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==Olfactory Communication==

==Olfactory Communication==

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 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.  

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 marks are not only a signal to other cats, but also carry meaning for the depositor. In the core territory, cats are more likely to be in a parasympathetic state of arousal, in preparation for activities such as resting, feeding and grooming. Outside the core territory, some degree of sympathetic arousal is always required, as the individual is either vulnerable or engaged in hunting. Facial and flank marks may provide a chemical signal to the cat that favours a parasympathetic state in all individuals that are present, reducing the likelihood of conflict in colonies where several cats share a common core territory.

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. 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.  

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.  

Scent signals carry both information and meaning. The information relates to the signaller; its identity, health and reproductive state. 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.  

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.

*'''Facial and flank marks''' are signals largely used in a cat’s '''core territory''', where it only expects to encounter other familiar cats. In areas where facial and flank marks are left, other forms of marking (spraying and clawing) are reduced or absent.

*'''Claw marking''' is 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.  

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.

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.

===Claw Marks===

===Claw Marks===

Claw marks are used to indicate firmer territorial boundaries that discourage the presence of non-resident cats.

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.

 

===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 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]] 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 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.

 

 

 

 

 

 

 

==References==

==References==