Changes

Cats are not asocial, they will form colonies where resources are plentiful and they do develop [[Feline Social Behaviour|affiliative relationships]] with each other when there is a mutual benefit. For example, groups of related females may form in an area where there is an excess of hunting and sheltering opportunities. The primary means of identification between cats in a social group is through the establishment of a group odour. This requires that cats approach close enough to groom and rub against each other (allogroom and allorub). It would not be possible to establish a group odour if cats did not have some communicative mechanism for reducing distance. So, distance reducing behaviours are an essential part of communication that allows cats to approach one another.

Cats are not asocial, they will form colonies where resources are plentiful and they do develop [[Feline Social Behaviour|affiliative relationships]] with each other when there is a mutual benefit. For example, groups of related females may form in an area where there is an excess of hunting and sheltering opportunities. The primary means of identification between cats in a social group is through the establishment of a group odour. This requires that cats approach close enough to groom and rub against each other (allogroom and allorub). It would not be possible to establish a group odour if cats did not have some communicative mechanism for reducing distance. So, distance reducing behaviours are an essential part of communication that allows cats to approach one another.

====Allogrooming and allorubbing====

====Allogrooming and Allorubbing====

Cats that are part of a social group will groom and rub against each other each other in order to transfer chemical odour signals between cats and establish a "group odour". This group odour is a common identifier that enables members of a social group to recognise each other. A significant benefit of this method of group recognition is that, since the odour is transferred from cat to cat throughout a group, not all cats need to groom each other, or even meet on a regular basis, in order to share enough elements of the group odour to recognise one another when they meet. It should be remembered that this system evolved to regulate interactions between related individuals, and the genetic contribution to personal odour in cats is not yet understood. It may be more difficult for unrelated cats to establish a group odour, if there is less commonality in their personal chemical identity.

Cats that are part of a social group will groom and rub against each other each other in order to transfer chemical odour signals between cats and establish a "group odour". This group odour is a common identifier that enables members of a social group to recognise each other. A significant benefit of this method of group recognition is that, since the odour is transferred from cat to cat throughout a group, not all cats need to groom each other, or even meet on a regular basis, in order to share enough elements of the group odour to recognise one another when they meet. It should be remembered that this system evolved to regulate interactions between related individuals, and the genetic contribution to personal odour in cats is not yet understood. It may be more difficult for unrelated cats to establish a group odour, if there is less commonality in their personal chemical identity.

====Affiliative Behaviour====

====Affiliative Behaviour====

When approaching to greet another cat, a cat will  move its '''tail into a vertical position''' (tail-up). A '''quiet trill or chirrup''' is issued, and after getting to within around 0.5 metres the approaching cat will '''sit down'''. '''Slow eye blinking''' is also used to indicate that this is a friendly approach. Physical contact only occurs between cats in in about 30% of these approaches. Two cats may approach each other with tail-up, in which case they will often sit down in close proximity, but still without physical contact. When cats do make contact during a greeting, it is often only briefly; one or two rubs or a brief period of grooming. Cats that are younger than 2 years of age are more sociable and also more likely to engage in closer contact after a greeting, possibly even playing.

When approaching to greet another cat, a cat will  move its '''tail into a vertical position''' (tail-up). A '''quiet trill or chirrup''' is issued, and after getting to within around 0.5 metres the approaching cat will '''sit down'''. '''Slow eye blinking''' is also used to indicate that this is a friendly approach. Physical contact only occurs between cats in about 30% of these approaches. Two cats may approach each other with tail-up, in which case they will often sit down in close proximity, but still without physical contact. When cats do make contact during a greeting, it is often only briefly; one or two rubs or a brief period of grooming. Cats that are younger than 2 years of age are more sociable and also more likely to engage in closer contact after a greeting, possibly even playing.

==Visual Communication==

==Visual Communication==

The function of the visual system of cats is highly biased in favour of [[Feline Predatory Behaviour|predatory behaviour]]. The cat’s [[Eye - Anatomy & Physiology#The Wall (retina, uvea and sclera)|retina]] has three times the rod density of the human eye, contributing to the light detection threshold in cats being eight times lower than in man. Although their vision is not monochromatic, cats have sixteen times fewer wavelength comparing retinal ganglia than primates and are '''behaviourally colour blind'''. The cat’s enhanced visual sensitivity to brightness, patterns and movement block learning associated with differences in colour; under normal conditions cats do not appear to learn associations based on colour discrimination. The [[Cranial Nerves - Anatomy & Physiology#Optic Nerve (II)|optic nerve]] in the cat has a much lower density of nerve fibres than in man, due to the much smaller amount of information transmitted from the cat’s retina. This is because the ratio of ganglionic cells to photoreceptors is very high in the cat; there is a far higher level of integration of information at the level of the ganglionic layer of the retina. The benefit of this is that '''movement detection is hard-wired into the sensory system''' and able to directly drive fast responding reflexive systems; the cat is therefore much more rapidly attentive and responsive to movement than a human. The overall effect is that cats are attentive to movement in the way that humans are attentive to colour.  

The function of the visual system of cats is highly biased in favour of [[Feline Predatory Behaviour|predatory behaviour]]. The cat’s [[Eye - Anatomy & Physiology#The Wall (retina, uvea and sclera)|retina]] has three times the rod density of the human eye, contributing to the light detection threshold in cats being eight times lower than in man. Although their vision is not monochromatic, cats have sixteen times fewer wavelength comparing retinal ganglia than primates and are '''behaviourally colour blind'''. The cat’s enhanced visual sensitivity to brightness, patterns and movement block learning associated with differences in colour; under normal conditions cats do not appear to learn associations based on colour discrimination. The [[Cranial Nerves - Anatomy & Physiology#Optic Nerve (II)|optic nerve]] in the cat has a much lower density of nerve fibres than in man, due to the much smaller amount of information transmitted from the cat’s retina. This is because the ratio of ganglionic cells to photoreceptors is very high in the cat; there is a far higher level of integration of information at the level of the ganglionic layer of the retina. The benefit of this is that '''movement detection is hard-wired into the sensory system''' and able to directly drive fast responding reflexive systems; the cat is therefore much more rapidly attentive and responsive to movement than a human. The overall effect is that cats are attentive to movement in the way that humans are attentive to colour.  

The cat's vision has therefore evolved to enable it to see in low light levels, break the camouflage of its prey and quickly detect movement. As a result of the wild cat’s camouflaged coat, [[Timing of Feline Activity|crepuscular and nocturnal activity]] and [[Feline Predatory Behaviour#Hunting Strategies|stealthy use of cover]], visual identification of affiliates is less important than the use of other sensory systems such as [[Feline Communication Behaviour#Olfactory Communication|olfaction]].

The cat's vision has therefore evolved to enable it to see in low light levels, break the camouflage of its prey and quickly detect movement. As a result of the wild cat’s camouflaged coat, [[Timing of Feline Activity|crepuscular and nocturnal activity]] and [[Feline Predatory Behaviour#Hunting Strategies|stealthy use of cover]], visual identification of affiliates is less important than the use of other sensory systems such as [[Feline Communication Behaviour#Olfactory Communication|olfaction]].

Visual communication involves '''facial and body postures''', as well as the '''visual aspects of''' certain behaviour such as '''spray marking''' and '''clawing'''. In general, visual communication is operates over a limited range, in line of sight and is rapidly modified. This gives the signaller a great deal of control over the signal, which can be changed according to the response it elicits. Visual signalling is less effective at night, in adverse weather conditions and dense undergrowth. The main disadvantage of visual communication is that it must be delivered face to face with a competitor, which increases the risk of conflict and injury.  

Visual communication involves '''facial and body postures''', as well as the '''visual aspects of''' certain behaviour such as '''spray marking''' and '''clawing'''. In general, visual signals operate over a limited range, in line of sight and are rapidly modified. This enables the signaller to alter its communication according to the response it elicits. Visual signalling is less effective at night, in adverse weather conditions and dense undergrowth. The main disadvantage of visual communication is that it must be delivered face to face with a competitor, which increases the risk of conflict and injury.  

Visual communication in cats therefore tends to be vivid. '''Whole body signals''' are used to make the signaller appear larger and more threatening, or smaller and less threatening. The most powerful signals are those at the extreme, such as the side-on body posture with arched back and tail erect.  

Cats use their '''whole body''' to convey information about their emotional and intentional state. These signals are vivid, but difficult to modify quickly.  The signaller attempts to make itself appear larger and more threatening, or smaller and less threatening. The most powerful signals are those at the extreme, such as the side-on body posture with arched back and tail erect.  

The intention of all body posture signalling during a conflict is to '''maintain distance between individuals''', either by intimidating an opponent into a retreat, or by indicating such a low level of threat that an opponent has no need for an approach. The roll is not a position of complete vulnerability in cats, and in this position the cat is able to make very good use of its teeth and all sets of claws. After a conflict, both parties go their separate ways and so there is no need for post-conflict resolution.  

'''Facial postures''' are less easily observed at a distance, but they can be changed more rapidly and therefore provide a more precise indication of the cat's state at a given moment. Aggressive facial postures are often supported by vocalisations, such as hissing and spitting, that further intensify their meaning. Changes in ear position, pupil dilation and the display of teeth are often not paralleled by body posture changes, since these are slower to accomplish, especially when the animal has adopted a fully erect or flattened body position.

'''Facial posture''' is extremely important in conveying momentary changes in the emotional state of the cat. Changes in ear position, pupil dilation and the display of teeth are not paralleled by body posture changes, since these are slower to accomplish, especially one the animal has adopted a fully erect or very low body position.

The intention of all postural signalling during a conflict is to '''maintain distance between individuals''', either by intimidating an opponent into a retreat, or by indicating such a low level of threat that an opponent has no attack and the signaller is able to make a discreet retreat. If two opponents engage in physical conflict, one may choose to roll on its back. In dogs, this can be interpreted as a display of vulnerability, and is an appeasement behaviour that diffuses tension and stops further violence. The roll does not carry the same meaning in cats; when in this position a cat is better able to use claws and teeth, and is not attempting to appease or diffuse conflict. After a conflict, there is no need for the kind of post-conflict reconciliation that is seen between dogs. Post conflict reconciliation between dogs reduces the risk of further conflict, and enables individuals to return to their role within the group. Cats that are not part of a social group are competitors and do not have any relationship that needs to be repaired or maintained.

==Vocal Communication==

==Vocal Communication==

Vocal communication includes a wide range of sounds from quiet purrs and trills to loud distress vocalisations. Vocal signals can vary from short to long range, but are always omnidirectional and carry a risk of being detected by cats other than the intended recipient. Vocal communication can be used at a distance in any weather or lighting conditions.

Vocal communication includes a wide range of sounds from quiet purrs and trills to loud distress vocalisations. Vocal signals can vary from short to long range, but are always omnidirectional and carry a risk of being detected by cats other than the intended recipient. Vocal communication can be used at a distance in any weather or lighting conditions, and are often used to support and strengthen the meaning of visual signals.

In general sounds that are '''produced with the mouth closed''', such as purrs, trills and chirrups, are '''distance-reducing signals'''. The same is true of vocalisations that are produced with the mouth initially open but closing whilst the sound is produced, such as the miaow. Strained intensity vocalisation that are '''made with the mouth open''' throughout the sound are generally '''distance increasing signals or signs of distress''', such as growls, hisses, spits, snarls and shrieks<ref>Kiley-Worthingon, M. (1984) Animal language? Vocal communication of some ungulates, canids and felids. ''Acta Zoologica Fennica''. 171, 83-88.</ref>.

In general sounds that are '''produced with the mouth closed''', such as purrs, trills and chirrups, are '''distance-reducing signals'''. The same is true of vocalisations that are produced with the mouth initially open but closing whilst the sound is produced, such as the miaow. Strained intensity vocalisation that are '''made with the mouth open''' throughout the sound are generally '''distance increasing signals or signs of distress'''. This includes growls, hisses, spits, snarls and shrieks<ref>Kiley-Worthingon, M. (1984) Animal language? Vocal communication of some ungulates, canids and felids. ''Acta Zoologica Fennica''. 171, 83-88.</ref>.

 

There is some evidence that cats are able to modify their vocalisations when communicating with people. In a study investigating human sensitivity to purring in cats, it was found that cats used a "solicitation purr" to obtain a response from people. This purr contains an embedded high pitched component that resembles sounds made by crying human infants, and which people experience as sounding urgent and disturbing<ref>McComb, K., Taylor, A.M., Wilson, C., Charlton, B.D. (2009) The cry embedded within the purr. ''Cell Current Biology''. 19(3), 507-508.</ref>.

==Olfactory Communication==

==Olfactory Communication==

Olfactory communication involves the deposition of scent marks in the environment to convey a signal indirectly to another individual. Scent signals carry both information and meaning. Apart from providing other cats with information about ownership and usage of territory, scent marks also convey information about the identity of the depositor, its sex, health and reproductive status. This enables more complex [[Feline Social Behaviour|social organisation]] than merely avoidance, with cats choosing to avoid an area or attempt to gain control of it, depending on the threat posed by the current resident and the need to gain access to the resources included in that area.  

Scent signals are localised, persistent and offer only an indirect form of communication. They cannot be removed by the signaller once they have been deposited and thus they can convey information about the presence of the signaller that may be to its detriment. For example, giving away information about the resident animal's territorial range and the timing of its activities.

Rodents have also been shown to exhibit unconditioned fear and avoidance behaviour in response to feline odours<ref>Dielenberg, R.A., Hunt, G.E., McGregor, I.S. (2001) When a rat smells a cat': The distribution of Fos immunoreactivity in rat brain following exposure to a predatory odor. ''Neuroscience''. 104(4), 1085-1097.</ref>. ''[[Toxoplasma gondii]]'' infection in rodents has been shown to have a highly specific effect on this aversive response, producing not merely a loss of fear of feline urine odours, but an attraction to them<ref>Vyas, A., Kim, S., Giacomini, N., Boothroyd, J.C.,. Sapolsky, R.M., (2007) Behavioral changes induced by Toxoplasma infection of rodents are highly specific to aversion of cat odors. ''PNAS''. 104(5), 6442-6447.</ref>. This enables the parasite to continue its life cycle by infection of its feline definitive host. The behavioural changes seen are completely specific to feline odours, with other fear and conditioning responses remaining unaffected.  

Scent signals are persistent, localised and indirect. They cannot be removed by the signaller once they have been deposited and thus they can convey information about the presence of the signaller that may be to its detriment. For example, rodents have been shown to exhibit unconditioned fear and avoidance behaviour in response to feline odours<ref>Dielenberg, R.A., Hunt, G.E., McGregor, I.S. (2001) When a rat smells a cat': The distribution of Fos immunoreactivity in rat brain following exposure to a predatory odor. ''Neuroscience''. 104(4), 1085-1097.</ref>. In a further twist, ''[[Toxoplasma gondii]]'' infection in rodents has been shown to have a highly specific effect on this aversive response, producing not just a loss of fear of feline urine pheromone odours, but also an attraction to them<ref>Vyas, A., Kim, S., Giacomini, N., Boothroyd, J.C.,. Sapolsky, R.M., (2007) Behavioral changes induced by Toxoplasma infection of rodents are highly specific to aversion of cat odors. ''PNAS''. 104(5), 6442-6447.</ref>. This enables parasite to continue its life cycle by infection of its feline definitive host. The behavioural changes seen are completely specific to feline odours, with other fear and conditioning responses remaining unaffected. Scent signals can also carry a considerable distance, as is the case in urinary scent marks by females during the reproductive season.  

Olfactory communication involves the deposition of scent marks in the environment to convey a signal indirectly to another individual. Apart from providing other cats with information about ownership and usage of territory, scent marks also convey information about the identity of the depositor, its sex, health and reproductive status. This enables more complex [[Feline Social Behaviour|social organisation]] than merely avoidance, with cats choosing to avoid an area or attempt to gain control of it, depending on the threat posed by the current resident and the need to gain access to the resources included in that area. Scent signals can also carry a considerable distance, as is the case in urinary scent marks by females during the reproductive season.  

The personal identification element of the signal has common features between claw marks, spray marks and facial/flank marks, so that the recipient of any of these signals will be aware of which individual is responsible. Previous experience with that individual adds value to this information. Meaning is conveyed in the position and type of signal used.

The personal identification element of the scent signal has common features with claw marks, spray marks and facial/flank marks, so that the recipient of any of these signals will be aware of which individual is responsible. Previous experience with that individual adds meaning to the location and type of signal.  

Claw, urine and facial/flank marks are generally deposited on vertical surfaces, often on objects that are close to an entry point to a particular space, so that other cats will notice them.

===Marking===

===Marking===

The function of marking behaviour is to identify the significance of certain locations to the ‘sender’ and ‘receiver’ of the mark. Scent marks, therefore, act both as a memento of previous experience in a location as well as a signal to others. When a cat encounters the facial and flank marks on inanimate objects in the [[Feline Territorial Behaviour#Core Territory|core part of the territory]], they signify that this location has been safe in the past and when a cat leaves another face or flank mark, it is relabelling that place as safe based upon its current experience.  

Claw, urine and facial/flank scent marks are generally deposited on vertical surfaces, often on objects that are close to an entry point to a particular space, so that other cats will notice them.  The function of this marking behaviour is to identify the significance of certain locations to the ‘sender’ and ‘receiver’ of the mark. Scent marks, therefore, act both as a memento of previous experience in a location as well as a signal to others. When a cat encounters facial and flank marks on inanimate objects in the [[Feline Territorial Behaviour#Core Territory|core part of the territory]], they signify that this location has been safe in the past and when a cat leaves another face or flank mark, it is relabelling that place as safe based upon its current experience.  

The odours that cats share when allogrooming and allorubbing help to identify the group so that these and the core territory odours are a memento of previous interactions.  

The odours that cats share when allogrooming and allorubbing help to identify the group so that these and the core territory chemical messages are a memento of previous interactions.  

Other odour marks are intended to enable cats to maintain distance from one another. Both claw marks and urine spray marks contain pheromone chemical signals that are intended to signal to cats outside the social group that they are entering an area that is also occupied by other cats.  

Other odour marks are intended to enable cats to maintain distance from one another; both claw marks and urine spray marks contain pheromone chemical signals that are intended to signal to cats outside the social group that they are entering an area that is also occupied by other cats.  

====Facial and Flank Marks====

====Facial and Flank Marks====

Spray marks are used to maintain '''temporal separation''' by indicating information about ownership of space as well as when a particular cat visits that location. The rate of spray marking by male and female cats is increased during the period when females are in [[Oestrus Behaviour - Anatomy & Physiology|oestrus]]. As a result, the rate of spray marking in males and females decreases after neutering, as does the pungency of the male spray mark.  

Spray marks are used to maintain '''temporal separation''' by indicating information about ownership of space as well as when a particular cat visits that location. The rate of spray marking by male and female cats is increased during the period when females are in [[Oestrus Behaviour - Anatomy & Physiology|oestrus]]. As a result, the rate of spray marking in males and females decreases after neutering, as does the pungency of the male spray mark.  

Both spray marking and claw marking also include an element of visual display. During spray marking, the cat will sniff the site to be marked, using a [[Vomeronasal Organ#The Flehmen Reaction|Flehmen behaviour]] to collect more of the pheromone signal. Then it will back up to the site to be sprayed, with its tail raised. During the act of spraying the cat’s tail twitches and moves, the cat has a glazed look in its eye and it will often make treading movements with its hind feet. This pattern is distinct from urination which is usually on horizontal surfaces (unless the cat has [[:Category:Urological Diseases - Cat|urinary tract disease]]) and is not accompanied by tail movements or treading.

Both spray marking and claw marking also include an element of visual display. During spray marking, the cat will sniff the site to be marked, using a [[Vomeronasal Organ#The Flehmen Reaction|Flehmen behaviour]] to collect more of the pheromone signal. Then it will back up to the site to be sprayed, with its tail raised. During the act of spraying the cat’s tail twitches and moves, the cat has a glazed look in its eye and it will often make treading movements with its hind feet. This pattern is distinct from urination which is usually on horizontal surfaces (unless the cat has [[:Category:Urological Diseases - Cat|urinary tract disease]]) and is not accompanied by tail movements or treading. The tail movements and treading are part of the process of expulsion of the chemical pheromone signal, but they are also a visual signal that other cats can observe.

Urine spray marks provide a signal between cats that a particular area is likely to be in use by different individuals at certain times of day. The degradation of a urine mark over time provides other cats with an indication of when the depositor was last in that location. With regular re-marking, this provides cats with a geographical and temporal map that enables them to pass through an area without encountering one another.

The degradation of a urine mark over time provides other cats with an indication of when the depositor was last in that location. Typically, spray marks are used at the outermost boundary of territory and in areas that may be shared with other cats. With regular re-marking, this provides cats with a geographical and temporal map that enables them to pass through an area without encountering one another.

====Claw Marks====

====Claw Marks====

Claw marks are used to indicate firmer territorial boundaries that discourage the presence of non-resident cats. It is also used to stretch back muscles after resting, as well as to maintain the condition of claws, so owners are often confused about its meaning as a form of communication. When scent is released from pedal glands during clawing, this creates a signal that indicates territorial residency.  

When scent is released from pedal glands during clawing, this creates a signal that indicates territorial residency. These scent marks are used to indicate a more fixed territorial boundary that is not affected by timing and discourages the entry of non-resident cats into their territory. Clawing behaviour is also used to stretch back muscles after resting, as well as to maintain the condition of claws, so owners are often confused about its meaning as a form of communication.  

===Pheromones===

===Pheromones===

The meaning of the chemical signal is conveyed using pheromones that form part of the spray, claw or facial/flank mark. Unlike the releaser pheromones that are found in insects, which have a direct and immediate effect on behaviour, mammalian chemical signals have a more '''subtle effect on mood, emotion and behavioural priorities'''. For example, in the presence of female sex pheromone a male cat will continue with its current behaviour but its priorities will shift toward seeking out a mate.

The meaning of the chemical signal is conveyed using pheromones that form part of the urine-spray, claw or facial/flank mark. Unlike the releaser pheromones that are found in insects, which have a direct and immediate effect on behaviour, mammalian chemical signals have a more '''subtle effect on mood, emotion and behavioural priorities'''. For example, in the presence of female sex pheromone a male cat will continue with its current behaviour but its priorities will shift toward seeking out a mate.

In cats, pheromone signals are composed of combinations of fatty acids that are deposited together on a scent marked site. Cat urine also contains the strong smelling compound 3-methyl-3-methylbutan-1-ol (MMB), which a chemical signalling function. This is produced by the action of the enzyme cauxin, which hydrolyses the chemical felinine that is synthesised in the [[Liver - Anatomy & Physiology|liver]] and excreted in urine.

In cats, pheromone signals are composed of combinations of fatty acids that are deposited together on a scent marked site. Cat urine also contains the strong smelling compound 3-methyl-3-methylbutan-1-ol (MMB), which is a pheromone precursor. This is produced by the action of the enzyme cauxin, which hydrolyses the chemical felinine that is synthesised in the [[Liver - Anatomy & Physiology|liver]] and excreted in urine.

The recipient of the signal will sniff the location, and may perform a [[Vomeronasal Organ#The Flehmen Reaction|Flehmen behaviour]] in order to draw a larger quantity of pheromone through ducts opening form the roof of the mouth and floor of the nasal cavity into a specialised chemosensory organ called the [[Vomeronasal Organ|vomeronasal organ (VNO)]]. The VNO consists of a pair of 1-2cm long tubular organs in the [[Hard Palate|hard palate]]. Once in the VNO pheromone chemicals bind to a pheromone binding protein before being presented to cell-surface receptors. Signals from the VNO are directed toward primitive brain structures such as the [[Forebrain - Anatomy & Physiology|amygdala]], that are involved in subconscious changes in mood, emotion and motivation.

The recipient of the signal will sniff the location, and may perform a [[Vomeronasal Organ#The Flehmen Reaction|Flehmen behaviour]] in order to draw a larger quantity of pheromone through ducts opening from the roof of the mouth and floor of the nasal cavity into a specialised chemosensory organ called the [[Vomeronasal Organ|vomeronasal organ (VNO)]]. The VNO consists of a pair of 1-2cm long tubular organs in the [[Hard Palate|hard palate]]. Once in the VNO pheromone chemicals bind to a pheromone binding protein before being presented to cell-surface receptors. Signals from the VNO are directed toward primitive brain structures such as the [[Forebrain - Anatomy & Physiology|amygdala]], that are involved in subconscious changes in mood, emotion and motivation.

==References==

==References==

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[[Category:Normal Feline Behaviour]]

[[Category:Normal Feline Behaviour]]