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Development of the predatory imminence continuum

The development of the predatory imminence continuum began with the description of species-specific defence reactions. Species-specific defence reactions are innate responses demonstrated by an animal when they experience a threat. Since survival behaviours are so vital for an animal to acquire and demonstrate rapidly, it has been theorized that these defence reactions would not have time to be learned and therefore, must be innate. While these behaviours are species-specific, there are three general categories of defence reactions - fleeing, freezing, and threatening. Species-specific defence reactions are now recognized as being organized in a hierarchical system where different behaviours are exhibited depending on the level of threat experienced. However, when this concept was first proposed, the dominant species-specific defence reaction in a certain context was thought to be controlled by operant conditioning. That is, if a species-specific defence reaction was unsuccessful in evading or controlling conflict, the hierarchical system would be rearranged because of the punishment, in the form of failure, experienced by an animal. It would then be unlikely for that species-specific defence reaction to be used in a similar situation again; instead, an alternative behaviour would be dominant. However, if the dominant behaviour was successful it would remain the recurring reaction for that situation. After experimentation, this theory was met with much opposition, even by the person who proposed it. One point of opposition was found through the use of shock on rats and the species-specific defence reaction of freezing. This experiment found that while punishment did seem to have an impact on freezing, it was not through response weakening but through the inducing of different levels of the behaviour. Other criticisms for this theory focused on the inability for species-specific defence reactions to effectively rearrange in this manner in natural situations. It has been argued that there would not be enough time for punishment, in the form of an animal being unsuccessful in its defence, to reorder the hierarchy of species-specific defence reactions. The rejection of the operant conditioning mechanism for the reorganization of species-specific defence reactions, lead to the development of the predatory imminence continuum where the organization of defensive behaviours can be attributed to level of threat an animal perceives itself to be in. This theory is one of adaptiveness, as the dominant defence reaction is the behaviour which is most effective in allowing the survival of the animal and the one which is most effective in preventing an increasing level of threat, also known as increasing imminence. The probability of being killed by a predator, known as predatory imminence, is what is responsible for the expressed defensive behaviour. The predatory imminence is dependent on many factors such as the distance from a predator, the potential for escape, and the likelihood of meeting a predator. Three general categories of defensive behaviours, based on increasing predatory imminence, have been identified. These are labelled as pre-encounter, post-encounter, and circa-strike defensive behaviours.

Stages of threat

Pre-encounter

Behaviours exhibited by animals when the threat of a predator is extremely low are known as preferred activity patterns. When the likelihood of predation increases above this point an animal enters into pre-encounter defensive behaviours. These behaviours are used to reduce the probability of having to exhibit avoidance behaviours, which are evoked when a predator is detected. Pre-encounter behaviours can be observed when an animal has left the safety of its residence to perform tasks such as mating or foraging, and occur before a predator has been identified. The animal does not perform these behaviours because of a predator, instead these behaviours are performed or altered depending on the probability of being detected by a predator when executing a task. One well studied example of a pre-encounter behaviour is that of varying meal frequency and size seen in foraging rats. When rats forage they do not typically hoard food, instead they either ingest the food where it was found or retreat to a nearby safe space to consume their meal. Depending on the level of perceived threat, a rat will vary the frequency of foraging and the size of its meal. In laboratory studies investigating this pre-encounter behaviour, density of shocks are often used to represent the risk of predation. A number of shocks are administered daily on a random schedule, the more shocks administered per day the higher the perceived predatory imminence. The shocks only affect the rats when they are outside of their safe nest area, but to get food they have to leave this area and press a lever to dispense their meal. This models the real-life situation of a rat needing to leave the relative safety of its nest to go foraging for food. The common finding is that, as the perceived threat of predation increases, the frequency of foraging decreases but the size of the consumed meal increases. The increased meal size ensures that even though the rat leaves the safe area in search of food less often, it still consumes a relatively constant amount of food each day, regardless of shock density. This organized modulation of foraging behaviour is consistent with a pre-encounter defensive behaviour. The frequency and meal size associated with foraging are reorganized depending on the perceived level of threat but they are not directly influenced by the interaction with a predator.

Post-encounter

Post-encounter defensive behaviours are avoidance behaviours performed when a predator is present and has been detected. When this stage of threat has been reached, behaviours are limited to species-specific defence reactions. These behaviours are commonly freezing, fleeing or threatening. The goal of a post-encounter defensive behaviour is to prevent the predatory imminence from further increasing. The dominant post-encounter defensive behaviour can depend on whether the predator has also detected the prey and how far away the two animals are from one another. Varying levels of predatory imminence, even in post-encounter situations, can impact the expressed defensive behaviour. If the prey has yet to be detected, the goal of the post-encounter behaviour will be to prevent the predator from detecting the prey. If both the predator and prey have detected each other, the goal will be to avoid making contact with the predator. Freezing behaviour in rats is an example of a post-encounter defensive behaviour which has been well studied. Freezing in rats is characterized by sudden, extended immobility, followed by a decreased heart rate and an increased respiration rate. This behaviour is often the dominant post-encounter defence behaviour in rats. In the laboratory setting, post-encounter defensive behaviours can be elicited by pairing a neutral stimulus, such as a light, with an aversive stimulus, such as a shock. The rat will engage in its post-encounter defensive behaviour when it becomes aware of the neutral stimulus, as the stimulus is acting as a predictor for the shock. Since the freezing behaviour is dominant in these situations, it can be assumed that it serves an adaptive function by preventing an additional increase in predatory imminence. Freezing is the dominant post-encounter behaviour because even when there are alternative defensive reactions available, freezing has been observed to be the rat's response the majority of the time. Even when a clear method of escape was made available, rats would freeze instead of fleeing. Freezing can prevent the perceived level of threat from increasing in several ways. If a rat becomes aware of a predator before the predator is aware of the prey, freezing can reduce the likelihood that the prey will be detected. As well, since many predators rely on motion to keep track of their prey, freezing may cause the predator to either lose site of their prey or shift their attention to a more active object.

Circa-strike

When a predator is prepared to strike or has stricken, the prey's behaviours change from post-encounter to circa-strike defensive behaviours. These reactions are employed if the post-encounter behaviours are unsuccessful. A predator making contact with its prey is the highest level of predatory imminence experienced before being killed by a predator. Therefore, the goal of circa-strike behaviours are to survive and escape from the predator during or after contact. These defence behaviours are often more reactive than post-encounter behaviours and are commonly in the form of jumping, vocalizing, striking or biting. If the animal is successful in evading its predator then, when it has found a safe space, it will eventually return to its preferred action patterns. One example of an extremely successful circa-strike defensive behaviour is the evasive leaping of the kangaroo rat. Kangaroo rats are desert dwelling mammals preyed upon by venomous sidewinder rattlesnakes. When rattlesnakes strike, kangaroo rats exhibit a quick and forceful circa-strike behaviour in the form of a leap using their hind legs. This leaping reaction serves several functions both to help escape and prevent envenomation. First, the kangaroo rat is able to jump within around 50ms after perceiving an incoming snake strike. This gives the rat the ability to avoid contact with the predator all together, thus allowing it to escape unharmed. If the leap is unsuccessful in completely avoiding the snake's strike, it still serves the purpose of preventing the fangs from embedding. The jumping action is so forceful that the rat may be able to disrupt the snake's striking movement. Finally, if all else fails and the fangs have made contact with the kangaroo rat, it is still able to employ its evasive leaping behaviour, and with the use of its hind legs, forcefully dislodge the snake's fangs and prevent envenomation. This leaping behaviour serves the function of preventing or escaping contact with a predator immediately before, during, or after an attack, therefore, it is consistent with a circa-strike defensive behaviour.

Draft your article - Predatory imminence continuum

The current predatory imminence continuum article only contains a lead section. While this lead section does introduce the topic adequately the expansion of the topic and various details within the topic is necessary.

• My first heading will be "Species-specific defence behaviours". These behaviours are important to discuss as they underly many of the defence responses seen in the predatory imminence continuum. These behaviours include freezing, fighting, and fleeing and are considered innate.
• My next heading will be "The development of the predatory imminence continuum". This section will discuss the process involved in creating the current theory behind why certain species-specific defensive behaviours are chosen in particular situations. The current theory is one of adaptiveness, where the behaviour chosen depending on threat level is the one which is most effective in allowing the survival of the animal. This however, was not the first theory proposed. The first theory was based on operant conditioning where defence behaviours are organized hierarchically and rearranged through punishment of certain behaviours when they resulted in failure.
• My next heading will be "Stages of Threats" this section will further breakdown the spectrum of predatory imminence. I will have a subheading for each of the three stages of threat: pre-encounter (low threat), post-encounter (moderate threat), and interaction (high-level threat). Under each subheading I will discuss the environment which leads to each threat level, the defence behaviours seen at that stage of threat, and also provide several examples of species demonstrating these behaviours.
• My final heading may be "Brain structures" which will explore the functions of brain structures in regards to defensive behaviour and the predatory imminence continuum. The different threat level responses are activated by varying types of fear. During a moderate threat the fear experienced is usually one of learned fear therefore it would rely on different brain structures than the fear experienced during a physical altercation. This will only be added if time permits and if the article feels like it is lacking. May not have enough information for full section. Will potentially add parts to stages of threat.

Predatory imminence continuum

This article's lead section presents a clear introduction to this topic so I will focus on expanding on several of the points already introduced. First I will talk about the spectrum of predatory imminence and its relation to the defensive behaviour system. Then I will focus on the specific stages of threat and the different defensive behaviours these stages cause. I will also mention species-specific behaviours and provide behavioural examples.

Article evaluation

The Wikipedia article on fixed action patterns, which is a part of the WikiProject Animals, has been given a “Start-Class” rating. Upon evaluation, the reasoning behind this rating becomes apparent. While the majority of information presented in this article is relevant to fixed action patterns and is presented in a neutral way, there is quite a lot of notable information missing surrounding the subject. Although the article does define the meaning of fixed action patterns and gives several examples of such behaviours, it does not go into the term’s history nor the criteria necessary for a behaviour to be considered a fixed action pattern. In class we spent time introducing the history of fixed action patterns and highlighted the six characteristics created by Konrad Lorenz that distinguish these behaviours. The article simply gives a weak definition and a brief look at terminology surrounding the principle before getting into several examples. As well, the article is not very well referenced. There are only nine references given for the entire article. While these references are reliable since the majority are from peer reviewed journals and textbooks and the links given are functioning, there are several statements throughout the article that are not sourced. For example, the entire section on Niko Tinbergen’s male stickleback fixed action pattern studies has no sources at all. As well, many of the statements the article makes surrounding the significance and exploitation of fixed action patterns are not sourced. Additionally, three out of the nine sources are for a one sentence example of lizard’s and snake’s use of a fixed action pattern. The talk page of this article presents several concerns and discussions. One post on the talk page explains their reasoning for removing a complete section on criticism of the term as it was based on only one source from an unreliable website. Another few posts on the talk page ask for clarification on the definition of a fixed action pattern and discuss whether or not the examples given are appropriate. It is hard to tell whether viewpoints in this article are overrepresented or underrepresented as there are not many viewpoints given. The bulk of the article is unsourced, vague examples, not actual valuable information on fixed action patterns or the varying viewpoints which surround this principle. Fixed action patterns are a very important topic within the fields of animal behaviour and classical ethology. In the future, this Wikipedia article should be reviewed so that more detailed information on the subject can be added and all statements made can be properly sourced.

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