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Wikipedia Topic/Article: Allelomimetic behaviour

italicized sentences in the beginning is copied directly from the page, other that that everything else is my own work.

Allelomimetic behaviour

Allelomimetic behaviour seen in ducks diving for food at the same time

Allelomimetic behaviour or allomimetic behaviour is a range of activities in which the performance of a behaviour increases the probability of that behaviour being performed by other nearby animals. Allelomimetic behaviour is sometimes called contagious behaviour and has strong components of social facilitation, imitation, and group coordination. It is usually considered to occur between members of the same species*. An alternate definition is that allelomimetic behaviour is a more intense or more frequent response or the initiation of an already known response, when others around the individual are engaged in the same behaviour. It is often referred to as synchronous behaviour, mimetic behaviour, imitative behaviour, and social facilitation.

Allelomimetic behaviour is displayed in all animals and can occur in any stage of life, but usually starts at a young age. This behaviour will continue throughout life, especially when an individual is living in a large group that emphasizes group cohesion. Cohesion is seen as a prerequisite for group living, with synchronous activity being crucial for social cohesion. However, animals in large cohesive groups face trade-offs when allelomimetic behaviour is adopted. If the behaviour is adopted then the risk of predation or capture decreases significantly but the inter-individual competition for immediate resources, such as food, mates, and space, will increase when cohesion is still stressed. Many collective group decisions in animals are the result of allelomimetism and can be explained by allelomimetic behaviours. Some examples are the cockroaches choosing a single aggregation site, schooling behaviours in fishes, and pheromone-based path selection in ants that allows all the workers to go down the same path to a specific food source.Allelomimetic behaviour can also be seen as an animal welfare indicator. For example, if cattle do not have enough room to all lie down simultaneously then it indicates that there are not enough resources present and this can result in lameness of the animals that are forced to stand. Allomimicry is affected by circadian rhythms and circadian cycles of activity within groups which can give the overall appearance of poor animal welfare, if allomimetic behaviour were to be used as a welfare indicator then it must be measured several times throughout the course of a day. It should be noted however, that most mechanisms involved in performing allelomimetic behaviour do not require circadian rhythms to function. Decisions at the individual level are, more often than not, enough to encourage allelomimetism. Patterns of allelomimetic behaviour can vary from species to species and can possibly explain other behaviours seen in the animal kingdom.

Threats to Allelomimetic Behaviour

Synchronous behaviour is also threatened when animals in a mixed-sex herd have differing nutritional or physical necessities. This causes group instability which often splits the herd up into two separate groups; generally all male and all female, to recreate the mimetic behaviour in a smaller, same-sex group that has more similar needs to the individual. Synchronized, allelomimetic behaviour is also affected by many factors, such as age, general group size, sex, space, resource availability, and domestication. Domestication can also be seen as a threat to allelomimetic behaviour. The process of domestication removes many threats like predation, food shortages, and competition from many individuals for breeding by providing basically unlimited food and resources while providing protection from outside predators. Domestication may favour less synchronization for animals and provide an adaptation to mimetic behaviour to save energy in domesticated animals. Domestication also changes the inter-individual distances between animals and behavioural synchrony in general, both of which are important for anti-predator strategies and responses.

In Horses

Allelomimetic behaviour shown in grazing horses. Postural imitation is seen in regards to the stance and position of the legs.

Horses, like many animals, learn through imitation of their parents, most often the mother or "broodmare", or the owner of the foal if the broodmare is not available. If the foal is paired with its mother it will not only mimic the mare's walking style and speed, but it will also mimic the mare's temperament and general behaviours. If a foal was to spend much of its early life around a mare that had an unstable temperament, irrational fears and was difficult to work with then it would grow to be an unstable adult that would develop behavioural abnormalities such as balking. Miller (1996) also adds that behavioural issues can also be removed through allelomimetic processes. Balking is a common behaviour in horses and mules that occur from insecurities and fears, and is observed as hesitance with a general fear of moving forward due to the possibility of attack or invasion of their personal space. This behaviour can be eliminated if the owner is fearless, looking forward with their head raised high and marching in place. This shows the horse that there is nothing to fear and eventually the horse will mimic the owner's behaviour, sync their gate to the owner and follow along. Repeating this behaviour as needed will eventually allow the horse to fully incorporate the owner's fearless behaviour into its repertoire and prevent balking from reoccurring. Bad habits are also learned through mimicry if the vast majority of the animals present a specific habit.Cribbing is an example of one such habit. If a foal is raised in an environment where this abnormal behaviour is performed by the older members of the team, then it will mimic the habit as well, even if the behaviour is dangerous to the individual's health. However, the same mimetic behaviour that produces these bad habits can help remove them as well. If a broodmare is unable to train her offspring then it falls to the owner to properly train the foal on proper walking gait, speed, and normal behaviours. If the owner spends enough time with their foal keeping a proper stance (ie, an erect, unmoving torso and flexed knees while walking) then the foal will mimic and perform the proper movements for the rest of its life.

In Sheep

Sheep herd spreading out and displaying foraging behaviour

Sheep provide a good basis for the evaluation of allelomimetic behaviour due to their large group sizes and social behaviour. Using them as an experimental subject allows for the determination of the imitative quality and intensity of allelomimetic behaviour within a specific herd. Merino sheep, or Ovis aries, are a prey species and a domesticated breed of sheep that require a healthy balance between predator avoidance and foraging space for each individual in the herd. They achieve this balance by spreading out to forage for a period of time then quickly running back to the centre of the herd, creating a fastpacking event. A fastpacking event is specifically when an individual on the outskirts of the herd changes its behaviour from grazing to running and moves towards the centre of the herd. Fastpacking events are seen as an adaptive behaviour for reducing predation due to the intensity of the response presented in other sheep when one individual is seen running from the outer edges to the centre of the herd. This behavioural change is also referred to as activation/inactivation rates or switching behaviour. This change influences other sheep to start running to the centre of the herd until all sheep simultaneously stop in the middle and form a tightly packed herd. The cycle will restart when the herd starts to spread out to forage again. The individual that is displaying the switching behaviour from inactivity (grazing in one spot) to activity (running towards the centre of the herd) or vice versa is generally referred to as the herd leader, and other individuals who mimic the switch from inactivity to activity or activity to inactivity are displaying allelomimetic behaviour. These activation and inactivation rates are generally more strongly presented in males compared to females and increase as the number of individuals performing an activity increase which is a good indication of imitation present in the herd. For example, if a herd had the majority of individuals active the likelihood of another random individual becoming active increases. The same goes for a herd with a majority of individuals being inactive. In experiments, these events are not specific to a singular spot or caused by any external stimuli that would warrant an alarm response in the sheep. In an experiment performed by Gautrais, members of the same sex more often performed mimetic behaviour with each other than when compared to members of a mixed-sex group. Gautrais also focused on behavioural synchrony of activity and inactivity since cohesion of a group involves individuals being active and inactive at the same time rather than syncing every activity to another individual.

Even in the absence of other animals, individuals will switch between behaviours that require activity and inactivity (such as digestion, inactivity is good to promote digestion or hunger outweighing the need for rest). Gautrais believes that this physiological need is what prompted the first individual in his herd experiment to switch from activity to inactivity or vice versa and the other members in the herd to follow suit. However, as the number of individuals in a heard decreases, the option to perform mimetic behaviour also decreases. This creates a higher rate of spontaneous switching between activity and inactivity-related behaviours. In contrast, the larger the herd is, the more likely it is for mimetic behaviours to occur with spontaneous switching behaviour occurring in the "herd leader", with other members following suit in a brief period of time. This is due to the number of con-specifics present at any given time. If the herd is a small group then there are fewer con-specifics which gives less opportunity for imitation to occur in any given individual and a higher rate of switching from inactivity to activity to inactivity will occur. Two ideas are proposed when the original running individual can recruit others into the running group and the herd as a whole can coordinate when to stop running during these fastpacking events. The first idea is that the initiation and inhibition of the packing event is based off of allelomimetic effects in local interactions, and the second idea was that a close enough distance between herd neighbours will stop running behaviour. These conclusions stem from an idea about optimization processes in sheep that, at an individual level, allows for the balance of exploring as much space as possible to avoid competition for food and keeping in contact with other herd members to avoid predation and reinforce herd cohesion. These optimization processes can adjust the allelomimetic interaction strengths between individuals to ensure both of the above necessities are met.

In Cattle

Social facilitation (allelomimetic behaviour) in cattle as seen from the pictured cows all resting at the same time

The synchronous behaviour that is displayed in cattle is what is typically thought of when discussing allelomimetic behaviour. When the necessary resources for survival, such as food, water, and shelter, are sufficient then cattle will display postural synchrony, lying down or standing while other members of the herd are doing the same. This phenomenon, also referred to as social facilitation, occurs in both free-range cattle that live in pastures, and cattle that live primarily in barns. Social facilitation (or allelomimicry) is the first of two main ideas behind cattle mimicking each other's behaviour until the herd acts synchronously, whether it be lying down, standing, or grazing together. According to Stoye and Porter, one member of the herd will alter its behaviour (ie. standing to lying down or vice versa) and its nearest neighbour will mimic the behaviour, this behaviour will travel across the herd until all members are performing one action. Stoye and Porter also noted that it was more likely for a cow to display the same behaviour as its nearest neighbour compared to a randomly selected cow from the herd. This behaviour continues despite the appearance of free will of an individual to graze/stand or lie down, even when the herd is stationary. The second idea behind synchronized behaviour is called concurrent or combined response. This dictates that the collective behaviour is a mere coincidence of each individual animal's decision to change their behaviour based on external (the discovery of food) or internal (exhaustion, the need for rest/food, or similar daily schedules as other members) factors. A human example of a concurrent behaviour is rush hour, many people are driving at the same time due to having a similar schedule to others, not because others are already driving and they decide to as well. Previous studies have identified artificial synchronization of (dairy) cattle behaviour because of consistent milking and feeding times. This creates a similar daily schedule for all the cattle to follow, which in turn creates collective behaviour between all members during these specific milking and feeding times. According to Stoye and Porter, cattle were most synchronous at night, least synchronous during the afternoon, and intermediately synchronous in the morning. This could be due to all the cattle being milked at similar times in the morning and feeding at similar times during the night, but having no "scheduled activity" to perform during the afternoon.

In Dogs

Feeding Habits

Dogs are one of the most common species to produce allelomimetic behaviour and have it go noticed by humans. Studies have been conducted across several breeds of dog regarding eating habits and stress vocalization when alone or with litter-mates. Many results have shown that most dogs will eat more when they are in large groups compared to when they eat alone. This has been coined as the "social facilitation of eating" and is not specific to certain breeds. In a study conducted by Compton and Scott, 80% of the dogs studied ate more in groups on the majority of trials and the overall group consumption together was larger than the summation of food consumption between each dog eating individually. When pups were considered food-satiated (would not eat more food even with a dish in front of them), they would resume eating when a hungry litter-mate was introduced into the room with them. It was believed that since everything other than the addition of the hungry litter-mate was held constant, the increased appetite in the food-satiated pup was caused by the presence of the hungry pup and allelomimetic behaviour. The allelomimetic behaviour that occurs in dogs develops directly from experiences that occur during the critical period of development. Scott's proposed theory of social motivation and the theory of the social facilitation of eating rely heavily on the allelomimetic behavioural system that occurs in these animals.

Isolation and Vocalization Responses

Isolation is a large stressor for many dogs and can result in separation anxiety if they are left alone from others for extended lengths of time during a critical period in their development. If a puppy is separated from their litter-mates, home pen, or owners during a critical period it will produce an intense negative emotional reaction and the reaction will subside when they return to their litter-mates, home pen, or owners. This reaction is a natural conditioned learning paradigm that will have to be constantly repeated during their infancy to remove the intensity of the response. When housed with other litter-mates the individual will be strongly motivated to mimic the behaviour of the animals around it. Introducing other litter-mates who do not experience a severe negative emotion to isolation is one of the ways to avoid negative reactions to isolation in the individual in question. This provides a base for mimetic behaviour to occur and the pup experiencing severe negative emotions will begin to mimic the behaviour of the litter-mates and adopt their neutral behaviour. Scott and Bronson replicated a study originally conducted by Fredericson done in 1952 about vocalization rates of beagle puppies when confined in a box alone or with a companion. Fredericson found that 75% of puppies would vocalize significantly less when confined with another puppy compared to alone, a result that was replicated and confirmed by Scott and Bronson in 1971. Many dogs were more distressed when they were placed alone in an unfamiliar environment compared to a known environment, or an unfamiliar environment with litter-mates. This increase in calm behaviour when multiple litter-mates are in an unfamiliar environment is caused by allelomimetic behaviour. If one pup is calm, many dogs will mimic that behaviour, creating a litter of relaxed dogs. Likewise, if there is one distressed pup, many dogs could also mimic the distressed behaviour but the former is more common among experiments.

In Chickens and Roosters

The domesticated white leghorn rooster

The wild red junglefowl

Allelomimetic behaviour can often be affected by domestication and lead to the evolution of new social behaviours, or subtle changes in current social behaviours. It is thought that domestication would reduce the level of allelomimetic behaviour in animals due to the removal of many important factors that create mimicries such as predation, food pressures and competition between species members. Since there is no need to worry about possible predation or a lack of resources in a domesticated environment, the allelomimetic behaviours seen in non-domesticated species evolved out and the adaptation to domestication became the new normal. Mimetic behaviours that once incorporated anti-predator strategies or mating strategies became unnecessary and the use of these behaviours decreased. An experiment was conducted by Eklund and Jensen using an ancestor of all domesticated chickens, the red junglefowl, and a domesticated breed, the white leghorn. They showed that allelomimetic behaviour was more prominent and used more frequently in the non-domesticated red junglefowl compared to the white leghorn, most likely due to the chance of predation, starvation, and the lack of shelter playing a role in producing these allelomimetic behaviours. Total synchronization only occurred in both species during comforting behaviours such as perching and dust bathing. In activities outside of comfort behaviour, there was little mimetic behaviour in the domesticated white leghorn and inter-individual distances presented by the chickens during perching was larger than the non-domesticated species. Perching in the red junglefowl occurred more frequently and was more mimetic than in the white leghorn. The social behaviour of the red junglefowl was also affected by allelomimetism, where behaviours such as feather pecking were more synchronized than in the white leghorn. A difference in feeding synchrony also appeared in females versus males. Females were more likely to mimic other females' eating behaviours compared to a mixed-sex group or males mimicking males.

• possibly use this article

Peer review of Allelomimetic behaviour by user Fresh oolong

I find the amount of content you added and citations substantially improved and expanded your article!

I do not personally see any major issues or challenges with your work the citations that I examined seem to be relevant.

The only advice i may offer could be another level of organization into sub-headings for fast easy access and review if someone wants to examine a specific concept related to one of your animal choices. Also, the pictures moved the paragraph structure into a for me unusual format but nothing bad.

If you are able adding one or two more citations to come of the animals may solidify some of the concepts compared to only having one source (may be challenging to find sources I am not sure)

Looks great!

Fresh oolong (talk) 03:15, 7 November 2018 (UTC)

Response to peer review

I will definitely look into reorganizing my information into sub-heading to allow for an easier time accessing the information. I'll also look into the possible formatting issues with the pictures and see if it can be fixed/less unusual. I am still looking for more citations and information to add, but since we are on a strict timeline I wanted to get what I had done, then look into adding more.

Thanks for the feedback! It's definitely helpful and it'll help me polish the work before the project is handed in.

Peer Review- by Amlftwix

Hi!

The amount of information that you added to the article is really impressive, and overall seems well written. I believe what you added to the opening really added to making the rest of the article make more sense. Everything was presented in a way that was relatively simple to understand which I think made it overall a stronger article!

One of the only criticisms that I have is that entire paragraphs, or multiple paragraphs are based on a single reference. Though these are credible sources, I do think that adding another source that supports the information you gave would increase the credibility of your article. Also, many of the pictures seem kind of randomly placed, and maybe for consistency potentially place them all on the same side of the article, but this is more of an aesthetic preference. This is a really interesting topic, and I am curious if anything can be found on it occurring in invertebrates or something of a 'lower level' of cognition.

Again really good job!

Peer Review: Cawhite88

Your contribution is very well put together with a solid structure and references that are both reliable and relevant to the subject. You obviously put a lot of work into your contribution and it is well written overall. I just have some suggestions, mainly just specific cases where the grammar was a little off.

The lead section is very well written but I'm concerned that it may not be as accessible to people without previous biological knowledge. Terms such as "inter-individual competition" and "group cohesion" may not make very much sense to a reader without giving the definitions.

The following is a list of the only grammar errors I found with suggested fixes for them (considering how much you wrote there were very few mistakes):

• "Domestication may favour" instead of "Domestication might favour"
• "a specific habit. Cribbing is an example of one such habit" instead of "a specific habit, cribbing is an example of one such habit."
• "Sheep provide a good basis for the evaluation of allelomimetic behaviour due to their large group sizes and social behaviour." instead of "Sheep provide a good basis for the evaluation of allelomimetic behaviour from their large group sizes and social behaviour."
• "They achieve this balance by spreading out to forage for a period of time then quickly running back to the center of the herd" instead of "They achieve this balance by spreading out to forage for a period of time and quickly running back to the centre of the herd"
• "allelomimetic behaviour was more prominent and used more frequently" instead of "allelomimetic behaviour was more prominent and used more frequently used"

There was only one spelling mistake: "allelomimesis"

Overall the piece was very neutral except for one statement: "The bright side of situations where bad habits or behaviours are learned and mimicked is that the same mimetic behaviour that developed these behaviours can remove them as well."

The only other issues I have are very minor structural problems:

• In the "In Horses" section: "Miller (1996) also adds that behavioural issues can also be removed through allelomimetic processes." and onward should be positioned with the part talking about behavioural issues in horses and not at the end of the section.
• In the "In Sheep" section: "Fastpacking events are seen as an adaptive behaviour for reducing predation due to the intensity of the response presented in other sheep when one individual is seen running from the outer edges to the centre of the herd." should be positioned within the section when fastpacking is first mentioned. Also, the large size of a herd effecting allelomimetic behaviour is mentioned twice in the section. "In contrast, the larger the herd is, the more likely it is for mimetic behaviours to occur with spontaneous switching behaviour occurring in the "herd leader", with other members following suit in a brief period of time. " is the better way of explaining the phenomena.

Again, this is a very well thought out and written contribution and I am very impressed with the time and effort you have put into this project!