Changes

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 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.  

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 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.  

Olfactory communication involves the deposition of scent marks in the environment to convey a signal indirectly to another individual. Scent signals carry information and can convey 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 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 odours 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==