Wednesday, 7 June 2017

Mounting Behaviour

All animals have a evolved a repertoire of innate behaviours and mounting behaviour is wide spread in species sufficient to designate it as an instinctive  behaviour.  Instinct or innate behaviour is the inherent inclination of a living  organism towards a particular complex behaviour. Which is an  unlearned, inborn behaviour characteristic of a species, this includes unconditioned autonomic responses and valuations of biological importance (Falk 1997) .

 Various studies have identified that naïve and isolates  and  animals which lack a sense of smell will also mount which suggests that the nervous system is wired to effect social and reproductive behaviours (Leopald et al 2002). When animals are closely associated for some period of time, they may occasionally briefly mount one another without thrusting, erection, or other signs of emotional arousal, and such behaviour has been interpreted as an expression of social affect or friendship rather than a sexual response (Beach 1976).

Mounting behaviour which occurs normally in mammals is not associated with any excess of heterologous gonadal hormones or deficiency of homologous ones. It is facilitated in both sexes by the presence of normal amounts of homologous hormones , but males react more strongly than females to androgen and the responsiveness of feminine mechanisms to estrogen is much greater in females than in males (Mason 2006). In male mammals, testosterone is released in a pulsatile fashion in which high “pulsatile” levels are periodically superimposed upon low “basal” levels. Two different types of pulsatile release occur: spontaneous and reflexive. Spontaneous release typically occurs multiple times per day in response to endogenous homeostatic conditions, which accounts for much of a male's circulating testosterone, and is centrally important in organizing and maintaining many aspects of male reproductive physiology and behaviour. At times of anxiety or emotional arousal there is a reflexive testosterone release in both males and females, testosterone has a fast action in affecting anxiety, reward learning  and analgesia. Testosterone has an anxiolytic effect, and an analgesic effect. At times of high arousal or anxiety the release of testosterone can lead  to an instinctive  response, the male animal may mount other animals or inanimate objects   as a result (Nyby 2008;van Honk et al 2005).

All mammals nose, lick, mouth, or otherwise manipulate their own genitals under various conditions. These responses constitute autogenital grooming. Males of many species also nose, sniff, or lick the genitals of another individual, but this investigative behavior is not accompanied by any indication of sexual excitement (Beach 1976).  

Male masturbation can be inferred when a  male animal repeatedly achieves erection and ejaculation by stimulating the penis or some other erogenous zone. An example of masturbation was observed  in a  male cat living in a laboratory cage, he  developed a technique of mounting his food dish and making copulatory movements that led to penile erection and ejaculation, this  behaviour usually occurred when the male could see other cats copulating on the floor below his cage. Rubbing the erect penis against inanimate objects or substrates is  frequently practiced by male porcupines and porpoises, and one captive monkey habitually employed this form of stimulation to achieve ejaculation. Captive monkeys and chimpanzees masturbate manually and orally, often inducing ejaculation in the process. This activity has sometimes been interpreted as a completely abnormal form of sexual outlet produced by heterosexual deprivation and other artificial conditions incident to a captive existence. However socially dominant male monkeys will copulate with an estrous female on one occasion and masturbate to ejaculation after a brief pause,  even though the female is still available. Therefore under these conditions, self-stimulation can scarcely be classified as a vicarious activity, or as a response to deprivation and sexual frustration. Self-stimulation to the point of ejaculation appears to be rare in feral males, but it is common in captivity. It is possible that the difference is owed to the lack of freedom and variety which characterizes a confined existence. Therefore, masturbation is in no sense abnormal, but its frequency may be affected by a male's living conditions. However,  sexual frustration may lead to masturbation. In one experiment, male rhesus monkeys were confronted with spayed females which had been rendered sexually attractive but not sexually receptive. When exposed to such females, some males masturbated to ejaculation (Beach 1976). Therefore it is possible that  male dogs may masturbate to ejaculation when he smells a neighbourhood bitch on heat but cannot access her. 

References

Beach, F. (1976). Cross-species comparisons and the human heritage. Archives of Sexual Behavior, 5(5), pp.469-485.
Chevalier-Skolnikoff, S. (1974). Male-Female, Female-Female, and Male-Male sexual behavior in the stumptail monkey, with special attention to the female orgasm. Archives of Sexual Behavior, 3(2), pp.95-116
Falk, J. (1977). The origin and functions of adjunctive behavior. Animal Learning & Behavior, 5(4), pp.325-335.
Leypold, B., Yu, C., Leinders-Zufall, T., Kim, M., Zufall, F. and Axel, R. (2002). Altered sexual and social behaviors in trp2 mutant mice. Proceedings of the National Academy of Sciences, 99(9), pp.6376-6381.
Mason, G. and Rushen, J. (2006). Stereotypic animal behaviour. 1st ed. Wallingford: CABI
Nyby, J. (2008). Reflexive testosterone release: A model system for studying the nongenomic effects of testosterone upon male behavior. Frontiers in Neuroendocrinology, 29(2), pp.199-210
van Honk, J. Peper, J.  Schutter, D. (2005). Testosterone reduces unconscious fear but not consciously experienced anxiety. Biological Psychiatry, 58, 218–225.

Sunday, 3 May 2015

Aggression in dogs

Dogs fulfil numerous roles within the human community. To early man, the main purpose for owning dogs is thought to have been their hunting prowess and the protection they conferred against other predators (Morris 2008). To date dogs continue to fulfil these roles  (Udell et al 2014), and for some they are  companions, confidants, and a member of the family (Overall 2013),  for others the dog is a work partner, an assistant, a status symbol, perhaps even a weapon (Morris 2008). These diverse, roles fulfilled by dogs, result in differing expectations, differing attitudes and differing husbandry, resulting with differing normative beliefs of aggression (Anderson et al 2003).

Aggression is defined as hostile behaviour, which is intended to harm (Davenport 1996; Gross et al 2004). Aggression in dogs includes: growling, snarling, snapping, biting, barking, or lunging (Abrantes 2005).

There are many different classifications of aggression within the literature and the definitions vary with each author, add on top the different classifications of aggressive behaviour in behavioural modification in dogs where some do not agree with each other, or they look at it from a different angle, or the labels change with each author. The intention is to select as much relevant information relating to aggression to ensure that most motivations are covered.

There are two types of aggression in mammalian species: predatory aggression (sometimes called quiet biting attack) and affective aggression. Predatory aggression can be deemed as an unemotional because behaviour is driven by food procurement at some level; inflicted on a different species; and is a self rewarding behaviour (Armony 2013). On the other hand affective aggression is driven by emotions and for the most part these are defensive behaviours, although if defensive behaviours are not working they can result in offensive behaviour. (Clutton-Brock et al 2013; Székely et al 2010).

Lorenz defines aggression as the "driving power,” an instinct toward the preservation of life, all mammals do not learn to be aggressive. It is in their nature. (Lorenz, K. 1963;2005).

Scott (1966) pointed out that:
“Aggression is a poor scientific term, and chiefly functions as a convenient handle to relate phenomena described in more objective terms to practical human problems. What we are really concerned with is agonistic behavior, a behavioral system composed of behavior patterns having the common function of adaptation to situations involving physical conflict between members of the same species. We cannot analyze fighting behavior without also studying the alternate behavior patterns of escape, threat, "freezing", defensive posture, dominance and subordination, etc” (page 683;Scott, 1966).

From an ethological point of view, aggression in dogs can be characterised as an intensive, emotional, expressive response oriented against the object of the dog's frustration. Aggression is a phylogenetic, pre-domestication, instinctive state of animals of self-preservation importance. Dogs are carnivorous animals and as such they exhibit various forms of aggression ; interspecific aggression is oriented against different animal species, which includes offensive and defensive elements. Its most frequent form is self-protective aggression (Kottferovi et al 2008; Ryan 2013). Predatory aggression is a part of feeding behaviour of carnivores, although domestication in dogs has resulted in relaxation of the predatory sequence, some dogs exhibit behaviour which is part of predatory behaviour, e.g. visual exploration, hunting small animals and occasionally humans. The stimuli that usually induce this behaviour include olfactory and visual (Udell et al 2014; Kottferovi et al 2008) . As behaviour is governed by rules that associate stimuli with responses and outcomes (Zhang et al 2013), treating predatory aggression is difficult, it is an instinctive behaviour, therefore good management is essential whilst behavioural modification is being commenced: use of leads, and muzzling if there is a risk of injury. Desensitization with counter-conditioning can be effective in some instances to change a dogs' perception of falsely identified prey. Satisfying the needs of the dog is paramount, I have used David Ryan’s ( 2013) two ball method successfully with dogs enjoy the pursuit and grab part of the predatory sequence, and I direct owners to read this which explains it simply http://www.dog-secrets.co.uk/how-do-i-stop-my-dog- chasing/ however owners need to be motivated and comply effectively for this to help.

Defence systems in the mammalian brain are aimed at making immediate responses to threatening stimuli; and a behavioural inhibition system responsible for the suppression of behaviour that could enhance danger, these conflicting systems work in concert and engage distinct hierarchic circuits depending on increasing demand for cognitive processing (Stein et al 2010). Defensive aggression is related to fear, pain or threat of punishment and includes defensive or submissive signals before or during the aggression (Overall 2013).

Territorial aggression:
Territoriality: is defined as behaviour by which an organism lays claim to an area and defends it against intruders, which is generally a fixed territory; personal territory on the other hand moves with the individual and can expand or contract depending on the circumstances. Territoriality is a basic behavioural system and is characteristic of all living organisms (Hall 1969,1990). Some breeds of dogs; and some individual dogs, regardless of breed are more territorial than others (Beaver 2009). Territorial behaviour usually appears as puppies mature into adolescence or adulthood, at one to three years of age (Beaver 2009; Overall 2013).

Territoriality behaviour can be a way to insure they have a safe place, because a dog with a territory can develop an inventory of reflex responses when perceived danger strikes, rather than having to think about the environment they are in (Székely et al 2010). So a dog with who is fearful might well defend a place he feels safe in.

Space requirements are influenced by the environment. One of the most fundamental parts of territoriality is proper spacing which protects against over exploitation of that part of the environment (Hall 1969;1990). So a dog that is displaying territorial aggression it could be because they are living in over crowded conditions, or that there is a high densitiy of dogs/humans/other animals living in their neighbourhood. Hediger (1965) inferred the need for proper spacing between individuals, were species specific, and environmental specific. Flight distance and critical distance are used when two difference species meet, where as personal distance and social distance can be observed during interaction between members of the same species (Hall 1969;1990). And for some dogs the mechanism of personal distance and social distance only applies to members of their immediate household, all other humans and animals are treated as a different species even if they have had effective socialisation (Hall 1969,1990; Coppinger et al 1996).

Hediger (1965) he also identified a variation in difference between species, some animals are contact species and some are non contact animals. Species like the Emperor Penguin huddle together and require physical contact with each other. Although dogs fall under the non contact species they begin their life in the contact phase, which is temporary, the dam will naturally reduce contact times even if man did not intervene. Some dogs will continue to seek out close contact with housemates other dogs will ensure regular spacing is adhered too (Hall 1969,1990; Hediger 1965). As dogs are deemed to be non contact animals they are likely to be more vulnerable to stresses of overcrowding.

How to help a dog who presents with territory aggression is he in over crowded conditions, are there lots of people and animals in the neighbourhood, if that is the case it is going to be more at looking at ways of ensuring that the dog has more space or alter the environment so that he has his own personal space that is not likely to be encroached upon by others, putting up double fences so their is dead space between his territory and passerbys. Desensitization and counter conditioning will help. Alter walks if possible going to wide open places, or places where there is less foot-fall, preventing the behaviour from occurring is the best way. It also ensures that trigger stacking is minimised

Protective aggression occurs as a result of territorial behaviour regarding their owners, handlers and the children of the family or other animals that they have developed social bonds with (McGrew 1983). Livestock Guardian breeds like the Anatolian, German Shepherd and Maremma’s dogs etc have a long history of being bred for their protective role. There is a behavioural distinction between protective aggression and territorial aggression, Instead of defending a geographical location, these dogs actually defend the space immediately around them, their personal space. These dogs are generally very attentive to the individual/s being protected, and consequently those individuals become included within this defended area resulting in the dog being equally protective anywhere it encounters a predator or a falsely identified predator, like the mother in law, or the friendly neighbour. (Mcgrew 1983). Protective behaviour does not occur in puppies but it can develop as early as 9 months old (Mcgrew 1983) or as late as 18 months old (Coppinger et al 1988).

Maternal aggression is a form of protective aggression, but it is stimulated by hormonal factors. At the end of pregnancy or Dioestrus, there is a sudden drop in progesterone and an increase in Prolactin and a reduction in serotonin levels (Dahl et al 2010). However hormones do not cause or inhibit behaviour by themselves, rather they affect the sensitivity of the neural pathways involved in protective aggression.

Bitches are unique because they are designed to be pregnant after every heat cycle, the whole period is a pre-programmed finite lifespan and is not subject to the regression that you can be observed in other mammals like humans, horses, cattle etc when they do not conceive (Razzaque et al 2008). So the unsuspecting owner two months after the heat cycle cannot understand how their lovely girl has turned in to an aggressive being. Or how their lovely girl not long after being spayed turns aggressive. So whether the bitch has just had a litter, or has just been spayed, or two months after heat cycle their is a likely hood that they can develop maternal aggression. The best thing is to prevent the dog from displaying aggression, leave them in peace, do not encroach, do not remove the very things they are protecting (in some circumstances removal of the things they are protecting can help for instance the dog is taken out for a walk, someone removed those items the dog does not have the visual stimuli to continue behaviour when they return. Remove as much stress as possible within the environment. This is a short term hormonal behaviour, what you don’t want the bitch to  do is to continually practice protective behaviour and consequently learn in the absence of the stimulating hormones protective behaviour because it works. If the bitch is displaying maternal aggression in the absence of pregnancy it might be worth discussing a suitable time to spay, this would usually be during Pro-oestrus, where the effects of spaying will have minimal, hormonal impact.

Paternal Aggression this is similar to maternal aggression however unrelated males and bitches can also display this aggression, protecting a young dog from others whether that is dogs or humans or other animals. This can be seen when a new puppy comes into the household and forms social bonds with a resident adult dog.

Possessive aggression develops when the dog appropriates some objects, for example a bone, food, a toy. Resource guarding is a normal behaviour if humans are constantly removing resources or not providing enough resources resource guarding is further enhanced. Hunting dogs like the Airdale Terrier are bred to protect kills from others, so are more likely to develop resource guarding behaviour. This behaviour can be seen in all ages from pups to adulthood (Beaver 2009; Overall 2013).

Social Aggression Social behaviour is driven by two antagonistic motivational incentives affiliation and competition. Competition theory was a fundamental part of Darwinian theory. Driven by their selfish gene, individuals seek to maximise their genetic contribution to the next generation, through affiliation and competing with others (Clutton-Brook et al 2013).

Competition requires aggression at some level. Lorenz believes that species develop a number of mechanisms for redirecting or inhibiting intraspecific aggression. Ritualized fighting in which no animal is really hurt which dissipates aggression, and submissive behaviour can block it. Dogs with their ability to kill easily, have developed rituals and inhibitions limiting and formalizing intraspecific aggression (Lorenz, K. 1963;2005). Competitive male displays are widespread in social animals and are used to attract breeding partners and to repel rivals. Visual; vocal ;and olfactory displays are often combined and frequently reflect the signallers hormonal status; condition; and physical strength.

Sex related aggression During development the brain is the target site for steroid hormones, during the neonatal period both males and females are exposed to variant amounts of testosterone, a female between two male neonates is going to be exposed to more testosterone than a female between two females (Trainor et al 2006). Exposing females to testicular hormones masculinises components of the central nervous system(CNS). Prenatal chemical castration or surgical castration of the male allows the development of a more female-like CNS. In mammals, the sexual differentiation of the CNS has a significant role in shaping sexual preference and other reproductive activities. In addition, it influences food intake and body weight, territorial marking and aggressive behaviour, learning strategies, and play behaviour (Birger et al 2003).

Testosterone can be deemed a pro-hormone, because it is a precursor to which when converted into 5-alpha-dihydrotestosterone (5α-DHT) acts on androgen receptors or when converted into estradiol by the enzyme aromatase, acts on oestrogen receptors There is overwhelming evidence that most of the effects of testosterone in mediating aggression occur after aromatization. For example, in rats testosterone induced aggression is concurrent with an elevated level of aromatization and nuclear oestrogenic receptor activity in the hypothalamic/preoptic area. Treatment with an aromatase inhibitor blocked this aggression and lowered nuclear activated oestrogen receptors Furthermore the intensity of aggressive behaviour was directly correlated with the aromatase activity in the posterior hypothalamus (Trainor et al 2006). Therefore we can surmise that the hormones do not cause or inhibit behaviour by themselves, rather they affect the sensitivity of the neural pathways involved in different behaviours. For instance testosterone sensitizes the responsiveness for stimuli which are related to aggression so they need less visual stimuli from a rival or from a threat which will start the aggressive motor programmes when testosterone levels are high. Furthermore testosterone strengthens the motor output so aggressive behaviour is performed at higher intensity (Jenson 2007). Prolactin and low levels of serotonin sensitizes the responsiveness for stimuli which are related to defence behaviour (Dahl et al 2010).

It has been identified that the interaction between low serotonin and high testosterone levels in the central nervous system has a significant effect on the neural mechanisms involved in the expression of aggressive behaviour. It seems that testosterone modulates serotonergic receptor activity in a way that directly affects aggression, fear and anxiety. Low serotonin exhibited high rates of aggression, high testosterone further augmented rates and intensity of aggression in subjects with low serotonin. Therefore testosterone concentrations and aromatase activity appear to be positively correlated with overall aggressiveness, but not with measures of impulsivity. it is the low levels of serotonin in combination with high levels of testosterone or aromatase which increase intensity of aggressive response. So depending on the individual dog, one dog might be more competitive than another dog, one dog might be more impulsive than another dog (Birger et al 2003).

Fights over access to territories resources and social status occur frequently in the males of most mammalian species. Independent males that compete differences in age, size, weight , and stamina play an important part of the outcome (Clutton-Brook et al 2013).

In most groups of dogs frequency and intensity of aggression in females is lower than in males, They use the same combination of visual, olfactory and vocal displays, and quality of signals, display their age, size and condition. Females will defend their territory from intruders and can be as aggressive as males and can lead to serious injury and death however female aggressive displays are usually directed at rivals. (Sherman et al 1996; Wrubel et al 2011). This type of aggression can begin around the time of sexual maturity to adulthood.

Fear aggression is a normal behaviour it is part of nature and affects a wide spectrum of causes of aggression: Stimuli that cause fear may be related to predators, intensive physical environmental stimuli and warning signals. Fear can sometimes be observed due to constitutional factors or temperament or in other cases because of insufficient experience with fearful stimuli (Overall 2013; Beaver 2010; Landsberg et al 2013). According to Hediger (1965) critical distance encompasses a narrow zone separating flight distance from attack distance, a dog might flee an approaching person/animal until it meets a barrier, if the person/animal continues to approach and the dog is cornered the dog is likely to attack. Effective socialisation will decrease critical distances, however some dogs despite good practices will be predisposed to maintain a distance (Hall 1969,1990; Coppinger et al 1996).

Dominance aggression, has definitions varying with nearly every A dog displaying aggression can display it self-confidently (dominant behaviour) however a dog that needs to control regardless of context is neither adaptive or normal. It is important not to blur the lines between normal behaviour i.e. the dog perceives he or his resources are at risk which is adaptive, or abnormal behaviour where there is no risk to the aggressor (Overall 2005). In addition a dog that has to control everything is not an evolutionary stable strategy (Dawkins 2006) and as a result true dominance aggression is rare (Overall 2013). Overall (2013) describes dominance aggression as a conflict aggression and is characterised by consistent atypical, out-of-context aggressive behaviour directed towards people. These behaviours include growling, snapping, and biting. Bites are usually not preceded by a vocal warning.

Frustration- elicited aggression The frustration-aggression hypothesis was proposed by Dollard et al (1939). According to this view, frustration, which is defined as "the state that emerges when circumstances interfere with a goal response," this can apply to the frustration that a dog feels when he can see but not reach his goal. This frustration occurs because there is a window, door, fence or a lead which prevents him achieving his goal, this often leads to aggression, fence fighting, lunging etc. This type of aggression occurs in both puppies and adults (Overall 2013 Beaver 2009 Landsberg et al 2013).

Redirected aggression is a lot like frustration-elicited aggression withthe exception that the dog need not be frustrated. Redirected aggression occurs when a dog is aroused by or displays aggression toward a person or animal, and someone or something else interferes with it. The dog redirects the aggression from the source that triggered it to the person or animal who has interfered with it . This is why people are often bitten when they try to break up dog fights. Another example is when two dogs are barking at someone from behind a fence, and one will turn and attack the other. Male and female dogs are equally prone to redirected aggression, and this type of aggression occurs in both puppies and adults (Overall 2013 Beaver 2009 Landsberg et al 2013).

Impulsive reactive aggression, Impulsivity is a trait related to inhibitory control Impulsive individuals show a decreased ability to tolerate delay of reinforcement (Lyndsay 2000). Impulsive aggression, is a lot like frustration-elicited aggression with the exception that the dog is being reactive; lacks self control and with the inability to control associated emotions (Abrantes 2005). Impulsive behaviour has been linked to decreased levels of serotonin and dopamine in a number of species. In domestic dogs, impulsivity is implicated in problem behaviours that result from a lack of self control this type of aggression occurs in both puppies and adults (van Rooney et al 2014).

Pain-Elicited Aggression An otherwise gentle, friendly dog can behave aggressively when in pain. A dog with a painful condition or an infection might bite with little warning, even if the reason they are being touched is to help the dog. Use of certain pieces of training equipment, such as the choke/prong / shock collars can inflict pain on a dog and prompt a pain-elicited bite to the handler. Male and female dogs are equally prone to pain-elicited aggression, and this type of aggression can occur in both puppies and adults (Overall 2013 Beaver 2009 Landsberg et al 2013).Relief of pain is important for reduction of misdirected aggression (Jenson 2007).

Loss of Rapid Eye Movement (REM) atonia leads to REM Sleep Behavioural Disorder (RBD) which causes dream enactment. If the dog is having combative dreams it can lead to vigorous and violent behaviours resulting in attacking the nearest object, person or animal. Most patients do not have structural lesions of the pons, but the disorder is thought to arise from an imbalance of neuronal regulation in this area,which is responsible for regulating REM and nREM sleep. The semi- purposeful behaviour with confusion may be impossible to distinguish from seizures or postictal behaviour. Unlike most partial seizures, REM behaviour disorder will be restricted to sleep. (Kaplin et al 2005: Brown et al 2012).

Medical problems should be considered for aggression : implicated in aggression are Hypothyroidism, epilepsy, metabolic disorders, Rabies ; Distemper, neoplasia, Hydrocephaly, hepatic disorders, endocrine disorders, poisoning amongst others (Kotteforovi et al 2008;Beaver 2009; Overall 2013; Landsberg et al 2013).

Drugs Certain medications could give rise to aggression. Selective serotonin re uptake inhibitor ( Prozac fluoxetine etc)   can result in paradoxical reactions resulting in increased aggression
towards individuals and so can Benzodiazepines, Barbituates, Chlorpromazine etc (Maddison et al 2008).

Nuture A deprived socio-cultural environment where both humans and their dogs are exposed to the same suboptimal conditions eg; homelessness (Gross et al 2004)

Punishment based training is more likely to result in a dog aggressing towards humans (Casey et al 2014)

Inconsistency in ownership style could also result in a dog aggressing towards humans. For example being allowed on the sofa one minute, being shouted at to get down next. Having food or valuable resources taken away constantly could result in a dog aggressing towards humans.

Constantly being disturbed whilst trying to sleep on his bed could result in the dog aggressing towards the person who disturbed him.

 In appropriate socialisation and insufficient exposure to humans and or other species. If the dam is aggressive towards humans the chances are the puppy will also be aggressive towards humans (Overall 2013; Beaver
2009).

Learning is also an important factor in the development of aggressive behaviour. The dog learns to be aggressive in order to achieve a goal, following the principal of instrumental conditioning, daily encounter s of the dog with people, can produce an important impact in the development of aggressiveness (lansdsberg et al 2013; Overall 2013).

Heritability of aggressive traits: Humans have selected behaviours in dogs for hundreds of years, and exploited the predatory sequence, Schavbert (2006) suggest that breeding selection for dog show use is positively correlated with social and non-social fearfulness, and negatively with playfulness, curiosity in potentially threatening situations and aggressiveness, whereas selection for working dog use is positively correlated with playfulness and aggressiveness. Furthermore, correlation analyses show that popular breeds have higher sociability and playfulness. However if you look at how the pedigree breed standard was developed for the Staffordshire Bullterrier. The full wild canine predatory sequence is: orient>eye>stalk>chase>grab bite>kill bite>dissect>consume (Udell 2014). Breeders of the Staffordshire bull terrier selected for the hypertrophied grab bite, which gave them their distinctive jaw and facial structure.

“The extraordinary features of a bully breed are reminiscent of the bull-baiting tactics and strategies, it made the Bulldog deadly effective in killing a bull. The quagmire of dogs, fighting bulls was intense and the way dogs used to grab the bulls nose needed some special features that could help in sticking to the bull’s nose to kill him. The Bulldogs were trained to crouch low to the ground to shield their bodies from the bull’s horns when it charged. The shoulders were placed on the outside of the body, the hind quarters of the Bulldog are not as well developed as the fore quarters, allowing the dog to be shaken violently without suffering any spinal injuries, The head of the Bulldog, like his body is also the part of the fighting strategies that men wanted to develop. The short jaws, lower jawbone is longer than the upper jawbone which allows the dog to hang onto whatever it wants with a surprising tenaciousness. The grip is very strong, and the structure of the jaws, surprisingly enables the dog to move jaws in a manner that it can hang onto the throat of the bull and shred the flesh, sinew, and muscle until it reaches the jugular. The Bulldog continues to hang on until the bull is brought down; bleeding to death. The skin folds on the face allow the Bull’s blood to flow down the dogs face under its chin rather than into its eyes. The Bulldog has a short snout on the upwards face to allow breathing, while retaining its grip on the bull’s nose. The bull is, in the meantime suffocating due to the loose jowls of the bulldog blocking the passage of air” (Page 15 Jenkins et al 1997).

The breed standard for the Staffordshire Bullterrier was developed to prevent the breed from ‘softening’ (Morris 2008; Jenkings et al 1997; Catherine 2011), If humans select for temperament the destabilizing effect of selection will result in changes to the phenotype and therefore not match the profile of the breed standard (Baelev 1979). Baelev suggest that the interface of physical and behavioural conformation means it is not possible to breed out aggressive behaviour of fighting dogs while retaining their shape and appearance. However it is clear that not all Staffordshire bullterriers carry the aggressive trait to make good fighting dogs (Jenkins 1997). Canine experts generally deny the existence that there is an automatic link between breed and aggression, but they point towards the existence of particular aggressive breeding lines (Serpell et al 2014), but even perfect breeding lines produce 'misfit' puppies (Coppinger et al 1988).

When it comes to assessing aggression It is not necessary to  see a dog doing the aggressing, and consequently giving the dog more practice at the behaviour. So roles should be more  of a  preventative model and the use of a good assessment tool that can identify triggers possibly use of video recordings. For example if the person referred says their dogs bark lunge or attack people coming into the house, but are fine out of the house, behavioural modification programme would begin away  from the environment that could trigger an aggressive response begin getting the skills in place before working in that environment.

It is clear from the above terms aggression, they can be applied to a variety of causes. Human directed aggression is not correlated with dog directed aggression but the cause of either human directed aggression or dog directed aggression or any other type of aggression can be due to nature , physiological disorders , neurological disorders, medical disorders, drugs , nurture, environment and any of the multitude of categories stated above. Therefore treatment of aggression includes identify the trigger, Eliminate all triggers, if you cant eliminate triggers manage it through well placed barriers, careful use of space and distance, good practice of resources, prevent from practicing the behaviour; and use desensitisation and counter-conditioning, satisfy the needs of the dog, and confidence building exercises. Occasionally re-homing is the best option particularly if it is a sex related aggression or owners are not able to manage and continue behavioural modification. And occasionally euthanasia might be the best option for the welfare of the dog and for safety of others.

We can use a functional definition of aggression but this adds our interpretation of the function of the animal’s so-called intent and as a result it focuses on the individual, but it forgets about the survival and fitness of the species and others. In addition is the interpretation actually related to the aim and function of the dogs behaviour, for example does chase behaviour have anything to do with predatory aggression or territorial, If it is unrelated to the aim and function, then our functional classification makes no sense. Many forms of behaviour contain representations of a desired end-point (Nelson 2006). If this is true, then we have to seriously consider cognition in the dog.

References and further reading

Abrantes,R. (2005) Evolution of Canine Social Behavior. 2nd edition. Wakan tanka Publishers, USA

Anderson,C. Rowell-heusmann, L. (2003) human aggression a social cognitive view.
Chapter 14 in Hogg, M. Cooper, j.( Eds)( 2003)The SAGE Handbook of Social Psychology Sage publications inc.

Belyaev, D. 1979. Destablising selection as a factor in domestication Journal of Heredity (1979) Vol 70 No 301 pp 301-308.

Beaver, B  (2009) Canine Behavior: Insights and Answers . 2nd edition.  Saunders, an imprint of Elsevier Inc.

Birger, M. Swartz,M. Cohen, D. Alesh,Y. Et al (2003) Aggression: The Testosterone-Serotonin Link Review article. Israel Medical Association Journal Vol 5 . Sept 2003 pp  653-658

Bradshaw, J.W.S. & Nott, H.M.R. 1995 Social and Communication Behaviour of Companion Dogs. In; Serpell, J.A. (Ed) The Domestic Dog; Its Evolution, Behaviour and Interactions With People. Cambridge University Press.

Brown, R.   Basheer, R.  McKenna. J. ,  Strecker, R , . McCarley, R, (2012) Control of Sleep and Wakefulness Physiological Reviews Published 1 July 2012 Vol. 92no. 1087 1187DOI: 10.1152/physrev.00032.2011. Avaliable from: http://physrev.physiology.org/content/92/3/1087

Casey, R. Loftus, B.  Bolster, C. Richards, G. Blackwell, E. (2014)  Human directed aggression in domestic dogs (Canis familiaris): Occurrence in different contexts and risk factors Applied Animal Behaviour Science vol:152 pp 52-63.

Catherine, M. (2011) “Bulldog breeds” Website of Bull dog information library. Catherine Marion-deluca 2003-2011 accessed (06/03/2015. http://bulldogbreeds.bulldoginformation.com/different-bulldog-breeds-types.html

Clutton-Brock, T. Huchard, E. (2013) Social competition and selection in males and females Philosophical Transactions of the Royal Society of Biological Science 5 December 2013 vol. 368 no. 1631. Available from:
Coppinger, R, coppinger, L. Langeloh, G. Gettler, Lorenz,J. (1988)  A Decade Of Use Of Livestock Guarding Dogs Proceedings of the Thirteenth Vertebrate Pest Conference paper 43 3-1-1988 avaliable from: http://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1042&cont

Coppinger, R. Coppinger, L. 2001. Dogs: A startling new understanding of canine origin, behavior & evolution. 1st edn. New York: Scribner

Coppinger, R., Coppinger, L., 1996. Biologic bases of behavior of domestic dog breeds. In: Voith, V.L., Borchelt, P.L. (Eds.), Readings in Companion Animal Behavior. Veterinary Learning Systems, Trenton, pp. 9–18.

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Wednesday, 29 April 2015

Inappropriate elimination in dogs

“Elimination (urination and defecation) is one of the fundamental biological activities of any species” (Wagner et al 2014). Maslow's(1943) hierarchy of needs, places elimination as a basic physiological need and a requirement for survival . Therefore a dog has a drive to urinate. However there is a whole lot going on in the dispersal behaviour of urine over and above the simple elimination of waste.
New born puppies urination depends on an exteroceptive somato- bladder reflex mechanism that is triggered when the mother licks the genital or perineal regional of the pup, this reflex diminishes as voluntary neural pathways develop between 7-12 weeks old ((animals with spinal injuries can be manipulated to urinate through this exteroceptive- somato reflex mechanism) (Fowler 2008). So it should be no surprise when a puppy under 3 months old eliminates shortly after someone picks them up because they have accidently stimulated this reflex.

Mature bladder function can be represented as a cycle, made up by a series of phases: Filling; desire to void; postponement; initiation of sphincter relaxation; and bladder contraction; and maintenance of both these until the bladder is empty/filling. To make use of the postponement phase three abilities are required:
to appreciate what is an acceptable micturition site;
to be able to get there;
and to adopt a micturition posture.
Each phase and each requirement in the postponement phase has its own disorders which may occur alone or in various combinations (Yeates 1974).

The desire to micturate and the symptom of urgency is produced by an increase in tension in the bladder wall, attenuated by bladder contractions. The contraction waves and resulting sensations are easily affected in either direction by cortical activities. Micturition is normally initiated by voluntary removal of its inhibition, then urination results from inhibition of the tone of the striated muscles in the pelvic floor. Therefore maturation of the control of micturition consists of reduction of the micturition reflex; development of consciousness bladder distension; and the development of conscious ability to postpone or to initiate micturition as required (Yeates 1974).

So it is important to think about the developmental stage of the dog understand that there is individual variation in bladder control age, and also to take note that there is variation in bladder size even between dogs of same breed and same age( Mantis 2008).

Over the last century scientists have studied elimination behaviour of various species including domestic dogs. Ross (1950) studied elimination behaviour of dogs under the specific conditions living conditions in laboratory colonies, where dogs existed in living-eating- sleeping quarters. It was identified that adult dogs preferred to eliminate away from the sleeping and eating areas, and selected a restricted area, and eliminated in that particular area daily. If house broken, they
preferred to eliminate outside, despite the requirement for more effort to be made to go outside. If dogs were disturbed by strange noises or by continued presence of observer the restricted behaviour of elimination was not continued.

Preferred elimination away from bed and food areas appears to be normal species specific behaviour which is not just confined to domestic dogs, other animals also have preferred locations away from food and bed area (Jenson 2002).

Puppies ceased eliminating in bed area around five weeks old, but do not develop selection and restricted elimination behaviour for another few weeks (Ross 1950; Wagner et al 2014). This is the time where voluntary neural pathways are being developed and where elimination behaviour begins to be learnt.

Identifying the site of elimination might well identify the reasons for inappropriate elimination, if the dog is eliminating in one area it is likely his preferred restricted site a sign that house training might have gone wrong, the dog has learnt indoor location and substrate preference (Beaver 2009). However Ross 1950 and Wagner et al 2014 identified that if a dog eliminates in one area the likely hood of another dog or the same dog eliminating in that area is high. So the elimination might have started as an accident, but if the smell is still in that area the dog is more likely to eliminate in that area again and as a result learns his preferred elimination site.

Often puppy owners use puppy pads inside the house, the pup learns to urinate on the puppy pads. Owners remove puppy pads without any further training, and the dog eliminates on the carpet instead because he has learnt that he can urinate in the house, and the
carpet feels just the same as a puppy pad to the dog.

If the dog has no preferred site and is urinating here there and anywhere the dog has either not developed preferred elimination areas, or is anxious or is continuously being disturbed by someone or something.

Anxiety can suppress bladder contraction waves or conversely exaggerate bladder contraction waves (Yeates 1974).

Weber (1939) suggested that providing a restricted crate with no more room than to lie down would ensure that the dog would inhibit defecation and urination in the cramped conditions, and many people still use this method of house training today. However leaving a dog in his crate for long periods without providing the chance to eliminate in appropriate areas, results in the overriding drive to urinate which is counterproductive because the dog learns that he has no choice but to
urinate in his crate area.

Physiological reasons (this list is not exhaustive):
The dog is in pain
Obesity is a common risk factor for incontinence in dogs.

Neutering is also one of the primary risk factors for incontinence, however, most animals do not develop any medical disorders as a result of neutering; complications are uncommon. If there is incontinence related to neutering, it will generally be temporary, as the dog learns to control its urinary muscles again during the recovery process (De Bleser 2011; Beauvais 2012) .

Disruption of the nerves around the bladder
Lesions in the brain
Damage to dogs nerves or spinal cord that innervate the bladder, trauma or neoplasia
 Overactive bladder syndrome
 Urinary tract infections
Chronic inflammatory disease
Pressure on the bladder caused by a mass
Underdevelopment of the bladder or other birth defects
Decreased sphincter control often seen in older females
This list as stated is not exhaustive but it shows how important it is to ensure that there is not a physiological reason for inappropriate elimination (Maddison et al 2008).
Medical reasons (the list is not exhaustive)
Diabetes, Cushings disease, Addisons disease, Hyperthyoidism, Hypercalcemia, Hypokalemia, Liver disease, generally due to
increased thirst and increased urine output ( Maddison et al 2008).
Kidney failure (maddison et al 2008)
ureta stones (Dalmations have a genetic mutation which interferes in uric acid transportation resulting in an increased risk of urate
stones) (Fascetti et al 2012),
Cancer within the urogenital system (Maddison et al 2008).
Epilepsy (Maddison et al 2008; Overall 2013).
Canine cognitive dysfunction (Osella et al 2007).

Pharmacological causes:
Steroids generally cause increase in thirst and consequently urination (Maddison et al 2008)
 Cholinergic drugs like acetylcholine, produce an urgent desire to void, and is associated with a bladder contraction wave. And anti cholinergic drugs decrease the desire to void (Maddison et al 2008, Yeates 1974). Diuretics causes increase in urination due to their effect of ridding the body of excess water (Maddison et al 2008).
Anticonvulsants usually due to increase in thirst resulting in increase in urination, however Anti convulsants can cause overflow urination, due to their depressive effects on the nervous system, dogs might not feel that their bladder is full and become incontinient ( Maddison et al 2008)
Antihistamines and antipsychotics which contain phenothiazine can cause increase in thirst and consequently increase in urination (Maddison et al 2008)
As a result of choosing just a few pharmacological products it shows how important it is to know what medications or over the counter supplements a dog is taking to identify if that is a likely candidate of causing inappropriate elimination.

If the dog is fearful of being outside he could hold onto his urine and only eliminate where he feels safe the house being his safe place to urinate.
If the dog is excited or absorbed in investigating his outside environment he might forget to eliminate outside, due to his other drives being more important at the time, then when the dog comes in to the house he has a full bladder and is then driven to urinate to reduce his discomfort.

If the dog is excited on homecomings he can urinate in the house this is a result of the activation of the entire autonomic nervous system in co- release resulting in the dog eliminating on greetings the same occurs in submissive weeing (Jenson 2002).

Male dogs are more likely to cock their legs on upright surfaces, particularly if, they are new items being brought in to the house hold. For some dogs this is way of making it smell like it belongs here, for other dogs, novel items in the household are a subject of anxiety and are likely to cause an increase in inappropriate elimination.

As highlighted by Ross (1950) any disturbances in their environment can cause the dog not to eliminate in their preferred area, so a house trained dog who starts to eliminate in the house, a likely candidate is a change in their environment, whether that is a new neighbour, new house, new resident dog or other pet, new baby, change in work schedules, change in feeding and exercise routines, building renovations, decorating, new carpets etc.

Dogs with separation anxiety have an increase in likelihood of eliminating when owners are absent.

Some dogs do not like eliminating in front of owners, and find it difficult to urinate whilst on lead. Occasionally this is caused by owners punishing the dog for urinating or defacating inappropriately, however for other dogs they have a bashful bladder in much the same way as humans suffer from Paruresis (unable to urinate in public) and as a result they come in from being outside and go under the coffee table and urinate. Dogs need to feel safe to eliminate whether that it is urine or
faeces. If they do not feel safe outside they are likely to eliminate where they feels safe and that being indoors.

To sum it all up, individual dogs develop at different rates, some dogs have small bladders some have a larger bladders. Some learn quickly and learn appropriate preferential substrate with little guidance, some dogs need me guidance than others. Some dogs are more anxious than others, some dogs are more busy than others. You hear that pups are housetrained at 8 weeks old, this is a bit farfetched, what it actually means is that there has been good husbandry and the pup has been guided to the appropriate elimination site at the right time. My personal opinion is that dogs are not reliably house trained until they are at least a year old, and even then if the dog has to rely on owners for his elimination needs, without good husbandry the dog can eliminate anywhere.

References

Beaver, B  (2009) Canine Behavior: Insights and Answers . 2nd edition.  Saunders, an imprint of Elsevier Inc.

Beauvais W, Cardwell J M, Brodbelt D C (2012). The effect of neutering on the risk of urinary incontinence in bitches – a systematic review, Journal of  Small Animal Practice 53: 198-204.

De Bleser B, Brodbelt D C, Gregory N G, Martinez T A (2011). The association between acquired urinary sphincter mechanism incompetence in bitches and early spaying: a case-control study, The Veterinary Journal 187: 42-47.

Fowler, C.  Griffiths, D. de Groat, W. (2008) The neural control of micturition Nature Reviews: Neuroscience 2008 June; 9(6): 453–466

Jenson,P. (2002) The Ethology Of Domestic Animals CABI publishing Oxon: UK. New york: USA
Maddison, J. Page,S. Church, D. (2008) Small animal clinical pharmocology. 2nd edition. Saunders elsevier: USA.

Mantis, P. (2008)  Ultrasonography of the Urinary and Genital System of the Dog and Cat. 2nd International Symposium of Veterinary Surgery and 7th Iranian Symposium of Veterinary Surgery, Anesthesia and Radiology, Kerman, Iran, 21-24 April, 2008. Iranian Journal Of Veterinary Surgery 2008 No. Supplement 2 pp. 63-71

Maslow, A. (1943) A Theory Of Human Motivation Psychological Review Vol :50  no : 4  pp 370-396.

Osella,M. Re,G. Odore,R. Girardi,C. Badino,P. Barbero,R. Bergamasco,L. (2007) Canine cognitive dysfunction syndrome: Prevalence, clinical signs and treatment with a neuroprotective nutraceutical Applied Animal Behaviour Science Volume 105, Issue 4, July 2007, Pages 297–310

Overall, K. (2013) Manual of clinical behavioural medicine for dogs and cats. Mosby and imprint of Elsevier Inc: Canada.

Ross, S. (1950) Some Observations on the Lair Dwelling Behavior of  dogs. Behaviour, Vol. 2, No. 3 (1950), pp. 144-162

Wagner, D. Newbury, S. Kass, P. Hurley,K. (2014)  Elimination Behavior of Shelter Dogs Housed in Double Compartment

 Yeates WK. (1974) Neurophysiology of the bladder. Paraplegia ; 12: 73–82

Monday, 12 January 2015

Inconsistency in ownership style and behavioural problems in dogs.

 A dog that receives contradictory and paradoxical information can be the root to behavioural disorders of a dog.  What this means is two opposite messages are emitted for  the exact same piece of information. The double message is a source of confusion and leads to disqualification of all the information, or a part of it, or the response by its receiver (Bateson 1971;1987). 
For instance the most common communication problem that I  come across is that the dog is rewarded or punished alternatively for recall, given a treat or alternatively shouted at. This alternating double message teaches the dog not to come when called, because he has no idea what to expect when he comes back.  The owner thinks that punishing him for the delay in recall means that he will come back immediately when called the next time.  However when the dog comes back from his adventures, those adventures are forgotten, he is attending to owner so the dog believes he is being punished for returning.  
Human communication involves verbal, vocal, postural, motor and contextual levels, successful communication ensures that these levels are congruent (Gross et al 2004).  If you call a dog in a soft voice, but your posture is threatening there is no congruence, this will result in a dog becoming anxious, because there is no consistency in the pattern of  human behaviour. 
Another scenario of inconsistency I often come across is between a couple and their dog. One owner is permissive the other a disciplinarian.  Although both owners might be consistent in the way they treat the dog independently it becomes incongruent for the dog when one minute he is allowed to do something and the next he is being punished for it. For example: allowed to get on the sofa one minute by one owner, shouted at to get down by the other. Therefore punishment is  more likely to be unpredictable for the dog. The stress response to an aversive event is influenced by its predictability and controllability (Arhant et al 2010; Gross et al 2004). There is a chance the  dog might learn not to get on the sofa in the presence of the disciplinarian, but also become anxious in the absence of the disciplinarian, because the dog is confused.

References

Arhant, C.  Bubna-Littitz,H.  Bartels,A.  Futschik, A. Troxler, J (2010) Behaviour of smaller and larger dogs: Effects of training methods, inconsistency of owner behaviour and level of engagement in activities with the dog. Applied Animal Behaviour Science 133 (2010) 131-142
Bateson G.(1971;1987) Steps To An Ecology Of Mind. Collected Essays In Anthropology,Psychiatry, Evolution, And Epistemology. Jason Aronson Inc.Northvale, New Jersey London
Gross,RMcIlveen, R. (2004) Psychology A new introduction. Holder and StroughtonEducational.:UK.

Sunday, 11 January 2015

How can chewing help with learning and behavioural problems?

Mastication involves rhythmic and voluntary movements of lower jaw by the masticatory muscles. Tooth loss and weakness of masticatory muscles causes impairment of masticatory function and chewing disability (Teixeira et al 2014).

Animal experiments have shown that tooth loss or long-term soft-diet feeding causes a decrease in learning and memory ability (Wanabe et al 2002) Another study also revealed that soft-diet feeding after a weaning period reduces synaptic formation in the cerebral cortex and impairs the ability of spatial learning in the adulthood hypothalamus ( Onozuka et al 2002; Teixeira et al 2014). ). Biting during restraint exposure suppresses stress-induced catecholamine concentrations and therefore responses are reduced. This  implies that masticatory activity induced a positive anti-stress effect in animals (cited in Frota de Almedia. Et al 2012). Therefore if a dog has been exposed to uncontrollable stressors, it might be beneficial to give the dog something to chew on.

Mastication is regulated by a neural population in the brainstem and a neural network including several brain regions. Chewing  has been shown to be associated with increased cerebral blood flow, and several studies have reported increased cerebral activity following gum chewing in humans.  Thus, the beneficial effects of chewing on cerebral activity have been suggested, because chewing uses the same neural pathways as cognition so pathways are less likely to decay if chewing is maintained (Sasaki-Otomaru et al 2012; Onozuka et al 2002).

In epidemiological studies, oral function and status have been shown to be related to physical, mental, and social health. In particular, chewing ability has been shown to influence activities of daily living, cognitive status and quality of life (Frota de Alemdia et al 2012).

Therefore chewing is  a fantastic way to ensure that the dogs  jaw muscles are exercised regularly.

Chewing may have  a relaxing effect for the dog aswell as the other benefits described above.  Play is defined by the oxford dictionary(2014) “to engage in an activity for enjoyment and recreation rather than a serious or practical purpose” therefore the chewing  of can be deemed play,  solitary play, and for interacting with as demonstrated in the following video.
http://www.youtube.com/watch?v=HCO4Hj5zbH8

A Kong is flexible dog toy  which the dog can bite down animal experiments have shown this activity induced a positive anti-stress effect in animals (cited in Frota de Almedia. Et al 2012).   Therefore if a dog has been exposed to uncontrollable stressors, it might be beneficial to give the dog a kong to carry with them,  it seems to act as a dummy and can be reassuring to a dog.

References

Frota de Almedia. Et al (2012) Spatial memory decline after masticatory deprivation and aging is associated with alterd laminar distribution of CA1 astrocytes. BMC Neuroscience (2012) volume 13: no 23

Onozuka M, Watanabe K, Fujita M, Tonosaki K, Saito S. (2002) Evidence for involvement of glucocorticoid response in the hippocampal changes in aged molarless SAMP8 mical e. Behavioural Brain Research. 2002;131:125-129

Oxford dictionary  (2014) Oxford Dictionary Of English .version 1.6  Anthony Lewis: Wordweb software.


Sasaki-Otomaru, A. Sakuma, Y Sato, C. (2011) Effect of Regular Gum Chewing on Levels of Anxiety, Mood, and Fatigue in Healthy Young Adults Clinical Practice & Epidemiology in Mental Health, 2011, Vol. 7, pp. 133-139.

Teixeira, F. Fernandes, L Noronha, P et al (2014) Masticatory Deficiency as a Risk Factor for Cognitive Dysfunction International Journal of Medical Sciences
2014; 11(2):209-214.

Wanabe Et al  The molarless condition in aged SAMP8 mice attenuates hippocampal Fos induction linked to water maze performance   Behavioural Brain Research 2002; 128:  19-25

Saturday, 3 January 2015

Separation Anxiety

The term Separation anxiety has its roots in the work of the ethologist Konrad Lorenz  who studied instinctive behaviour particularly imprinting. which is how bond formation occurs between individuals and groups of  mammalian species and birds. Konrad’s theory states that bonds are instinctive and adaptive in nature ( Gross 2004; Garcia 2005).

Bowlby a psychoanalyst, in the 1950’s was inspired by Konrad and applied ethological theory to human attachments.  The key concepts of this theory was that;-
• innate behaviours are essentially the same in all members of a species.
• Fixed action patterns, which are the complex innate behaviours that promote species survival, for example: the attachment behaviours of infant and mother.
Sensitive periods which are specific periods where an animal is biologically ready to acquire a particular new behaviour (Bowlby 1983: Parthasarathy et al 2005: Davenport 1993 ;Gross Et al  2004; Garcia 2005).

Therefore attachment theory has a strong evolutionary base, although exploratory behaviour in young animals is necessary for them to learn about their physical and social world, wandering too far away from the mother would leave that infant vulnerable to predatory and conspecific attack (Gross et al 2004). Therefore it is of evolutionary benefit for the infant to seek the proximity of its mother when it is afraid, and behaviours such as crying; following; and clinging; facilitates gaining and maintaining proximity .  Bowlby termed this behaviour as Separation Anxiety and was a normal reaction of an infant, to maintain close proximity with the attachment figure (Bowlby 1983).

Bowlby describes how the   infant tries to withdraw or escape from a situation or object that they find alarming, and attempt  to go towards or remain with some person or in some place that makes them feel secure. The first type of behaviour is commonly accompanied by a sense of fright or alarm; the second type of behaviour is, of course, what is termed as attachment behaviour. So long as the required proximity to the attachment-figure can be maintained, no unpleasant feeling is experienced. When, however, proximity cannot be maintained because either the figure is lost or some barrier intervenes, the consequent searching and striving are accompanied by a sense of discomfort, and the same is true when loss is threatened. In this discomfort of separation and at a threat of separation , separation anxiety is termed (Bowlby 1983).

Ainsworth (1978,2014) was further inspired and developed a way to test attachment styles through the ‘stranger situation’ and identified that the quality of attachment relationships between infants and caregivers is dependent, upon the degree to which caregivers serve as a secure base for infants. Further analysis, characterised caregivers’ ability to provide a secure base for their infants as emotional availability. Ainsworth described the emotionally available caregiver as promoting an atmosphere of ‘quiet supportiveness’ for autonomous play, thus encouraging infants’ exploration and  also emphasised caregivers’ emotional availability as an important antecedent factor for infants’ attachment security. According to Ainsworth et al, the emotionally available caregiver is accessible and appropriately responsive as well as tuned in to the infant’s signals (Ainsworth et al (1978; Parthasarathy et al 2005; Davenport 1993 ;Gross Et al 2004; Garcia 2005).
Therefore Separation anxiety is not exclusive to dogs or even to veterinary medicine. It is a psychological term to describe the stress and anxiousness in an individual which is brought on by the leaving (perceived or imagined) of another individual (Davenport 1995; Gross et al 2004;Bowlby 1983: Bowlby 1980; Parthasarathy et al: Garcia 2005; Ainsworth et al 1978; Topal et al 1998; Hett 2008).
A more specific definition of separation anxiety requires ongoing attachment to the maternal or primary caregiver . In the case of a dog, separation anxiety is usually an anxiety brought on by separation from the primary caretaker. it is a normal developmental phase and it is necessary for survival.   Separation anxiety disorder occurs later in development  and interferes with normal activities (Landsberg et al 2013; Overall 2013).

What classifies as a "separation”  varies greatly between dogs : some must have "their person" within their line of sight, other dogs  are fine as long as the owner is within a comfortable distance (i.e. somewhere else in the house), and still others are fine until the owner leaves. Even finer distinctions would be dogs that are fine for a certain period of time after their owner leaves, but then start to show signs of anxiousness some time later ( Topal et al 1998; Parathasay 2006).
Just like the variance in what stresses out each individual, the signs of separation anxiety vary greatly as well, from pacing, to learnt helplessness, to barking and  howling, incontinence, and destructive behaviour, just to name a few (0verall 2013; Landsburg et al 2013).

Separation anxiety in dogs was once thought to be a hyper-attachment to the owner. But studies have shown that dogs with separation anxiety and dogs without separation anxiety displayed same behaviours around owners, i.e. follow owner around the house (Mariti et al 2013; Horn et al 2013). Behaviour modification protocols for separation anxiety that recommend owners ignore their dogs and also the behaviours from the dog  that are designed to promote social contact,  is a harmful recommendation because it damages the human-animal bond and produces anxiety and frustration, because the dog is powerless to initiate social interaction, and the owner is prevented from doing so (Hett et al 2008; Parathasy et al 2006) In addition  the  hyper-attachment behaviour is more likely to be due to an insecure attachment to the owner.  Ainsworth demonstrated in the ‘strange situation’ different forms of attachment.  Secure attachments when the attachment figure was compassionate to the needs of the child; insecure attachments seen in children whose attachment figure was not responsive to the needs of the child, and ambivalent attachments, where children showed no attachment to the attachment figure, due to a caregiver being inconsistent in emotional and physical availability to the child.   Many years ago it was once thought that children should be seen and not heard, that they need to learn to have a backbone, and responding to the childs needs immediately was deemed that the child would become too reliant on the caregivers.
Children brought up this way resulted in a child who cried more, explored less, and clung more to caregivers and were not easily reassured, than the children who had secure attachments to their caregivers (Ziv et al 2013).

The strange situation has been repeated with dogs  and found the same findings secure attachments, insecure attachments as is avoidance and ambivalent attachments  (Topal et al 1998; Hett et al 2008; Mariati et al 2013,Horn et al 2013).

Attachments are not just for caregivers, they can be for place, objects whatever makes mammals feel safe, usually attachments occur first with primary caregivers, which gives the individual the safety and security to  explore their environment, when they become comfortable after exploration, the attachment to that place becomes another  secure base. The BBC episodes Monkey planet aired on the 2nd April 2014 demonstrates a place as an attachment,  the baby gibbon was too scared to leave the safety of the enclosure to go and investigate  the big outside world, mum goes and reasures him, the infant comes out of the enclosure, Mum cuddles him again, and hay presto he is swinging from the trees free, the video clip below is not from that programme and does not demonstrate the cuddle but the calm reassurance from mum, giving the infant the confidence to begin his exploration of the outside world.  http://www.youtube.com/watch?v=IT3aLsTxbnQ

Most researchers assume that the acceptance of human beings as conspecific alongside the mother-like and effective security-providing role of humans for distressed puppies are results of domestication. For 10,000 years, artificial selection in dogs favoured socialisation with humans as if they were conspecific.  The genetic changes regarding a dog's capacity for conspecific recognition, might have played a key role in this process that resulted in a preference for humans. A dog's preparedness for forming a bond with humans, a bond that is rooted in the evolutionary past, would be a prerequisite for the development of attachment between a particular person (the owner) and the dog itself (Topal et al 2011).

Although the ability to form attachments is usually associated with an early sensitive period, Gasci et al (2001) demonstrated that in certain conditions a short responsive interaction with an unfamiliar human individual may result in attachment behaviour even in the case of dogs that are more than 1 year old. Gasci et al (2001) study in rescue centres show that dogs of low or restricted contact with humans may retain their ability to form new attachment relationships with humans. Probably the extreme separation from human social contact (i.e., shelter conditions) has a crucial role in this sensitisation process. In other words, dogs living in poor social conditions become more responsive to humans, which results in a remarkable readiness to form attachment relationships. It should be noted that in the case of rhesus monkeys, a similar effect has been shown because most abnormally socialised monkeys could be rehabilitated to a certain extent by appropriate exposure to conspecific groups and individuals (Harlow et al1971).

Panskepp  (2004) looks at the neurocircuitry and neurochemistry of social bonds, and is assumed that the arousal of the panic system is one of the many driving forces that guide the construction of social bonds and identifies that the attachments and bonds increase brain opoids,  and on separation, the neurotransmitters glutamate and corticotrophin releasing hormone are the neurotransmitters that are responsible for the panic response of separation distress.  According to Panskepp the panic system is distinct from the fear system because the administration of  opiates  to new born rat pups proved effective in reducing separation distress but has no effect on fearful behaviours. This nil effect of opoid administration on fearful behaviours was demonstrated when pups were separated from mother and put in a strange environment, so the addition of strange environment provoked the fear system.  The distress calls of young animals are to seek reunion with individuals who help create the feeling of a ‘secure neurochemical base’. Panskepp hypothesises  the distress calls from the infant arouses distress circuits in the parents which facilitates the social bond. Harlow (1971) demonstrated that socially deprived monkeys who’s behaviour to other monkeys was sociopathic could be driven to demonstrate maternal behaviour after incessant separation distress calls from the infant.

Having an understanding and compassion of the legitimate reasons why dogs suffer from separation anxiety and not treating it as a problem behaviour , will go along way to helping the dog be happy alone.
• So if a dog is showing anxiety after separation but is easily settled when owner returns, the owner has done a fantastic job of ensuring that the dog feels safe in their company.
• If a dog is showing anxiety after separation but is not easily settled by the owners return this shows that the dog is insecurely attached to the owner.
• If the dog shows no outward signs of separation anxiety but greets the owner on return from separation, the dog has learnt to be happy alone in his environment , owners done a fantastic job.
If the dog shows no outward signs of separation anxiety and does not greet the owner on return there is no attachment. There has to be an attachment in order for separation anxiety to exist (Bowlby 1983).

So ensure that puppies are well socialised, that the dog has a secure attachment to the owner, and other members of the family,  this is done with quality time spent in each others company, touching, playing feeding etc, once the dogs understands that the owner is the safe base, the dog can explore his environment, and make an attachment to his environment, this environment becomes a secure base (Scott 1958: Morgan 2010).  A routine gives them knowledge that food and walks and cuddles and playtime etc will be always there and life becomes more predicable, eventually they learn to cope with changes in routines, but in order for that to happen the attachments and their safety needs have to be met first.

References


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