. In both cases, cows show an increase in activity, but it is not possible to determine if this activity is a result of the outdoor access itself or surrounding variables. Factors that have been found to modulate cow motivation for outdoor access are the individual conditions of the access themselves. Features such as paddock size, outing duration, and handling during trips to and from outdoor areas can all impact how cows perceive and make use of the spaces provided to them [ 17 ]. In the present study, several controlled conditions for outings (indoor vs outdoor, size, duration) were tested, with the aim of maximizing the opportunity for movement and, more specifically, the locomotor behaviour, measured in number of steps, of dairy cows when provided access to an exercise area. One factor that was found to affect locomotor behaviour was the type of exercise area (indoor vs outdoor): we found that cows took around 20% more steps when they had access to an outdoor exercise area compared to an indoor one. There was a clear difference in the activities and level of stimulation offered when indoors vs outdoors. The indoor exercise area consisted of a loose-pen, without water or feed, which was already known to the animals from other contexts (for example, the loose-pen also serves as calving pen). In contrast, the outdoor exercise area, although it is of equivalent size, represented an entirely new and unpredictable environment which contains different stimuli than the cows may experience inside (e.g., rain showers, vehicles driving past the area), with associated behavioural responses specific to these stimuli (e.g., searching for shelter during rain), as well as a slightly more complex alleyway to traverse (curved rather than a straight line indoors). These factors, in addition to potential differences in handling from handlers equally responding to the two areas, may explain the slight discrepancy in step activity between the two areas. Our study also assessed the impact of the duration of the outing as well as the size of the exercise area. Increasing the size of the outdoor exercise area led cows to take a higher number of steps, but this increase was not proportional to the increase in area. Doubling the size of the exercise area (20 m² vs 40 m²) led to no increase in the number of daily steps. Comparatively, quadrupling the size of the exercise area (20 m² vs. 80 m²) only increased the number of daily steps by 16%. As such, our findings corroborate results from previous studies conducted in other housing environments (indoor free-stall housing [ 18 ]; pasture [ 19 ]) showing that the animals moved more in a bigger environment, but that this increase in locomotor behaviour was modest. In the smallest exercise area (20 m²), the available space per cow was 10 m². At this density, it is unlikely that other cows could represent a physical barrier if a cow decided to move freely [ 18 ], which allowed cows to meet the same needs as in the larger areas. Indeed, if the smallest area was large enough to meet baseline needs for locomotion, then no significant difference with space augmentation would be expected. Rather, the moderate difference observed could be accounted for by the extra space being of interest, but not necessary for the baseline locomotion needs of cattle. In terms of outing duration, our results showed a 9% increase in the number of daily steps with 2-hour outings compared with 1-hour outings. Previous studies found that cows given a 3-hour period of free access to an outdoor yard would choose to spend approximately 1 hour outdoors before choosing to return indoors, and thus were able to manage their outings to meet their needs [ 12 , 20 ]. Moreover, cows tend to return to the barn faster after a second hour of outdoor access compared to the first, emphasizing a higher motivation to return to their stalls when outing time is doubled [ 17 ]. Given these results, as well as the moderate increase in locomotive behaviour, one might assume that the second hour did not serve to meet any additional needs of locomotion for the cows. However, in contrast to step data, time budgets detailing the activities conducted during the first hour of outing and the second hour of outing appeared numerically similar, indicating that cows performed the same types of behaviours with similar occurrence during the second hour as they did during the first hour of outing. The fact that the same behaviours continue to be practiced in the second hour at the same rate supports the idea that behavioural needs remain, and cows are managing their activities to meet them. The lack of additional resources in the areas may serve to explain why step data did not also double. Though cows were motivated to go outside, and are clearly meeting needs while outdoors (evidenced by the fact that they continue the same activities), there are no other elements in the exercise areas that would serve to motivate the cows to move even more once there, as has been observed in other studies when such resources were provided [ 12 , 14 , 15 , 16 ]. Conclusion Our study found that the provision of daily exercise, regardless of conditions, has a major impact on the amount of locomotor behaviour performed by cows, leading them to take more daily steps while engaging in a greater range of other natural behaviours. Providing access to an exercise area for at least 1 hour allows cows to meet their behavioural needs for locomotion, and in our study, led cows to double their number of daily steps, with outing conditions such as the type of access, the amount of space provided, and the duration of the outing playing a role in both step activity and behaviours performed. Additional resources such as shelter structures, feed, or water could provide a more enriching experience to the cows brought to the exercise area, further enhancing the benefits of such a practice on cow locomotor activity and welfare. Materials and Methods Ethical Notes The use of animals in these projects and all procedures were reviewed and approved by the Animal Care Committee of McGill University and affiliated hospitals and research institutes for the experimentations conducted at McGill University (protocol #2016–7794; trials W19, S19, F19) and by the Animal Care Committee of the CRSAD (Deschambault Animal Science Research Centre) for the experimentations conducted at the CRSAD (protocol #1920-BL-387; trials W20, S20, S21). All aspects of this study met the standards established by the Canadian Council on Animal Care to ensure the humane and ethical use of animals in research, and all experiments followed relevant guidelines and regulations. Animals and Housing The present study was composed of six independent trials, each with different contexts (exercise area location and size, duration of outing, substrate, etc.), in order to test different exercise access conditions intended to increase the opportunity for movement provided to cows housed in a movement-restricted environment (i.e., tie-stall cows with no routine exercise access). While this precluded the possibility of analyzing all six trials together in one single model, it allowed for an in-depth comparison of specific conditions of the outings across trials. Outing conditions tested were as follows: 1) exercise vs non-exercise treatment, 2) indoor vs outdoor exercise area, and 3) a combination of outing duration (1h vs 2h) and size of exercise area (20, 40, 60 or 80 m²). In this section, the experimental setups for each of the six trials are briefly described. An overview of the different setups is shown in Table 1. In the W19 and S19 trials, 24 naïve, tie-stall-housed Holstein dairy cows from the McGill herd were selected and grouped in eight trios by days in milk and parity. Cows from each trio were then randomly assigned to one of three treatments: Exercise treatment cows (cows provided with outdoor access), non-Exercise treatment cows (cows remained at their stalls all day), and companion cows (cows for which data was not recorded but which accompanied Exercise treatment cows to avoid isolation and allow for social interaction). In the F19 trial, 30 cows from the McGill herd were blocked by days in milk and parity into five groups of six cows each -three Exercise treatment cows (provided with outdoor access) and three non-Exercise treatment cows (who remained at their stalls). One of the three cows in each trio had previous experience outdoors during an earlier trial. In the W20 and S20 trials, 27 and 30 cows respectively, were selected from the CRSAD herd and grouped into trios by days in milk and parity. Cows from each trio were randomly assigned into one of three treatments: Indoor Exercise (cows provided with indoor area access), Outdoor Exercise (cows provided with outdoor area access) and non-Exercise (cows remaining in their stalls for the duration of the treatment). Finally, in the S21 trial, 18 cows from the CRSAD herd were selected and grouped into trios by days in milk and parity, and randomly assigned to either: Outdoor-Biofilter1 Exercise treatment (cows providing with outdoor access, the ground of the exercise area covered by the biofilter1 (see biofilter details below)), Outdoor-Biofilter2 Exercise treatment (cows providing with outdoor access, the ground of the exercise area covered by the biofilter2) and non-Exercise treatment (cows remaining in the stall for the duration of the treatment). All the animals selected to participate in the different trials were healthy and non-lame lactating Holstein cows (evaluated based on a visual locomotion score adapted from [21]). During the study, all cows were housed in a mechanically-ventilated barn consiting of cubicle tie-stalls (stall area 2.7m 2 (W19, S19, F19) or 2.9m 2 (W20, S20, S21)) fitted with rubber mats and either wood chips (W19, S19, F19) or straw (W20, S20, S21). Cows had ad libitum access to water and were fed a total mixed ration (TMR) 2-4 times daily which was pushed up throughout the day to ensure feed accessibility. Milking was conducted twice daily at 9.5-14.5 h intervals. Procedures General process For the W19, S19, and F19 trials, during 8 weeks (W19 and S19) or 5 weeks (F19) cows went out every morning, five days per week (weekends excluded) for 1-2 h (treatment dependent) at 10AM, to an outside grassland exercise yard (1344 m²) adjacent to the barn (as demonstrated in [17], figure 1) in pairs in the winter and summer, and in trios in the fall. The trip distance from the stall to the exercise area measured approximately 56 m and was concrete inside and dirt outside. The outdoor exercise area consisted of paddocks delimited by electric fencing. Neither feed, water nor shelter were provided during outdoor access. In the W19 and S19 trials, the size of and time spent in the paddock changed weekly according to a Latin square design, such that each week each pair was assigned to one of the eight treatments: 1h-20m 2 , 1h-40m 2 , 1h-60 m 2 , 1h-80 m 2 , 2h-20m 2 , 2h-40m 2 , 2h-60 m 2 , or 2h-80 m 2 . In the F19 trial, cow trios were assigned to a different paddock each week, all paddocks measuring 117 m² (9 m x 13 m; 39 m² per cow). The ground cover was grass and snow in fall, wood chips and snow in winter, and grass during summer. For the S20, W20, and S21 trials, cows were provided with exercise access either 2 days per week for 1 hour per day (W20), or 5 days per week (weekends excluded) for 1.5 h per day (S20, S21) at the same time daily (morning or afternoon). In the W20 and S20 trials, cows were brought to either one of two outdoor exercise yards located adjacent to the barn (28.73 m² each (5.36 m x 5.36 m; one cow per area)), or to an indoor exercise area (29.60 m² (7.29 m x 4.06 m; one cow allowed at a time)), composed of three full walls and one half-height wall, allowing cows to see other cows in their stalls. The ground surface consisted of a concrete floor covered with a rubber mat and wood shavings. In the S21 trial, cows were brought to one of the two outdoor yards, consisting in this trial of different biofilter substrates composed of either wood chips and gravel, or wood chips, biochar, and sphagnum moss. These trials were designed as Latin square designs, with each cow experiencing one treatment (outdoor, indoor, or non-exercise in W20 and S20 or outdoor biofilters1, 2, or non-exercise in S21) for either 3 (W20) or 1 (S20, S21) week(s) before being rotated to the next treatment. During winter, snow covered the outdoor areas most of the time. The trip distances from the stall to the outdoor and indoor exercise area measured approximately 34 m and 19 m, respectively, and was concrete inside and dirt outside. Specific Notes by trial Winter 2020. The original setup of this trial was designed as a 3x3 Latin square in which the same treatment was applied for three consecutive weeks before switching to a new treatment. The trial was intended to last nine weeks but, due to the COVID-19 pandemic, the last 3-week period had to be dropped. Thus, the trial lasted six weeks, each cow receiving only two treatments for three weeks each. Summer 2021. The trial was initially designed as a 3x3 Latin square in which each cow would spend one week in each of the three treatments: Outdoor-Biofilter1 Exercise treatment, Outdoor-Biofilter2 Exercise treatment, and non-Exercise treatment. However, due to difficulties encountered in having some of the cows adapt to the outdoor exercise areas, the final dataset included six cows for three weeks, blocked in two groups of second and third lactation respectively. Measures Step Activity Locomotor behaviour (evaluated as the number of steps) was recorded continuously during each trial, beginning the first day of the trial, using a 3D pedometer (Peacock Technology Ltd, Stirling, Scotland, UK), attached to either rear leg of the cows. IceManager software (IceRobotics, Edinburgh, Scotland) was used to extract the raw data in 1-min intervals. The use of these pedometers to record step activity in tie-stall housed cows has previously been validated [22]. Other Behaviours Behavioural observations were conducting using either live observations (W19, S19, F19 and S21) or video recordings (S20) when the cows were in the exercise areas (indoor and outdoor). The frequency of the behaviours observed was recorded (Table 2). Observations began once the cows were calm and the barn staff no longer visible, and lasted either 30 min (F19), 1h (S20), 1.5h (S21), or 2h (W19 and S19). Data was collected by one to three observers, either by scan sampling every 30 s (S20 and S21) or every 4 min (W19, S19 and F19). These observations took place once a week for a maximum of one day (S20), two days (F19), three days (S21), six days (W19), or eight days (S19) of observations per cows. Statistical Analysis Data handling All statistical data handling and analyses were carried out using R (version 1.4.1103, R Core Team, 2021) and its specific packages. Four data sets were created. The first set included all trials described in the “Animals and Treatments” section and allowed us to investigate our primary objective, which was to know if exercise access, regardless of the conditions under which it was provided, had an influence on the number of daily steps. The second dataset included the trials conducted in the W20 and S20 and investigated the influence of the type of exercise area (indoor vs. outdoor) on the number of daily steps. The third dataset included trials conducted in the W19 and S19 and served to evaluate the influence of the duration of access to exercise and the size of the exercise area on the number of daily steps. Finally, the fourth dataset included information on the behaviours performed by the cows while in the exercise areas during all trials. For all analyses, week was coded to correspond to the week of treatment application at the cow level, and not the calendar week. The cleaning data method is available in Supplementary material. Dataset 1: Exercise vs non-Exercise treatments. For each trial, the lme4 package [23] was used to perform the following generalized linear mixed models ( GLMMs ) based on a Poisson distribution in order to evaluate the effect of exercise vs non-exercise within each trial. For each model, we considered the following factors: (1) treatment as a fixed effect: exercise treatments dependent on trial (outdoor exercise, indoor exercise, outdoor exercise biofilter 1 or 2) vs non-exercise; (2) week as a fixed effect: each week according to the trial; (3) block as a fixed effect (cows blocked by parity and days in milk); (4) Cow nested within block as a random effect; and, for trials W20, S20, S21, the order in which treatment was applied (as it changed weekly) as a random effect. For all these GLMMs, the independence between the residuals of the model and the predicted values, as well as the absence of over-dispersion of the residuals were assessed. In case of over-dispersion of the residuals, the model was adjusted by replacing the Poisson distribution by a negative binomial distribution using the glmer.nb function from the lme4 package. From each model, the Least Square means ( LSmeans ) and standard deviation ( SD ) were extracted using the emmeans function of the emmeans package [24]. The LSmeans were used to provide adjusted probabilities by accounting for the effect of each significant variable. We used these data to investigate the number of steps taken by cows with and without access to an exercise area. As the number of steps was recorded in the same way in all trials, a between-Group Mean difference ( MD ) meta-analysis was performed using the metacont function of the meta package [25]. Each trial could contribute more than one comparison (if different treatments were used in one trial), but the number of steps in non-Exercise treatment was always used as the control. As between-study heterogeneity was possible due to the trials’ different seasons and localizations, a random-effects model was used. The restricted maximum likelihood estimator [26] was used to calculate the heterogeneity variance τ². We used Knapp-Hartung adjustments [27] to calculate the confidence interval around the pooled effect. Dataset 2: Type of exercise area To determine if the type of exercise area, outdoor vs. indoor, had an impact on the daily number of steps, trials S20 and W20 containing these two outing conditions were grouped in the same dataset. We repeated a similar process to above, using the lme4 package [23] to perform a generalized linear mixed model based on a Poisson distribution, considering the following factors: (1) treatment as a fixed effect: outdoor exercise vs indoor exercise; (2) week as a fixed effect: each week according to the trial; (3) block as a fixed effect (cows blocked by parity and days in milk); (4) Cow nested within block as a random effect; and order in which treatment was applied (as it changed weekly) as a random effect. Independence between the residuals of the model and the predicted values, as well as absence of over-dispersion of the residuals, were assessed. Due to over-dispersion of the residuals, the model was adjusted by replacing the Poisson distribution by a negative binomial distribution using the glmer.nb function from the lme4 package. A type II Anova with Wald tests was carried out and LSmeans were used to perform post-hoc pairwise Bonferroni corrected comparisons. Results are presented as least square means with SEs unless otherwise stated. The significance threshold used was P < 0.05. Dataset 3: Duration of outing and size of the exercise area To determine if conditions of access to exercise such as the duration of the outings or the size of the outdoor exercise areas had an impact on the number of daily steps taken by the cows, the trials S19 and W19 containing these outing conditions were grouped in the same dataset. The lme4 package [23] was used to perform a generalized linear mixed model based on a Poisson distribution, considering the following factors: (1) duration as a fixed effect: 1 vs 2 hours; (2) week as a fixed effect: each week according to the trial; (3) block as a fixed effect (cows blocked by parity and days in milk); (4) Cow nested within block as a random effect; and order in which treatment was applied (as it changed weekly) as a random effect. Independence between the residuals of the model and the predicted values, as well as absence of over-dispersion of the residuals were assessed. In case of over-dispersion of the residuals, the model was adjusted by replacing the Poisson distribution by a negative binomial distribution using the glmer.nb function from the lme4 package. A type II Anova with Wald tests was carried out and LSmeans were used to perform post-hoc pairwise Bonferroni corrected comparisons. Results are presented as least square means with SEs unless otherwise stated. The significance threshold used was P < 0.05. Dataset 4: Other Behaviours Descriptive statistics were used to obtain the percentage of time allocated to each of the behavioural activities by each animal within each trial. Declarations Funding Funding for conducting research was provided by the Natural Sciences and Engineering Research Council of Canada, Novalait, Dairy Farmers of Canada, and Valacta as a part of the Industrial Research Chair in the Sustainable Life of Dairy Cattle, as well as Agriculture and AgriFood Canada and Dairy Farmers of Canada through AgriScience Clusters program, Growing Forward III. Competing Interests Statement The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. Author Contribution M. Cellier: conceptualization, data curation - lead, format analysis - lead, methodology, writing original draft – lead, writing review & editing ;E. Shepley: conceptualization, methodology, writing – review and editing ; N. Aigueperse: conceptualization, methodology, writing – review and editing ;M.V. Robichaud: conceptualization, funding acquisition, methodology, writing – review and editing ;E. Vasseur: conceptualization, funding acquisition, investigation, methodology, project administration, resources, supervision, writing original draft – lead, writing – review and editing. Acknowledgement The authors would like to acknowledge the following individuals who conducted the animal trials: McGill University M. Sc. graduates Jordan Tonooka, Catherine Lussier, Amir Nejati, Rachel Chiasson, and Université Laval research assistant Béatrice Dupont-Fortin. We extend our gratitude to the staff at the McGill Macdonald Campus Dairy Unit and Dairy Facility of the Centre de Recherche en Sciences Animales de Deschambault (CRSAD), as well as Véronique Boyer (McGill University) and Janie Lévesque (CRSAD), who assisted in animal trial planning and data collection. We would like to thank Rachel VanVliet (McGill University) for the editorial suggestion and corrections, and Véronique Boyer (McGill University) for the proofreading and corrections of the article; Tania Wolfe (McGill University), Roger Cue (McGill University) and Sébastien Buczinski (University of Montréal) for their help with the statistical reflection. We would like to acknowledge the contribution of environmental engineers Stéphane Godbout (Institut de recherche et de développement en agro-environnement IRDA), Sébastien Fournel (Université Laval) and Pierre Ruel (CRSAD) in funding acquisition, experimental project planning and implementation at CRSAD. Data Availability The raw data supporting the conclusions of this article are available here : https://github.com/WELL-E-chair/data-movement-behavior?tab=readme-ov-file References Haskell, M. J., Brotherstone, S., Lawrence, A. B. & White, I. M. S. Characterization of the Dairy Farm Environment in Great Britain and the Effect of the Farm Environment on Cow Life Span. J Dairy Sci. 90(11) , 5316-5323 (2007). Beaver, A., Ritter, C. & von Keyserlingk, M. A. G. The dairy cattle housing dilemma: Natural behavior versus animal care. Vet Clin North Am Food Anim Pract . 35(1) , 11-27 (2019). Charlton, G. L. & Rutter, S. M. The behaviour of housed dairy cattle with and without pasture access: A review. Appl Anim Behav Sci. 192 , 2-9 (2017). Kresnye, K. C., Chung, C., Martin, C. F. & Shih, P. C. Survey on the Past Decade of technology in Animal Enrichment: A Scoping Review. Animals . 12(14) , 1792. 10.3390/ani12141792 (2022). Boissy, A. et al . Assessment of positive emotions in animals to improve their welfare. Physiol Behav. 92(3) , 375-397 (2007). Fraser, D. Understanding animal welfare. Acta Vet Scand. 50 , S1. 10.1186/1751-0147-50-S1-S1 (2008). Krohn, C. C. 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(2) , 1-16 (1994). Shepley, E., Lensink, J. & Vasseur, E. Cow in Motion: A review of the impact of housing systems on movement opportunity of dairy cows and implications of locomotor activity. Appl Anim Behav Sci. 230 , 105026. 10.1016/j.applanim.2020.105026 (2020). Newberry, R. C. Environmental enrichment: Increasing the biological relevance of captive environments. Appl Anim Behav Sci. 44 , 229-243 (1995). Loberg, J., Telezhenko, E., Bergsten, C. & Lidfors, L. Behaviour and claw health in tied dairy cows with varying access to exercise in an outdoor paddock. Appl Anim Behav Sci. 89(1-2) , 1-16 (2004). Veissier, I., Andanson, S., Dubroeucq, H. & Pomiès, D. The motivation of cows to walk as thwarted by tethering. J Anim Sci. 86(10) , 2723-2729 (2008). Shepley, E., Bergeron, R., Bécotte, F. & Vasseur, E. Dairy cow preference for outdoor access during winter under Eastern Canada climatic conditions. Can J Anim Sci. 97(1) , 1-5. 10.1139/CJAS-2016-0028 (2016). Mattiello, S., Battini, M., De Rosa, G., Napolitano, F. & Dwyer, C. How can we assess positive welfare in ruminants? Animals . 9(10) , 758. 10.3390/ani100758 (2019). Zeeb, K. Locomotion and space structure in six cattle units. In Farm Animal Housing and Welfare (eds. Baxter, S. H., Baxter, M. R. & MacCormack, J. A. D.). 129-136 (Springer Science & Business Media, 1983). Gustafson, G. M. Effects of daily exercise on the health of tied dairy cows. Prev Vet Med. 17(3-4) , 209-223 (1993). Palacio, S., Bergeron, R., Lachance, S. & Vasseur, E. The effects of providing portable shade at pasture on dairy cow behavior and physiology. J Dairy Sci. 98(9) , 6085-6093 (2015). Aigueperse, N., Boyer, V. & Vasseur, E. How outing conditions relate to the motivation of movement-restricted cattle to access an outdoor exercise yard. Behav Processes. 213 , 104957. 10.1016/j.beproc.2023.104957 (2023). Telezhenko, E., von Keyserlingk, M. A. G., Talebi, A. & Weary, D. M. Effect of pen size, group size, and stocking density on activity in freestall-housed dairy cows. J Dairy Sci. 95(6) , 3064-3069 (2012). Henkin, Z. et al . The spatial distribution and activity of cattle in response to plot size. J Anim Feed Sci. 16 (Suppl. 2) , 399-404 (2007). Shepley, E., Bergeron, R. & Vasseur, E. Daytime summer access to pasture vs free-stall barn in dairy cows with year-long outdoor experience: A case study. Appl Anim Behav Sci. 192 , 10-14 (2017). Flower, F. C. & Weary, D. M. Gait assessment in dairy cattle. Animal . 3(1) , 87-95 (2009). Shepley, E., Berthelot, M. & Vasseur, E. Validation of the Ability of a 3D Pedometer to Accurately Determine the Number of Steps Taken by dairy Cows When Housed in Tie-Stalls. Agriculture . 7(7) , 53. 10.3390/agriculture7070053 (2017). Bates, D., Maechler, M., Bolker, B. & Walker, S. Fitting Linear Mixed-Effects Models Using lme4. J Stat Softw. 67(1) , 1-48. 10.18637/jss.v067.i01 (2015). Lenth, R., Singmann, H., Love, J., Buerkner, P. & Herve, M. Emmeans: Estimated marginal means, aka least-squares means . R Package Version 1 (2018). Balduzzi, S., Rücker, G. & Schwarzer, G. How to perform a meta-analysis with R: a practical tutorial. Evid Based Ment Health. 22(4) , 153-160 (2019). Viechtbauer, W. Bias and Efficiency of Meta-Analytic Variance Estimators in the Random-Effects Model. J Educ Behav Stat. 30(3) , 261-295 (2005) Knapp, G. & Hartung, J. Improved tests for a random effects meta-regression with a single covariate. Stat Med. 22(17) , 2693-2710 (2003). Tables Table 1. Summary of the characteristics of trials Trial Winter 2019 Summer 2019 Fall 2019 Winter 2020 Summer 2020 Summer 2021 Duration of the trial (weeks) 8 8 5 6 3 3 Number of cows 24 24 30 27 18 18 Number of blocks 8 8 5 9 6 6 Parity per block 2.0 ± 1.46 2.5 ± 1.68 1.9 ± 0.79 2.2 ± 0.93 2.0 ± 0.84 2.7 ± 1.00 DIM block 149 ± 63.5 112 ± 63.5 163 ± 66.4 117 ± 65.8 187 ± 99.5 173 ± 102 Treatments Duration x Size Duration x Size Outdoor Outdoor Indoor Outdoor-Biofilter1 Outdoor-Biofilter2 Control Stall Stall Stall Stall Stall Stall Outings/Week 5 5 5 2 5 5 Duration per outing (hours) 1 or 2* 1 or 2* 1 1 1.5 1.5 Size of the area (m²) 20, 40, 60, 80* 20, 40, 60, 80* 117 28.73(outdoor) 29.60(indoor) 28.73 Localization McGill McGill McGill CRSAD CRSAD CRSAD * depending on the treatment (combination between duration and size) applied Table 2. Detailed ethogram used to assess cow activity during the different trials Category Behaviour Description Trial Idle Idle Body relaxed, immobile with head down or in the extension of her body. Cow can do other activities such as ruminating or standing alert, while idling F19, S19, W19, S20, S21 Rumination Rhythmic/repeated chewing motions of partly digested food not associated with head down in feed F19, S19, W19 Vigilance Vigilance Body immobile with head up. Cow standing alert, with the ears facing forward F19, S19, W19 Lying Lying Cows were considered lying when the flank of the animal was in contact with a surface F19, W19, S20, S21 Exploration Exploring Cow is sniffing or licking the ground, feed, stall structure, or any other object of her surroundings. Cow is not walking during exploration. F19, S19, W19, S20, S21 Locomotion Walking Movement forward of the feet, with one- or two-feet lift. Walking is considered when the cow takes at least 3 steps. Cow can ruminate and/or explore the ground or any object on her surroundings while walking F19, S19, W19, S20, S21 Running Fast movement forward of the parallel feet which touch and lift simultaneously with a phase of suspension in air S20, S21 Jumping Four legs lifted from the ground S20 Social Social interactions Any positive or negative behaviour involving counterparts (e.g., mutual play headbutt, allo-grooming...) F19, S19, W19 Eating Eating Cow is actively grabbing and chewing feed with head down, or any chewing movement after gathering a mouthful of feed. Cow can walk while eating F19, S19, W19 Maintenance Self-grooming Tongue in contact with own skin or fur/scratches with hind legs or any sort of structure to scratch against F19, S19, W19, S20, S21 Defecation Elimination of feces from the body S19, S20, S21 Urination Elimination of urine from the body S19, S20, S21 Other Others Any other activity not described here F19, S19, W19, S20, S21 W19: Winter 2019 trial; S19: Summer 2019 trial; F19: Fall 2019 trial; W20: Winter 2020 trial; S20: Summer 2020 trial; S21: Summer 2021 trial Table 3: Descriptive statistics for each trial, after the data cleaning steps Winter 2019 Summer 2019 Fall 2019 Winter 2020 Summer 2020 Summer 2021 N of cows Non-Exercise 7 8 15 18 18 5 Exercise 7 7 15 - - - Treatment A - - - 19 18 4 Treatment B - - - 18 17 5 LSmean number of daily steps ± SD Non-Exercise 543 ± 155 616 ± 671 442 ± 228 547 ± 197 725 ± 207 528 ± 207 Exercise 777 ± 264 1233 ± 482 763 ± 393 - - - Treatment A - - - 835 ± 314 1239 ± 354 770 ± 279 Treatment B - - - 765 ± 282 954 ± 266 968 ± 380 For Winter 2020 and Summer 2020, treatment A and B corresponded to Outdoor Exercise and Indoor Exercise, respectively. For Summer 2021, treatment A and B corresponded to Outdoor-Biofilter1 Exercise and Outdoor-Biofilter2 Exercise, respectively. Note that for Winter 2020, Summer 2020 and Summer 2021, the cows went through each treatment (Exercise vs. non-Exercise) consecutively, while in Winter 2019, Summer 2019 and Fall 2019, Exercise and non-Exercise cows were different individuals. Additional Declarations No competing interests reported. Supplementary Files Supplementarymaterial.docx Cite Share Download PDF Status: Published Journal Publication published 18 Feb, 2025 Read the published version in Scientific Reports → Version 1 posted Editorial decision: Revision requested 13 Oct, 2024 Reviews received at journal 11 Oct, 2024 Reviewers agreed at journal 30 Sep, 2024 Reviews received at journal 25 Sep, 2024 Reviewers agreed at journal 23 Sep, 2024 Reviewers invited by journal 21 Sep, 2024 Editor assigned by journal 18 Sep, 2024 Editor invited by journal 22 Aug, 2024 Submission checks completed at journal 21 Aug, 2024 First submitted to journal 21 Aug, 2024 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-4953052","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":356194988,"identity":"0651aea3-f5e4-4a9d-93ca-0f9453e4dec0","order_by":0,"name":"Marjorie Cellier","email":"","orcid":"","institution":"McGill University","correspondingAuthor":false,"prefix":"","firstName":"Marjorie","middleName":"","lastName":"Cellier","suffix":""},{"id":356194989,"identity":"03077904-07c5-4301-825d-03e85a12ed4b","order_by":1,"name":"Elise Shepley","email":"","orcid":"","institution":"University of Minnesota","correspondingAuthor":false,"prefix":"","firstName":"Elise","middleName":"","lastName":"Shepley","suffix":""},{"id":356194990,"identity":"325510e6-e40a-4604-b282-ab23193b0aa4","order_by":2,"name":"Nadège Aigueperse","email":"","orcid":"","institution":"Université Clermont Auvergne, INRAE, VetAgro Sup, UMR Herbivores","correspondingAuthor":false,"prefix":"","firstName":"Nadège","middleName":"","lastName":"Aigueperse","suffix":""},{"id":356194991,"identity":"36848ef2-2371-4d66-84fb-52e2e7ee3e11","order_by":3,"name":"Marianne Villettaz Robichaud","email":"","orcid":"","institution":"Université de Montréal","correspondingAuthor":false,"prefix":"","firstName":"Marianne","middleName":"Villettaz","lastName":"Robichaud","suffix":""},{"id":356194992,"identity":"fe39be2b-8895-409d-a396-f176942fb766","order_by":4,"name":"Elsa Vasseur","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA9ElEQVRIiWNgGAWjYDACdmTOByBmY2duwK+FGYnNOAOkhZmRBC3MPGCSgBZ+ZvaHHxjb7Ozl23vMPtvm2Nn1gbT8qMGtRbKZx1iCsS05ccOZM8azc7clJ7cBtTD2HMOtxeAwD4MEwxnmBAOJHGPm3G0HkkF+YWZgw63F/jD74x8MZ+rt5ee/MWa2hGv5h8cWZgYzCYaKw4wNN3iMmRm3HbADa2Fsw61F4jCPmUVCxXGgX9KKGXu3JSeAtBzs7cOthb+9/fGNDwbVwBA7vJnh5zZQ0DUffPDjG24tYJCAxE5sABIHCGhABfYkqR4Fo2AUjIIRAQCR50XvQLjvrQAAAABJRU5ErkJggg==","orcid":"","institution":"McGill University","correspondingAuthor":true,"prefix":"","firstName":"Elsa","middleName":"","lastName":"Vasseur","suffix":""}],"badges":[],"createdAt":"2024-08-21 16:37:45","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4953052/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4953052/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1038/s41598-025-89891-4","type":"published","date":"2025-02-18T15:56:51+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":64896102,"identity":"5dce3827-e725-4cf4-a37b-3b062ab9cd56","added_by":"auto","created_at":"2024-09-20 07:05:18","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1517199,"visible":true,"origin":"","legend":"\u003cp\u003eForest plot of nine modalities (distributed within six trials) associated with number of daily steps. An increase of the number of daily steps compared to the control (non-Exercise treatment) is represented by a deviation of the square (difference in the means) to the right. The length of the horizontal line represents the 95% confidence interval, and the diamond represents the summarized effect. W20_Indoor: Winter 2020 trial with an indoor exercise area, W20_Outdoor: Winter 2020 trial with an outdoor exercise area; S20_Indoor: Summer 2020 trial with an indoor exercise area; S20_Outdoor: Summer 2020 trial with an outdoor exercise area; W19: Winter 2019 trial; S19: Summer 2019 trial; F19: Fall 2019 trial; S21_Biofilter1: Summer 2021 trial with the ground covered by the biofilter 1 (composed of either wood chips and gravel); S21_Biofilter: Summer 2021 trial with the ground cover by the biofilter 2 (composed of either wood chips, biochar and sphagnum moss); MD: Group Mean difference; CI: Confidence Interval\u003c/p\u003e","description":"","filename":"Figure1highres.png","url":"https://assets-eu.researchsquare.com/files/rs-4953052/v1/aeec39e6e7eecbabb6e691ab.png"},{"id":64895431,"identity":"dd33e90f-8b46-4945-82e9-a001ac57a387","added_by":"auto","created_at":"2024-09-20 06:57:18","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":433279,"visible":true,"origin":"","legend":"\u003cp\u003eNumber of daily steps (LSmeans ± SE) taken when cows had access to an outdoor exercise area for 1-hour or 2-hours (a) and when the size of the exercise area varied from 20 m² to 80 m² (b). Significant differences indicated by the model (P \u0026lt; 0.05) are represented by different letters.\u003c/p\u003e","description":"","filename":"Figure2highres.png","url":"https://assets-eu.researchsquare.com/files/rs-4953052/v1/9886a0ba718bb6b0e2c19333.png"},{"id":64895430,"identity":"92061bec-d629-48e4-97f0-41644956f477","added_by":"auto","created_at":"2024-09-20 06:57:18","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1844805,"visible":true,"origin":"","legend":"\u003cp\u003eTime budgets of different behaviours’ categories (definitions in Table 2) that cows displayed during the different trial: A) the first hour of outing during Winter 2019 trial, B) the second hour of outing during Winter 2019 trial, C) the first hour of outing during Summer 2019 trial, D) the second hour of outing during Summer 2019 trial, E) Summer 2020 trial with an indoor exercise area, F) Summer 2020 trial with an outdoor exercise area, G) Fall 2019 trial, and H) Summer 2021 trial. Social, eating, and vigilance behaviours were not recorded in Summer 2020 and Summer 2021 trials, whereas lying behaviour was not recorded in Summer 2019 trial.\u003c/p\u003e","description":"","filename":"Figure3highres.png","url":"https://assets-eu.researchsquare.com/files/rs-4953052/v1/dcbaa1b32ecf81a673aab67b.png"},{"id":77052443,"identity":"77883f29-2fd8-4d6a-b9cb-1a666d8fab5b","added_by":"auto","created_at":"2025-02-24 16:00:24","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":4923503,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4953052/v1/798275e3-a7a2-4fe2-9f87-b8b496f2f7f7.pdf"},{"id":64896827,"identity":"1fc156b4-ef99-4e4b-ba3f-1dd909c369bc","added_by":"auto","created_at":"2024-09-20 07:13:18","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":22003,"visible":true,"origin":"","legend":"","description":"","filename":"Supplementarymaterial.docx","url":"https://assets-eu.researchsquare.com/files/rs-4953052/v1/3ac0dd5f492d80169d53ac73.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Enhancing movement opportunity and fulfilling behavioural needs for movement- restricted cattle by providing different conditions of access to exercise","fulltext":[{"header":"Introduction","content":"\u003cp\u003eIn recent decades, the animal production, including dairy industry, has undergone an intensification characterized by changes in animal housing, notably an increase in indoor housing [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e] with limited to non-existent outdoor access. Arguments put forward for the use of this type of system are, among others, individualized care, feed rationing, and reduced risk of agonistic behaviours [\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]. This change has been met with public criticism by a society increasingly concerned about farm animal welfare [\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e]. What is often missing from these debates, however, is the perspective of the cow. As we cannot ask her directly, other means must be utilized in order to shed light on this important stakeholder within this debate. How does she use the space provided to her and what utility does it have? Can other elements be introduced to mitigate the negative effects of certain housing systems, or bring additional positive ones? If so, how does she perceive and make use of these elements? We often rely on what is known as environmental enrichment to add to the captive environment. This is a broad term that covers any change or addition to the living environment or routine of an individual with the goal of providing more opportunities for choice, agency, and natural behaviours. The effectiveness and utility of any enrichment can then only be evaluated through the perception of the individual and will thus depend on how the cow perceives and uses what is provided to her [\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e]. Moreover, evidence suggests that the greatest positive benefit to animals comes from additions that fulfil existing behavioural needs [\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e], further emphasizing our need to understand the cow\u0026rsquo;s existing needs and perception of her environment.\u003c/p\u003e \u003cp\u003eOne such need impacted by the increase of indoor housing is the need for movement and particularly the need for locomotion. The provision of exercise area access by increasing movement opportunity, is a potential solution to meet a cow\u0026rsquo;s needs for movement opportunity, as well as a variety of other natural behaviours not available to her in her home restricted environment [\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e]. For example, cows provided with regular access to an exercise area get more opportunities to engage in social interactions, to explore their environment, and to groom hard-to-reach body parts such as their hindquarters [\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e]. Movement opportunity can be defined as the level and ease of movement provided to an animal in its housing environment [\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e]. This includes the number of steps achieved by the cow [\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e], freedom of movement, and locomotor behaviours [\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e]. The average number of daily steps taken by cows, differs across housing systems: it ranges from 700 steps/day in tie-stall housing to 4000 steps/day on pasture, with free-stall housing sitting in-between at 1500 steps/day (reviewed by [\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e]). These quantitative data support the idea that systems offering more opportunity of movement can positively influence the level of locomotor behaviour performed by a cow (i.e. the cow will make use of the additional space provided). However, new scientific knowledge is necessary relating to the benefits of providing exercise access and the conditions of exercise access, like outdoor or indoor access, required to enhance such benefits. In particular, we need to know if and how cows actually use the exercise area as a movement opportunity to meet their locomotion needs by increasing their locomotor activities.\u003c/p\u003e \u003cp\u003eMoreover, cow motivation to engage in locomotor behaviours such as walking, trotting, or running has been studied both in individuals kept in tie-stalls given access to an exercise area [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e]and in those usually housed on pasture (summer) or free-stalls (winter) kept tied for the duration of the experiment [\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e]. Though being tied reduced locomotor behaviour for these cows, when additionally provided with daily access to an exercise area (e.g., indoor exercise area, outdoor paddock, or pasture), they showed an increase in locomotor behaviours [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e]. However, it is not known to what extent access to an exercise area for cows continuously living in a movement-restrictive environment (i.e., cows that are not used to moving) affects their level of locomotor behaviour, most notably in terms of number of steps. It is further unknown how the conditions of access to exercise, such as the duration of access or the size of the exercise area, can influence these outcomes.\u003c/p\u003e \u003cp\u003eOur study aimed to 1) quantify how providing movement opportunity outside of the stall impacts the locomotor behaviour of cows housed in a movement-restricted environment; 2) evaluate which conditions of access to exercise optimize locomotor behaviour: indoor vs outdoor, duration of outing, size of the exercise area; and 3) investigate which other behaviours cows perform when granted access to an exercise area.\u003c/p\u003e \u003cp\u003eWe hypothesized that regardless of the specific conditions, providing access to exercise would lead cows to take more steps than when confined to their stalls. We also hypothesized that cows would take a similar number of steps regardless of the conditions of access to the exercise area and that it is the overall experience provided to the cow outdoors (companionship, etc.) rather than the exercise area per se that would have a greater influence on the increase in locomotor behaviour in cows provided with regular exercise access. Finally, we hypothesized that cows would engage in a variety of behaviours (as opposed to solely engaging in locomotor behaviours) when provided with opportunity for movement outside of their stalls.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cem\u003eDataset 1. Exercise vs non-Exercise treatments: Meta-analysis\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eDescriptive statistics for each trial are shown in Table 3.\u003c/p\u003e\n\u003cp\u003eThe forest plot of the meta-analysis is presented in Figure 1. The meta-analysis of all comparisons between Exercise and non-Exercise treatments resulted in a between-study heterogeneity variance estimated at \u0026tau;\u0026sup2; = 2686 (95% Confident Interval (\u003cstrong\u003eCI\u003c/strong\u003e): 0-26859), with an I\u0026sup2; value of 9.5% (95% CI: 0% \u0026ndash; 68.1%), meaning that less than 10% of the variation is due to between-study heterogeneity. The nonsignificant Q score suggests that the studies represent samples from a common population and that the coalescing\u0026nbsp;of\u0026nbsp;their data was justified (Q = 8.84, degrees of freedom (\u003cstrong\u003edf\u003c/strong\u003e) = 8, P = 0.356).\u0026nbsp;The prediction interval ranged from g = 151 to 457. The number of daily steps was significantly higher when cows had access to an exercise area, with an average increase of 304 steps per day (95% CI: 215-393; \u003cem\u003eP\u003c/em\u003e\u003cem\u003e\u0026nbsp;\u0026lt;\u003c/em\u003e 0.001), equivalent to 53% more steps (923 \u0026plusmn; 65\u0026nbsp;daily steps) compared to the 578 (\u0026plusmn; 32\u0026nbsp;daily steps) mean daily steps taken in the stall.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eDataset 2.\u003c/em\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003cem\u003eType of exercise area\u003c/em\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe number of daily steps differed depending on whether cows had access to an indoor or outdoor exercise area. Cows took 167, or around 20%, more daily steps in an outdoor exercise area (987 \u0026plusmn; 50.3 daily steps) than in an indoor one (820 \u0026plusmn; 42.4 daily steps; \u0026chi;\u0026sup2; = 44.980, df = 1, P \u0026lt; 0.001).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eDataset 3. Duration of outing and size of the exercise area\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe number of daily steps was influenced by the duration of the outing (\u0026chi;\u0026sup2; = 10.090, df = 1, P = 0.002) and by the size of the exercise area (\u0026chi;\u0026sup2; = 18.317, df = 3, P \u0026lt; 0.001). Cows took 84, or around 9%, more daily steps when staying in the exercise area for 2 hours (1009 \u0026plusmn; 22.8 daily steps) compared with 1 hour (925 \u0026plusmn; 21.3 daily steps; Figure 2A). They also took more daily steps when provided access to a larger exercise area compared to smaller areas. More precisely, they took 16% more daily steps in the large area compared to the smaller (20 m\u0026sup2;: 902 \u0026plusmn; 28.4 daily steps; 40 m\u0026sup2;: 929 \u0026plusmn; 27.8 daily steps; 60 m\u0026sup2;: 992 \u0026plusmn; 27.6 daily steps; 80 m\u0026sup2;: 1048 \u0026plusmn; 31.0 daily steps; Figure 2B).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eDataset 4. Other Behaviours\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eWhen granted access to an exercise area, cows spent more than 2/3 of their time idle (Figure 3). In the trials where rumination behaviour was recorded (W19, F19 and S19), the cows were observed spending between 25 and 70% of their idle time ruminating. This behaviour was observed 10 to 50% of the time when a second hour of outing was provided. Cows dedicated between 5 and 30% of their time to locomotor behaviours and between 5 and 20% to exploration behaviours. When they were with other individuals in the same exercise area, the cows spent about 5% of their time engaging in social behaviours. Cows also spent about 15% of their time lying down in the indoor area (S20), while there were little (3% in S21) to no records of lying down in the outdoor exercise areas.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eWhile cows spent only 4% of their daily time in an exercise area, their locomotor behaviour increased by 53%. The use of exercise areas, such as the ones used in our study for one hour, led cows to take an extra 300 steps per day compared to when they were kept in tie-stalls for the whole day (700 daily steps, review by [\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e]). Animals perform locomotor behaviours when granted with the opportunity, indicating that it might represent an important behavioural need [\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]. Therefore, providing an exercise area with sufficient space for locomotion, regardless of the other conditions of access, allows cows to increase their level of locomotor activity.\u003c/p\u003e \u003cp\u003eWhen in the exercise area, cows were observed spending one third of their time engaging in locomotor behaviours, a result which corroborates the extra number of steps recorded by the accelerometers. In addition to walking, however, providing access to the exercise areas allowed cows to engage in a number of additional behaviours unavailable (or less available) to them in their home pens. Exploration, for one, is a behaviour for which animals generally show a strong motivation, indicating another possible need [\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e, \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e, \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e]. Similarly, cows exercised in dyads or triads and allowed to have visual and vocal contact with cows in other paddocks were observed performing social behaviours. Grazing (when grass was available in summer trials), grooming, and other maintenance behaviours were also observed, albeit to a lesser extent. Access to exercise areas therefore allowed cows to improve the quality of expression of a number of natural behaviours [\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e], beyond those tied directly to locomotion, indicating that the outdoor area offered more than just the fulfilment of the need of locomotion. Moreover, some of their behaviours also involve a degree of locomotion. Exploration, for example, is often carried out while walking, to better scan the environment. Certain social behaviours, such as following one another, also increase the proportion of animal\u0026rsquo;s locomotion. As such, these behaviours also contribute to increasing the animals' overall locomotion. With this in mind, it is important to note that our study likely under-estimated non-locomotive behaviours. In fact, some behaviours occurred only rarely, and/or lasted for a short duration of time; the scan sampling methodology employed to observe the cows in our study was thus unlikely to capture every occurrence of such behaviours (exploration for example). It is clear, however, that the cows outdoors engaged in a number of behaviours unavailable to them in stall. Moreover, the presence of such behaviours, despite a method that was not made to capture them, indicates a strong motivation to engage, and thus a potential behavioural need being met.\u003c/p\u003e \u003cp\u003eNevertheless, in exercise areas, we observed cows spent two-thirds of their time idle, despite the noted increase in both locomotory and other behaviours. This does not imply, however, that the cows did not benefit from or did not need exercise area access. The noted increase in locomotor activity, regardless of the conditions of access, indicates an important improvement in locomotor activity. The high level of inactivity may show that moving for the sake of moving is not necessarily what the cow wants. Indeed, in our study, the exercise areas were empty, i.e. without food, water, shelter or any additional objects. Only access to the exercise area itself was offered to the cows. This made it possible to observe the cows' behaviour in relation to access to the exercise area only, without any other interference. Having the opportunity to perform other behaviours could therefore further increase locomotor activity. Other studies had shown that when additional resources, such as food or shade, are provided in an outdoor area, cows will make use of them [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e]. However, such studies do not distinguish between the benefits provided from the outing itself and from additional items added to the outing environment. Cow behaviour outdoors is moderated based on their needs for the resource in question. When provided with feed at a manger, cows will spend more time at the manger [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]. When provided with shade, cows will modify behaviour while outside to access it [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e]. In both cases, cows show an increase in activity, but it is not possible to determine if this activity is a result of the outdoor access itself or surrounding variables.\u003c/p\u003e \u003cp\u003eFactors that have been found to modulate cow motivation for outdoor access are the individual conditions of the access themselves. Features such as paddock size, outing duration, and handling during trips to and from outdoor areas can all impact how cows perceive and make use of the spaces provided to them [\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e]. In the present study, several controlled conditions for outings (indoor vs outdoor, size, duration) were tested, with the aim of maximizing the opportunity for movement and, more specifically, the locomotor behaviour, measured in number of steps, of dairy cows when provided access to an exercise area. One factor that was found to affect locomotor behaviour was the type of exercise area (indoor vs outdoor): we found that cows took around 20% more steps when they had access to an outdoor exercise area compared to an indoor one. There was a clear difference in the activities and level of stimulation offered when indoors vs outdoors. The indoor exercise area consisted of a loose-pen, without water or feed, which was already known to the animals from other contexts (for example, the loose-pen also serves as calving pen). In contrast, the outdoor exercise area, although it is of equivalent size, represented an entirely new and unpredictable environment which contains different stimuli than the cows may experience inside (e.g., rain showers, vehicles driving past the area), with associated behavioural responses specific to these stimuli (e.g., searching for shelter during rain), as well as a slightly more complex alleyway to traverse (curved rather than a straight line indoors). These factors, in addition to potential differences in handling from handlers equally responding to the two areas, may explain the slight discrepancy in step activity between the two areas.\u003c/p\u003e \u003cp\u003eOur study also assessed the impact of the duration of the outing as well as the size of the exercise area. Increasing the size of the outdoor exercise area led cows to take a higher number of steps, but this increase was not proportional to the increase in area. Doubling the size of the exercise area (20 m\u0026sup2; vs 40 m\u0026sup2;) led to no increase in the number of daily steps. Comparatively, quadrupling the size of the exercise area (20 m\u0026sup2; vs. 80 m\u0026sup2;) only increased the number of daily steps by 16%. As such, our findings corroborate results from previous studies conducted in other housing environments (indoor free-stall housing [\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e]; pasture [\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e]) showing that the animals moved more in a bigger environment, but that this increase in locomotor behaviour was modest. In the smallest exercise area (20 m\u0026sup2;), the available space per cow was 10 m\u0026sup2;. At this density, it is unlikely that other cows could represent a physical barrier if a cow decided to move freely [\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e], which allowed cows to meet the same needs as in the larger areas. Indeed, if the smallest area was large enough to meet baseline needs for locomotion, then no significant difference with space augmentation would be expected. Rather, the moderate difference observed could be accounted for by the extra space being of interest, but not necessary for the baseline locomotion needs of cattle.\u003c/p\u003e \u003cp\u003eIn terms of outing duration, our results showed a 9% increase in the number of daily steps with 2-hour outings compared with 1-hour outings. Previous studies found that cows given a 3-hour period of free access to an outdoor yard would choose to spend approximately 1 hour outdoors before choosing to return indoors, and thus were able to manage their outings to meet their needs [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e]. Moreover, cows tend to return to the barn faster after a second hour of outdoor access compared to the first, emphasizing a higher motivation to return to their stalls when outing time is doubled [\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e]. Given these results, as well as the moderate increase in locomotive behaviour, one might assume that the second hour did not serve to meet any additional needs of locomotion for the cows. However, in contrast to step data, time budgets detailing the activities conducted during the first hour of outing and the second hour of outing appeared numerically similar, indicating that cows performed the same types of behaviours with similar occurrence during the second hour as they did during the first hour of outing. The fact that the same behaviours continue to be practiced in the second hour at the same rate supports the idea that behavioural needs remain, and cows are managing their activities to meet them. The lack of additional resources in the areas may serve to explain why step data did not also double. Though cows were motivated to go outside, and are clearly meeting needs while outdoors (evidenced by the fact that they continue the same activities), there are no other elements in the exercise areas that would serve to motivate the cows to move even more once there, as has been observed in other studies when such resources were provided [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e].\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eOur study found that the provision of daily exercise, regardless of conditions, has a major impact on the amount of locomotor behaviour performed by cows, leading them to take more daily steps while engaging in a greater range of other natural behaviours. Providing access to an exercise area for at least 1 hour allows cows to meet their behavioural needs for locomotion, and in our study, led cows to double their number of daily steps, with outing conditions such as the type of access, the amount of space provided, and the duration of the outing playing a role in both step activity and behaviours performed. Additional resources such as shelter structures, feed, or water could provide a more enriching experience to the cows brought to the exercise area, further enhancing the benefits of such a practice on cow locomotor activity and welfare.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cp\u003e\u003cem\u003eEthical Notes\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe use of animals in these projects and all procedures were reviewed and approved by the Animal Care Committee of McGill University and affiliated hospitals and research institutes for the experimentations conducted at McGill University (protocol #2016\u0026ndash;7794; trials W19, S19, F19) and by the Animal Care Committee of the CRSAD (Deschambault Animal Science Research Centre) for the experimentations conducted at the CRSAD (protocol #1920-BL-387; trials W20, S20, S21). All aspects of this study met the standards established by the Canadian Council on Animal Care to ensure the humane and ethical use of animals in research, and all experiments followed relevant guidelines and regulations.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eAnimals and Housing\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe present study was composed of six independent trials, each with different contexts (exercise area location and size, duration of outing, substrate, etc.), in order to test different exercise access conditions intended to increase the opportunity for movement provided to cows housed in a movement-restricted environment (i.e., tie-stall cows with no routine exercise access). While this precluded the possibility of analyzing all six trials together in one single model, it allowed for an in-depth comparison of specific conditions of the outings across trials. Outing conditions tested were as follows: 1) exercise vs non-exercise treatment, 2) indoor vs outdoor exercise area, and 3) a combination of outing duration (1h vs 2h) and size of exercise area (20, 40, 60 or 80 m\u0026sup2;). In this section, the experimental setups for each of the six trials are briefly described. An overview of the different setups is shown in Table 1.\u003c/p\u003e\n\u003cp\u003eIn the W19 and S19 trials, 24 na\u0026iuml;ve, tie-stall-housed Holstein dairy cows from the McGill herd were selected and grouped in eight trios by days in milk and parity. Cows from each trio were then randomly assigned to one of three treatments: Exercise treatment cows (cows provided with outdoor access), non-Exercise treatment cows (cows remained at their stalls all day), and companion cows (cows for which data was not recorded but which accompanied Exercise treatment cows to avoid isolation and allow for social interaction).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn the F19 trial, 30 cows from the McGill herd were blocked by days in milk and parity into five groups of six cows each -three Exercise treatment cows (provided with outdoor access) and three non-Exercise treatment cows (who remained at their stalls). One of the three cows in each trio had previous experience outdoors during an earlier trial.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn the W20 and S20 trials, 27 and 30 cows respectively, were selected from the CRSAD herd and grouped into trios by days in milk and parity. Cows from each trio were randomly assigned into one of three treatments: Indoor Exercise (cows provided with indoor area access), Outdoor Exercise (cows provided with outdoor area access) and non-Exercise (cows remaining in their stalls for the duration of the treatment).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eFinally, in the S21 trial, 18 cows from the CRSAD herd were selected and grouped into trios by days in milk and parity, and randomly assigned to either: Outdoor-Biofilter1 Exercise treatment (cows providing with outdoor access, the ground of the exercise area covered by the biofilter1 (see biofilter details below)), Outdoor-Biofilter2 Exercise treatment (cows providing with outdoor access, the ground of the exercise area covered by the biofilter2) and non-Exercise treatment (cows remaining in the stall for the duration of the treatment).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAll the animals selected to participate in the different trials were healthy and non-lame lactating Holstein cows (evaluated based on a visual locomotion score adapted from [21]). During the study, all cows were housed in a mechanically-ventilated barn consiting of cubicle tie-stalls (stall area 2.7m\u003csup\u003e2\u003c/sup\u003e (W19, S19, F19) or 2.9m\u003csup\u003e2\u003c/sup\u003e (W20, S20, S21)) fitted with rubber mats and either wood chips (W19, S19, F19) or straw (W20, S20, S21). Cows had ad libitum access to water and were fed a total mixed ration (TMR) 2-4 times daily which was pushed up throughout the day to ensure feed accessibility. Milking was conducted twice daily at 9.5-14.5 h intervals.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eProcedures\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eGeneral process\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eFor the W19, S19, and F19 trials, during 8 weeks (W19 and S19) or 5 weeks (F19) cows went out every morning, five days per week (weekends excluded) for 1-2 h (treatment dependent) at 10AM, to an outside grassland exercise yard (1344\u0026nbsp;m\u0026sup2;) adjacent to the barn (as demonstrated in [17], figure 1) in pairs in the winter and summer, and in trios in the fall. The trip distance from the stall to the exercise area measured approximately 56 m and was concrete inside and dirt outside. The outdoor exercise area consisted of paddocks delimited by electric fencing. Neither feed, water nor shelter were provided during outdoor access. In the W19 and S19 trials, the size of and time spent in the paddock changed weekly according to a Latin square design, such that each week each pair was assigned to one of the eight treatments: 1h-20m\u003csup\u003e2\u003c/sup\u003e, 1h-40m\u003csup\u003e2\u003c/sup\u003e, 1h-60 m\u003csup\u003e2\u003c/sup\u003e, 1h-80 m\u003csup\u003e2\u003c/sup\u003e, 2h-20m\u003csup\u003e2\u003c/sup\u003e, 2h-40m\u003csup\u003e2\u003c/sup\u003e, 2h-60 m\u003csup\u003e2\u003c/sup\u003e, or 2h-80 m\u003csup\u003e2\u003c/sup\u003e. In the F19 trial, cow trios were assigned to a different paddock each week, all paddocks measuring 117 m\u0026sup2; (9 m x 13 m; 39 m\u0026sup2; per cow). The ground cover was grass and snow in fall, wood chips and snow in winter, and grass during summer.\u003c/p\u003e\n\u003cp\u003eFor the S20, W20, and S21 trials, cows were provided with exercise access either 2 days per week for 1 hour per day (W20), or 5 days per week (weekends excluded) for 1.5 h per day (S20, S21) at the same time daily (morning or afternoon). In the W20 and S20 trials, cows were brought to either one of two outdoor exercise yards located adjacent to the barn (28.73 m\u0026sup2; each (5.36 m x 5.36 m; one cow per area)), or to an indoor exercise area (29.60 m\u0026sup2; (7.29 m x 4.06 m; one cow allowed at a time)), composed of three full walls and one half-height wall, allowing cows to see other cows in their stalls. The ground surface consisted of a concrete floor covered with a rubber mat and wood shavings. In the S21 trial, cows were brought to one of the two outdoor yards, consisting in this trial of different biofilter substrates composed of either wood chips and gravel, or wood chips, biochar, and sphagnum moss. These trials were designed as Latin square designs, with each cow experiencing one treatment (outdoor, indoor, or non-exercise in W20 and S20 or outdoor biofilters1, 2, or non-exercise in S21) for either 3 (W20) or 1 (S20, S21) week(s) before being rotated to the next treatment. During winter, snow covered the outdoor areas most of the time. The trip distances from the stall to the outdoor and indoor exercise area measured approximately 34 m and 19 m, respectively, and was concrete inside and dirt outside.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eSpecific Notes by trial\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eWinter 2020.\u0026nbsp;\u003c/em\u003eThe original setup of this trial was designed as a 3x3 Latin square in which the same treatment was applied for three consecutive weeks before switching to a new treatment. The trial was intended to last nine weeks but, due to the COVID-19 pandemic, the last 3-week period had to be dropped. Thus, the trial lasted six weeks, each cow receiving only two treatments for three weeks each.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eSummer 2021.\u0026nbsp;\u003c/em\u003eThe trial was initially designed as a 3x3 Latin square in which each cow would spend one week in each of the three treatments: Outdoor-Biofilter1 Exercise treatment, Outdoor-Biofilter2 Exercise treatment, and non-Exercise treatment. However, due to difficulties encountered in having some of the cows adapt to the outdoor exercise areas, the final dataset included six cows for three weeks, blocked in two groups of second and third lactation respectively.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eMeasures\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eStep Activity\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eLocomotor behaviour (evaluated as the number of steps) was recorded continuously during each trial, beginning the first day of the trial, using a 3D pedometer (Peacock Technology Ltd, Stirling, Scotland, UK), attached to either rear leg of the cows. IceManager software (IceRobotics, Edinburgh, Scotland) was used to extract the raw data in 1-min intervals. The use of these pedometers to record step activity in tie-stall housed cows has previously been validated [22].\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eOther Behaviours\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eBehavioural observations were conducting using either live observations (W19, S19, F19 and S21) or video recordings (S20) when the cows were in the exercise areas (indoor and outdoor). The frequency of the behaviours observed was recorded (Table 2). Observations began once the cows were calm and the barn staff no longer visible, and lasted either 30 min (F19), 1h (S20), 1.5h (S21), or 2h (W19 and S19). Data was collected by one to three observers, either by scan sampling every 30 s (S20 and S21) or every 4 min (W19, S19 and F19). These observations took place once a week for a maximum of one day (S20), two days (F19), three days (S21), six days (W19), or eight days (S19) of observations per cows.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eStatistical Analysis\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eData handling\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eAll statistical data handling and analyses were\u0026nbsp;carried out using\u0026nbsp;R (version 1.4.1103, R Core Team, 2021) and its specific packages.\u0026nbsp;Four data sets were created. The first set included all trials described in the \u0026ldquo;Animals and Treatments\u0026rdquo; section and allowed us to investigate our primary objective, which was to know if exercise access, regardless of the conditions under which it was provided, had an influence on the number of daily steps. The second dataset included the trials conducted in the W20 and S20 and investigated the influence of the type of exercise area (indoor vs. outdoor) on the number of daily steps. The third dataset included trials conducted in the W19 and S19 and served to evaluate the influence of the duration of access to exercise and the size of the exercise area on the number of daily steps. Finally, the fourth dataset included information on the behaviours performed by the cows while in the exercise areas during all trials. For all analyses,\u0026nbsp;week was coded to correspond to the week of treatment application at the cow level, and not the calendar week.\u0026nbsp;The cleaning data method is available in Supplementary material.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eDataset 1: Exercise vs non-Exercise treatments.\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eFor each trial,\u0026nbsp;the \u003cem\u003elme4\u003c/em\u003e package [23]\u0026nbsp;was used to perform the following generalized linear mixed models (\u003cstrong\u003eGLMMs\u003c/strong\u003e) based on a Poisson distribution in order to evaluate the effect of exercise vs non-exercise within each trial. For each model, we considered the following factors: (1) treatment as a fixed effect: exercise treatments dependent on trial (outdoor exercise, indoor exercise, outdoor exercise biofilter 1 or 2) vs non-exercise; (2) week as a fixed effect: each week according to the trial; (3) block as a fixed effect (cows blocked by parity and days in milk); (4) Cow nested within block as a random effect; and, for trials W20, S20, S21, the order in which treatment was applied (as it changed weekly) as a random effect.\u003c/p\u003e\n\u003cp\u003eFor all these GLMMs, the independence between the residuals of the model and the predicted values, as well as the absence of over-dispersion of the residuals were assessed. In case of over-dispersion of the residuals, the model was adjusted by replacing the Poisson distribution by a negative binomial distribution using the \u003cem\u003eglmer.nb\u003c/em\u003e function from the \u003cem\u003elme4\u003c/em\u003e package.\u003c/p\u003e\n\u003cp\u003eFrom each model, the Least Square means (\u003cstrong\u003eLSmeans\u003c/strong\u003e) and standard deviation (\u003cstrong\u003eSD\u003c/strong\u003e) were extracted using the \u003cem\u003eemmeans\u003c/em\u003e function of the \u003cem\u003eemmeans\u003c/em\u003e package [24].\u0026nbsp;The LSmeans were used to provide adjusted probabilities by accounting for the effect of each significant variable. We used these data to investigate the number of steps taken by cows with and without access to an exercise area. As the number of steps was recorded in the same way in all trials, a between-Group Mean difference (\u003cstrong\u003eMD\u003c/strong\u003e) meta-analysis was performed using the \u003cem\u003emetacont\u003c/em\u003e function of the \u003cem\u003emeta\u003c/em\u003e package [25]. Each trial could contribute more than one comparison (if different treatments were used in one trial), but the number of steps in non-Exercise treatment was always used as the control. As between-study heterogeneity was possible due to the trials\u0026rsquo; different seasons and localizations, a random-effects model was used. The restricted maximum likelihood estimator [26] was used to calculate the heterogeneity variance \u0026tau;\u0026sup2;. We used Knapp-Hartung adjustments [27] to calculate the confidence interval around the pooled effect.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eDataset 2: Type of exercise area\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eTo determine if the type of exercise area, outdoor vs. indoor, had an impact on the daily number of steps, trials S20 and W20 containing these two outing conditions were grouped in the same dataset.\u0026nbsp;We repeated a similar process to above, using the\u0026nbsp;\u003cem\u003elme4\u003c/em\u003e package [23] to perform a generalized linear mixed model based on a Poisson distribution, considering the following factors: (1) treatment as a fixed effect: outdoor exercise vs indoor exercise; (2) week as a fixed effect: each week according to the trial; (3) block as a fixed effect (cows blocked by parity and days in milk); (4) Cow nested within block as a random effect; and order in which treatment was applied (as it changed weekly) as a random effect.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIndependence between the residuals of the model and the predicted values, as well as absence of over-dispersion of the residuals, were assessed. Due to over-dispersion of the residuals, the model was adjusted by replacing the Poisson distribution by a negative binomial distribution using the \u003cem\u003eglmer.nb\u003c/em\u003e function from the \u003cem\u003elme4\u003c/em\u003e package.\u0026nbsp;A type II Anova with Wald tests was carried out and LSmeans were used to perform post-hoc pairwise\u0026nbsp;Bonferroni corrected comparisons. Results are presented as least square means with SEs unless otherwise stated. The significance threshold used was P \u0026lt; 0.05.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eDataset 3: Duration of outing and size of the exercise area\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eTo determine if conditions of access to exercise such as the duration of the outings or the size of the outdoor exercise areas had an impact on the number of daily steps taken by the cows, the trials S19 and W19 containing these outing conditions were grouped in the same dataset. \u0026nbsp;The\u0026nbsp;\u003cem\u003elme4\u003c/em\u003e package [23] was used to perform a generalized linear mixed model based on a Poisson distribution, considering the following factors: (1) duration as a fixed effect: 1 vs 2 hours; (2) week as a fixed effect: each week according to the trial; (3) block as a fixed effect (cows blocked by parity and days in milk); (4) Cow nested within block as a random effect; and order in which treatment was applied (as it changed weekly) as a random effect.\u003c/p\u003e\n\u003cp\u003eIndependence between the residuals of the model and the predicted values, as well as absence of over-dispersion of the residuals were assessed. In case of over-dispersion of the residuals, the model was adjusted by replacing the Poisson distribution by a negative binomial distribution using the \u003cem\u003eglmer.nb\u003c/em\u003e function from the \u003cem\u003elme4\u003c/em\u003e package.\u0026nbsp;A type II Anova with Wald tests was carried out and LSmeans were used to perform post-hoc pairwise\u0026nbsp;Bonferroni corrected comparisons. Results are presented as least square means with SEs unless otherwise stated. The significance threshold used was P \u0026lt; 0.05.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eDataset 4: Other Behaviours\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eDescriptive statistics were used to obtain the percentage of time allocated to each of the behavioural activities by each animal within each trial.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eFunding\u003c/h2\u003e\n\u003cp\u003eFunding for conducting research was provided by the Natural Sciences and Engineering Research Council of Canada, Novalait, Dairy Farmers of Canada, and Valacta as a part of the Industrial Research Chair in the Sustainable Life of Dairy Cattle, as well as Agriculture and AgriFood Canada and Dairy Farmers of Canada through AgriScience Clusters program, Growing Forward III.\u003c/p\u003e\n\u003ch2\u003eCompeting Interests Statement\u003c/h2\u003e\n\u003cp\u003eThe authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.\u003c/p\u003e\n\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\n\u003cp\u003eM. Cellier: conceptualization, data curation - lead, format analysis - lead, methodology, writing original draft \u0026ndash; lead, writing review \u0026amp; editing ;E. Shepley: conceptualization, methodology, writing \u0026ndash; review and editing ; N. Aigueperse: conceptualization, methodology, writing \u0026ndash; review and editing ;M.V. Robichaud: conceptualization, funding acquisition, methodology, writing \u0026ndash; review and editing ;E. Vasseur: conceptualization, funding acquisition, investigation, methodology, project administration, resources, supervision, writing original draft \u0026ndash; lead, writing \u0026ndash; review and editing.\u003c/p\u003e\n\u003ch2\u003eAcknowledgement\u003c/h2\u003e\n\u003cp\u003eThe authors would like to acknowledge the following individuals who conducted the animal trials: McGill University M. Sc. graduates Jordan Tonooka, Catherine Lussier, Amir Nejati, Rachel Chiasson, and Universit\u0026eacute; Laval research assistant B\u0026eacute;atrice Dupont-Fortin. We extend our gratitude to the staff at the McGill Macdonald Campus Dairy Unit and Dairy Facility of the Centre de Recherche en Sciences Animales de Deschambault (CRSAD), as well as V\u0026eacute;ronique Boyer (McGill University) and Janie L\u0026eacute;vesque (CRSAD), who assisted in animal trial planning and data collection. We would like to thank Rachel VanVliet (McGill University) for the editorial suggestion and corrections, and V\u0026eacute;ronique Boyer (McGill University) for the proofreading and corrections of the article; Tania Wolfe (McGill University), Roger Cue (McGill University) and S\u0026eacute;bastien Buczinski (University of Montr\u0026eacute;al) for their help with the statistical reflection. We would like to acknowledge the contribution of environmental engineers St\u0026eacute;phane Godbout (Institut de recherche et de d\u0026eacute;veloppement en agro-environnement IRDA), S\u0026eacute;bastien Fournel (Universit\u0026eacute; Laval) and Pierre Ruel (CRSAD) in funding acquisition, experimental project planning and implementation at CRSAD.\u003c/p\u003e\n\u003ch2\u003eData Availability\u003c/h2\u003e\n\u003cp\u003eThe raw data supporting the conclusions of this article are available here : https://github.com/WELL-E-chair/data-movement-behavior?tab=readme-ov-file\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eHaskell, M. J., Brotherstone, S., Lawrence, A. B. \u0026amp; White, I. M. S. 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How to perform a meta-analysis with R: a practical tutorial. \u003cem\u003eEvid Based Ment Health.\u003c/em\u003e \u003cstrong\u003e22(4)\u003c/strong\u003e, 153-160 (2019).\u003c/li\u003e\n\u003cli\u003eViechtbauer, W. Bias and Efficiency of Meta-Analytic Variance Estimators in the Random-Effects Model. \u003cem\u003eJ Educ Behav Stat.\u003c/em\u003e \u003cstrong\u003e30(3)\u003c/strong\u003e, 261-295 (2005)\u003c/li\u003e\n\u003cli\u003eKnapp, G. \u0026amp; Hartung, J. Improved tests for a random effects meta-regression with a single covariate. \u003cem\u003eStat Med.\u003c/em\u003e \u003cstrong\u003e22(17)\u003c/strong\u003e, 2693-2710 (2003).\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003eTable\u0026nbsp;1. Summary of the characteristics of trials\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"837\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"21.479713603818617%\"\u003e\n \u003cp\u003eTrial\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.365155131264917%\" colspan=\"2\"\u003e\n \u003cp\u003eWinter 2019\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.007159904534607%\" colspan=\"2\"\u003e\n \u003cp\u003eSummer 2019\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.54653937947494%\"\u003e\n \u003cp\u003eFall 2019\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.723150357995227%\" colspan=\"2\"\u003e\n \u003cp\u003eWinter 2020\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.410501193317423%\" colspan=\"2\"\u003e\n \u003cp\u003eSummer 2020\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.46778042959427%\" colspan=\"2\"\u003e\n \u003cp\u003eSummer 2021\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"21.479713603818617%\"\u003e\n \u003cp\u003eDuration of the trial (weeks)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.365155131264917%\" colspan=\"2\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.007159904534607%\" colspan=\"2\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.54653937947494%\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.723150357995227%\" colspan=\"2\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.410501193317423%\" colspan=\"2\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.46778042959427%\" colspan=\"2\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"21.479713603818617%\"\u003e\n \u003cp\u003eNumber of cows\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.365155131264917%\" colspan=\"2\"\u003e\n \u003cp\u003e24\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.007159904534607%\" colspan=\"2\"\u003e\n \u003cp\u003e24\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.54653937947494%\"\u003e\n \u003cp\u003e30\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.723150357995227%\" colspan=\"2\"\u003e\n \u003cp\u003e27\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.410501193317423%\" colspan=\"2\"\u003e\n \u003cp\u003e18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.46778042959427%\" colspan=\"2\"\u003e\n \u003cp\u003e18\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"21.479713603818617%\"\u003e\n \u003cp\u003eNumber of blocks\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.365155131264917%\" colspan=\"2\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.007159904534607%\" colspan=\"2\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.54653937947494%\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.723150357995227%\" colspan=\"2\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.410501193317423%\" colspan=\"2\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.46778042959427%\" colspan=\"2\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"21.479713603818617%\"\u003e\n \u003cp\u003eParity per block\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.365155131264917%\" colspan=\"2\"\u003e\n \u003cp\u003e2.0 \u0026plusmn; 1.46\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.007159904534607%\" colspan=\"2\"\u003e\n \u003cp\u003e2.5 \u0026plusmn; 1.68\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.54653937947494%\"\u003e\n \u003cp\u003e1.9 \u0026plusmn; 0.79\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.723150357995227%\" colspan=\"2\"\u003e\n \u003cp\u003e2.2 \u0026plusmn; 0.93\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.410501193317423%\" colspan=\"2\"\u003e\n \u003cp\u003e2.0 \u0026plusmn; 0.84\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.46778042959427%\" colspan=\"2\"\u003e\n \u003cp\u003e2.7 \u0026plusmn; 1.00\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"21.479713603818617%\"\u003e\n \u003cp\u003eDIM block\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.365155131264917%\" colspan=\"2\"\u003e\n \u003cp\u003e149 \u0026plusmn; 63.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.007159904534607%\" colspan=\"2\"\u003e\n \u003cp\u003e112 \u0026plusmn; 63.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.54653937947494%\"\u003e\n \u003cp\u003e163 \u0026plusmn; 66.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.723150357995227%\" colspan=\"2\"\u003e\n \u003cp\u003e117 \u0026plusmn; 65.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.410501193317423%\" colspan=\"2\"\u003e\n \u003cp\u003e187 \u0026plusmn; 99.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.46778042959427%\" colspan=\"2\"\u003e\n \u003cp\u003e173 \u0026plusmn; 102\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"21.479713603818617%\"\u003e\n \u003cp\u003eTreatments\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.365155131264917%\" colspan=\"2\"\u003e\n \u003cp\u003eDuration x Size\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.007159904534607%\" colspan=\"2\"\u003e\n \u003cp\u003eDuration x Size\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.54653937947494%\"\u003e\n \u003cp\u003eOutdoor\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.723150357995227%\" colspan=\"2\"\u003e\n \u003cp\u003eOutdoor\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.410501193317423%\" colspan=\"2\"\u003e\n \u003cp\u003eIndoor\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.46778042959427%\" colspan=\"2\"\u003e\n \u003cp\u003eOutdoor-Biofilter1\u003c/p\u003e\n \u003cp\u003eOutdoor-Biofilter2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"21.428571428571427%\"\u003e\n \u003cp\u003eControl\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.095238095238095%\"\u003e\n \u003cp\u003eStall\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.095238095238095%\" colspan=\"2\"\u003e\n \u003cp\u003eStall\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.095238095238095%\" colspan=\"3\" style=\"width: 13.5006%;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Stall\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.095238095238095%\" colspan=\"2\" style=\"width: 13.0227%;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; Stall\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.095238095238095%\" colspan=\"2\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp;Stall\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.095238095238095%\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; Stall\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"21.479713603818617%\"\u003e\n \u003cp\u003eOutings/Week\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.365155131264917%\" colspan=\"2\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.007159904534607%\" colspan=\"2\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.54653937947494%\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.723150357995227%\" colspan=\"2\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.410501193317423%\" colspan=\"2\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.46778042959427%\" colspan=\"2\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"21.479713603818617%\"\u003e\n \u003cp\u003eDuration per outing (hours)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.365155131264917%\" colspan=\"2\"\u003e\n \u003cp\u003e1 or 2*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.007159904534607%\" colspan=\"2\"\u003e\n \u003cp\u003e1 or 2*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.54653937947494%\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.723150357995227%\" colspan=\"2\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.410501193317423%\" colspan=\"2\"\u003e\n \u003cp\u003e1.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.46778042959427%\" colspan=\"2\"\u003e\n \u003cp\u003e1.5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"21.479713603818617%\"\u003e\n \u003cp\u003eSize of the area (m\u0026sup2;)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.365155131264917%\" colspan=\"2\"\u003e\n \u003cp\u003e20, 40, 60, 80*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.007159904534607%\" colspan=\"2\"\u003e\n \u003cp\u003e20, 40, 60, 80*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.54653937947494%\"\u003e\n \u003cp\u003e117\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.723150357995227%\" colspan=\"2\"\u003e\n \u003cp\u003e28.73(outdoor)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.410501193317423%\" colspan=\"2\"\u003e\n \u003cp\u003e29.60(indoor)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.46778042959427%\" colspan=\"2\"\u003e\n \u003cp\u003e28.73\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"21.479713603818617%\"\u003e\n \u003cp\u003eLocalization\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.365155131264917%\" colspan=\"2\"\u003e\n \u003cp\u003eMcGill\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.007159904534607%\" colspan=\"2\"\u003e\n \u003cp\u003eMcGill\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.54653937947494%\"\u003e\n \u003cp\u003eMcGill\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.723150357995227%\" colspan=\"2\"\u003e\n \u003cp\u003eCRSAD\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.410501193317423%\" colspan=\"2\"\u003e\n \u003cp\u003eCRSAD\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.46778042959427%\" colspan=\"2\"\u003e\n \u003cp\u003eCRSAD\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e* depending on the treatment (combination between duration and size) applied\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eTable 2. Detailed ethogram used to assess cow activity during the different trials\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"938\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"9.913793103448276%\"\u003e\n \u003cp\u003eCategory\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.452586206896552%\"\u003e\n \u003cp\u003eBehaviour\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"67.13362068965517%\"\u003e\n \u003cp\u003eDescription\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.5%\"\u003e\n \u003cp\u003eTrial\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"0%\" height=\"21\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"9.913793103448276%\" rowspan=\"2\"\u003e\n \u003cp\u003eIdle\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.452586206896552%\"\u003e\n \u003cp\u003eIdle\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"67.13362068965517%\"\u003e\n \u003cp\u003eBody relaxed, immobile with head down or in the extension of her body. Cow can do other activities such as ruminating or standing alert, while idling\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.5%\"\u003e\n \u003cp\u003eF19, S19, W19, S20, S21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"0%\" height=\"41\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"11.602870813397129%\"\u003e\n \u003cp\u003eRumination\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"74.52153110047847%\"\u003e\n \u003cp\u003eRhythmic/repeated chewing motions of partly digested food not associated with head down in feed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.875598086124402%\"\u003e\n \u003cp\u003eF19, S19, W19\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"0%\" height=\"21\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"9.913793103448276%\"\u003e\n \u003cp\u003eVigilance\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.452586206896552%\"\u003e\n \u003cp\u003eVigilance\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"67.13362068965517%\"\u003e\n \u003cp\u003eBody immobile with head up. Cow standing alert, with the ears facing forward\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.5%\"\u003e\n \u003cp\u003eF19, S19, W19\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"0%\" height=\"41\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"9.913793103448276%\"\u003e\n \u003cp\u003eLying\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.452586206896552%\"\u003e\n \u003cp\u003eLying\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"67.13362068965517%\"\u003e\n \u003cp\u003eCows were considered lying when the flank of the animal was in contact with a surface\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.5%\"\u003e\n \u003cp\u003eF19, W19, S20, S21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"0%\" height=\"41\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"9.913793103448276%\"\u003e\n \u003cp\u003eExploration\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.452586206896552%\"\u003e\n \u003cp\u003eExploring\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"67.13362068965517%\"\u003e\n \u003cp\u003eCow is sniffing or licking the ground, feed, stall structure, or any other object of her surroundings. Cow is not walking during exploration.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.5%\"\u003e\n \u003cp\u003eF19, S19, W19, S20, S21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"0%\" height=\"41\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"9.913793103448276%\" rowspan=\"4\"\u003e\n \u003cp\u003eLocomotion\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.452586206896552%\"\u003e\n \u003cp\u003eWalking\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"67.13362068965517%\"\u003e\n \u003cp\u003eMovement forward of the feet, with one- or two-feet lift. Walking is considered when the cow takes at least 3 steps. Cow can ruminate and/or explore the ground or any object on her surroundings while walking\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.5%\"\u003e\n \u003cp\u003eF19, S19, W19, S20, S21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"0%\" height=\"52\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"11.602870813397129%\" rowspan=\"2\"\u003e\n \u003cp\u003eRunning\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"74.52153110047847%\" rowspan=\"2\"\u003e\n \u003cp\u003eFast movement forward of the parallel feet which touch and lift simultaneously with a phase of suspension in air\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.875598086124402%\" rowspan=\"2\"\u003e\n \u003cp\u003eS20, S21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"0%\" height=\"31\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd height=\"21\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd\u003e\n \u003cp\u003eJumping\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eFour legs lifted from the ground\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eS20\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd height=\"21\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd\u003e\n \u003cp\u003eSocial\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eSocial interactions\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eAny positive or negative behaviour involving counterparts (e.g., mutual play headbutt, allo-grooming...)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF19, S19, W19\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd height=\"35\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd\u003e\n \u003cp\u003eEating\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eEating\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eCow is actively grabbing and chewing feed with head down, or any chewing movement after gathering a mouthful of feed. Cow can walk while eating\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF19, S19, W19\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd height=\"35\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"3\"\u003e\n \u003cp\u003eMaintenance\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eSelf-grooming\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eTongue in contact with own skin or fur/scratches with hind legs or any sort of structure to scratch against\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF19, S19, W19, S20, S21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd height=\"41\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd\u003e\n \u003cp\u003eDefecation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eElimination of feces from the body\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eS19, S20, S21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd height=\"41\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd\u003e\n \u003cp\u003eUrination\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eElimination of urine from the body\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eS19, S20, S21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd height=\"21\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd\u003e\n \u003cp\u003eOther\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eOthers\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eAny other activity not described here\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF19, S19, W19, S20, S21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd height=\"41\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003eW19: Winter 2019 trial; S19: Summer 2019 trial; F19: Fall 2019 trial; W20: Winter 2020 trial; S20: Summer 2020 trial; S21: Summer 2021 trial\u003c/p\u003e\n\u003cp\u003eTable 3: Descriptive statistics for each trial, after the data cleaning steps\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd colspan=\"2\" valign=\"bottom\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eWinter 2019\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eSummer 2019\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eFall 2019\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eWinter 2020\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eSummer 2020\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eSummer 2021\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eN of cows\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003eNon-Exercise\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003eExercise\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eTreatment A\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e19\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eTreatment B\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e17\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eLSmean number of daily steps \u0026plusmn; SD\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003eNon-Exercise\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e543 \u0026plusmn; 155\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e616 \u0026plusmn; 671\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e442 \u0026plusmn; 228\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e547 \u0026plusmn; 197\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e725 \u0026plusmn; 207\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e528 \u0026plusmn; 207\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003eExercise\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e777 \u0026plusmn; 264\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e1233 \u0026plusmn; 482\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e763 \u0026plusmn; 393\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eTreatment A\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e835 \u0026plusmn; 314\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e1239 \u0026plusmn; 354\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e770 \u0026plusmn; 279\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eTreatment B\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e765 \u0026plusmn; 282\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e954 \u0026plusmn; 266\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e968 \u0026plusmn; 380\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003eFor Winter 2020 and Summer 2020, treatment A and B corresponded to Outdoor Exercise and Indoor Exercise, respectively.\u0026nbsp;\u003cbr\u003e\u0026nbsp;For Summer 2021, treatment A and B corresponded to Outdoor-Biofilter1 Exercise and Outdoor-Biofilter2 Exercise, respectively.\u0026nbsp;\u003cbr\u003e\u0026nbsp;Note that for Winter 2020, Summer 2020 and Summer 2021, the cows went through each treatment (Exercise vs. non-Exercise) consecutively, while in Winter 2019, Summer 2019 and Fall 2019, Exercise and non-Exercise cows were different individuals.\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"behavioural needs, locomotor behaviour, movement opportunity, steps","lastPublishedDoi":"10.21203/rs.3.rs-4953052/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4953052/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eIntensification in animal productions has led to increased indoor confinement for animals, thereby limiting their opportunities to fulfill some behavioural needs such as exploration and locomotion. This poses a crucial question as to how these restrictions can be alleviated by providing additional space designed with that specific purpose in mind. Working with cows housed in a movement-restricted environment, our study aimed to: 1) quantify how providing an exercise area outside the normal housing environment affects locomotor behaviour; 2) evaluate how exercise access conditions can contribute to optimize benefits for animals; 3) investigate the cows\u0026rsquo; time budgets during exercise access. Six trials involving different exercise access conditions (indoor vs outdoor, outing duration, exercise area size) enrolled 141 tie-stall-housed lactating Holsteins. A meta-analysis compared daily steps for exercise vs non-exercise treatments, while generalized linear mixed models determined exercise conditions\u0026rsquo; impact on daily steps. Providing 1-hour exercise access increased daily steps by 53% (304 more steps), influenced by type of access (167 more steps outdoor vs indoor), larger space (146 more steps) and longer outings (84 more steps). Cows spent 50\u0026ndash;85% of exercise time idle, exploring (5\u0026ndash;20%) and socializing (5%). Our results highlight the significant impact of 1h daily exercise on movement-restricted cattle.\u003c/p\u003e","manuscriptTitle":"Enhancing movement opportunity and fulfilling behavioural needs for movement- restricted cattle by providing different conditions of access to exercise","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-09-20 06:57:13","doi":"10.21203/rs.3.rs-4953052/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2024-10-14T01:38:14+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-10-11T09:29:07+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"168324675388876398917226230931221948205","date":"2024-09-30T07:04:30+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-09-25T13:16:10+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"45809711849994348891304612332550354460","date":"2024-09-23T10:46:55+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-09-21T14:36:42+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-09-18T12:14:31+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2024-08-22T07:52:38+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2024-08-22T03:07:49+00:00","index":"","fulltext":""},{"type":"submitted","content":"Scientific Reports","date":"2024-08-21T16:36:29+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"635f90f0-4ad2-4f2d-a01b-257b465a25ca","owner":[],"postedDate":"September 20th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[{"id":37873680,"name":"Biological sciences/Zoology/Animal behaviour"},{"id":37873681,"name":"Biological sciences/Zoology"}],"tags":[],"updatedAt":"2025-02-24T15:58:16+00:00","versionOfRecord":{"articleIdentity":"rs-4953052","link":"https://doi.org/10.1038/s41598-025-89891-4","journal":{"identity":"scientific-reports","isVorOnly":false,"title":"Scientific Reports"},"publishedOn":"2025-02-18 15:56:51","publishedOnDateReadable":"February 18th, 2025"},"versionCreatedAt":"2024-09-20 06:57:13","video":"","vorDoi":"10.1038/s41598-025-89891-4","vorDoiUrl":"https://doi.org/10.1038/s41598-025-89891-4","workflowStages":[]},"version":"v1","identity":"rs-4953052","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4953052","identity":"rs-4953052","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}