Pheromones: cat

Although the structures that secrete pheromones are still poorly known, it seems to have been established that the production and secretion of pheromones are involuntary.

In carnivores the pheromones are secreted by various glandular structures in the skin and mucous membranes around the natural body orifices. The main secretory structures in the cat are the sebaceous glands in the intermammary sulcus, the perioral glands (spread across the chin, lips and skin of the face near the vibrissae and the cheeks, extending to the area of glabrous skin at the temples), the anal glands (including the hepatoid glands, the sebaceous glands in the cutaneous part of the anus, rectal mucosa and the anal sacs), the supracaudal glands (on the dorsal surface of the base of the tail; it seems that the activity of these glands is closely related to the secretion of sex steroids, such that orchidectomy or ovariectomy leads to a notable reduction in the secretion of pheromones), the subcaudal glands (on the ventral surface of the tail, which, although poorly developed, are explored at length during interactions) and the footpad glands (located in the plantar footpads and in the skin of the interdigital region). Furthermore, there are pheromones which appear to be in solution in the amniotic fluid (called pheromones ‘of adoption’), in the saliva, in the urine (urinary marking) and, according to P. Pageat, in the faeces.

PHEROMONES IN THE CAT

Only very few pheromones have so far been identified and related to a precise behavioural function. As for canine pheromones, the known feline pheromones have been classified according to the structures that secrete them and on the basis of their action. Therefore, there are pheromones of adoption, appeasement, “territory structuring” and alarm as well as sexual pheromones. At birth, the kitten is covered by a pheromone dissolved in the amniotic fluid whose action lasts 3 days and promotes bonding between the mother and kitten. This pheromone “of adoption” can trigger maternal behaviours also in a queen (belonging to the “social” group) which has not given birth because the young of the feline species are often raised “in a nursery”.

The development of mother-kitten “bonding” guarantees the survival of the young animal through the administration of food, the presence of a source of heat and protection from predators. Following a Caesarean section or in order to facilitate the adoption of a newly born kitten by another female, it can be useful to save part of the amniotic fluid to sprinkle over the neonate before presenting it to the mother. Pluriparous females are less sensitive to pheromones of adoption because only seeing the neonatal kitten is often sufficient to trigger maternal behaviour. From the third to fourth day after delivery, the sebaceous glands of the intermammary sulcus of lactating females secrete “appeasement pheromones”, or the apaisines, which in the cat are called cat appeasing pheromones (C.A.P.). These secretions facilitate the development of bonding between the kitten and its mother. The chemical composition of the C.A.P. is very similar to that of facial pheromones (F3 fraction) even though some studies have shown that kittens less than 3 weeks old cannot recognize the structural analogues.

Through the deposition of pheromones secreted from the perioral glands and glands in the skin of the interdigital region as well as the contents of urinary marking, the cat “structures” the environment in which it lives, subdividing it into numerous zones called territorial fields, which are connected to each other by “invisible” routes called paths. The pheromones contained in the secretion of some glands in the lateral part of the face consist of several fractions and are deposited by rubbing the cheek, from the labial commissure to the area of hairless skin at the temple (Fig. 1). Olfactory marking is conducted mainly for the benefit of the individual that performs it; in fact, these secretions are deposited on “explored” objects, individuals belonging to the same social group, and vertical substrates (just before emission of urine marks during sexual behaviour); the marking indicates that the object, individual or substrate is considered not dangerous. Urine spraying not related to courting behaviour is performed at the junction between a territorial field and a path.

The pheromones secreted by the interdigital glands are deposited during scratching and mainly delimit the fields of isolation (e.g. resting areas) and elimination (Fig. 2).

The “alarm” pheromones act on both the individual that emits them and on other animals of the same genus. They are deposited in the case of notable stress (for example, during a clinical examination) and are secreted by the glands in the plantar footpads and anal sacs. The perception of these pheromones by both animals of the same species and those of different species (for example, dogs) causes avoidance of the marked area, tachycardia, tachypnoea, mydriasis and tremors. The alarm pheromones emitted by the glands in the plantar footpads are “linked” to a highly adhesive protein. The examination table and carry box must, therefore, be cleaned carefully with hot water and neutral soap in order to remove these pheromones definitively, otherwise the emotional state of the cat will be immediately disturbed the next time it undergoes an examination or is put into the carry box again after some time.

The pheromones secreted by the supracaudal and subcaudal glands seem to be involved in social interactions, in particular in the recognition of individuals and sexual behaviour. The pheromones contained in the anal secretions are evacuated during defecation and seem to be implicated in sexual behaviour, like those emitted during urinary marking by both males and females, during the courting behaviour prior to mating. It is very probable that there are other pheromones but so far it has not been possible to identify them.

THE CAT AND ITS TERRITORY

The territory in which a cat lives is a complex structure made up of multiple fractions called “territorial fields” (Waser & Wiley, 1979). The space used by the animal is subdivided into a variable number of areas, of different sizes, each related to a precise behavioural activity. According to P. Pageat, three types of territorial fields can be distinguished:

• Activity fields: these are areas in which the cat carries out a precise activity: hunting (feeding), playing, elimination;
• Isolations fields: in these areas, the animal isolates itself and avoids contact. There are usually two or three isolation fields, preferably located at a height: the most classic is the resting place, which also serves as a refuge in the case of need. In poorly sociable animals, the site of elimination can be considered an isolation field. According to J. Dehasse the reproduction field can also be considered an isolation field and when another male individual enters it, this immediately produces aggressive behaviour;
• Aggression field: this is a space of varying size, centred on the individual (“the bubble”). Any intrusion causes aggressive behaviour almost immediately. The size of the field varies in relation to the emotional and physiological state of the animal: when a cat is wounded or frightened the area is large, whereas when the cat is peaceful the area is much smaller.

The territorial fields are connected to each other by invisible paths that the cat organizes during its different exploratory activities. The stabilisation and functionality of the set of fields in the territory are ensured by precise signals that constitute territorial communication. Vocalisations, postures and visual and chemical signals enable the cat to “orientate itself” in the environment and at the same time, provide a means of communication with animals of the same species.

As far as concerns visual and chemical signals, there are territorial signals, signals of identification or familiarity and alarm signals. The territorial signals are scratching and urine spraying. Scratching is primarily a method of communication through the combination of visual signals (the scratches) and olfactory ones (the pheromones secreted by the interdigital glands). Scratching is performed, independently of the presence of conspecifics, in strategic places such as clearly visible vertical supports. The scratches are made from above downwards with the front paws and with the animal in a particular posture (extended limbs, erect back) called the “underscoring posture” because it highlights the presence of a pheromonal message. The scratches indicate to other like animals that there is a habitual occupant of that territory and encourage conspecifics to avoid the marked zone. They are generally made near the fields of isolation, hunting (or feeding), elimination and the passages between the inside and outside of the house.

Urinary marks are made by leaving a spot of urine, with a diameter of 10-20 centimetres (visual signal) at a height of about 30-50 centimetres from the ground on a vertical support. The marking is characterized by a specific behavioural sequence which differentiates it from other types of micturition: the cat sniffs the place to make the spot, then remains standing (it does not hunker down), moving its paws in alternation (“kneading”) and, with the tail held vertically and quivering, makes the spot. Subsequently, through the Flehmen, it smells the pheromones deposited. The posture described is called an “underscoring posture” because it highlights the presence of a pheromonal message.

Every change in emotional state, whether positive (sexual excitement) or negative (anxiety related to the presence of a new cat, fear) can provoke this behaviour. Reactive-type urinary marking indicates the presence of a habitual occupant of the territory and is carried out at the intersection between a passage (path) and an activity field, while sexual-type marking is performed near exits to the exterior (doors and windows) and is often accompanied by vocalisation.

Urinary marking is performed by both males and females; only the frequency of the behaviour is different, being higher among males. Orchidectomy and ovariectomy can only inhibit the onset of the behaviour if performed before puberty. Urine marks are emitted sporadically and clumsily by kittens during the phases of playing and “fighting” around the age of 4 weeks and then this behaviour disappears from the cat’s ethogram until puberty. The definitive urine marking behaviour, which will last lifelong, appears after the hormone peak. “Late” sterilisation can cause a decrease in the frequency of marking and reduce the intensity of the scent emitted by males (since it is related to the deterioration of some aromatic compounds in the sperm).

The signals of identification or familiarity are formed of the “facial pheromones”. These are deposited by the cat by it rubbing the lateral part of the face, from the labial commissure to the hairless region of skin at the temples, against explored objects or living beings. In this way they become known and no longer represent a danger. The studies carried out so far have revealed the existence of numerous secretions formed of about 40 different chemical components of which only 13 are common to all cats. Some pheromonal combinations form a chemical message. P. Pageat identified at least five, named F1 to F5, which appear to be released in different circumstances, although only three combinations can be related to a precise functional situation:

• F2 is the secretion deposited during sexual excitation, above all by the male in the presence of a female in pro-oestrus or oestrus. The deposition of this pheromonal fraction precedes sexual-type urinary marking.
• F3 is the secretion deposited on objects in the environment in which the cat lives, particularly on objects along the paths jointing territorial fields. Deposition and subsequent perception of this fraction produce a “reassuring” effect and decrease the probability of the appearance of fear-related behaviours. Furthermore, the F3 fraction encourages feeding and exploratory behaviours while it inhibits reactive-type urinary marking and, to a lesser extent, sexually related marking.
• F4 is the secretion deposited on conspecifics of the same clowder and on animals and humans which make up part of the “family group” (allomarking). The deposition of this fraction inhibits aggressive behaviour related to irritation or protection of the territory and promotes the development of relationships within the group.

The alarm signals are pheromones secreted from the anal sacs and glands in the plantar foot pads. The perception of these scents causes reactions of avoidance and flight. They act on both the individual that discharges them and on conspecifics; they may also be a form of communication between species.

PHEROMONE TREATMENT: THE USE OF SYNTHETIC PHEROMONES

Pheromone therapy initiated in the 1990s as a result of the research by Pageat on the relationship between the F3 facial pheromones and urinary marking. At first the facial pheromones were collected by rubbing the cat’s face with a gauze and were then placed where there were urine spots. The results obtained encouraged Pageat to isolate the F3 fraction and create a synthetic analogue, which was marketed for administration via a spray or diffuser.

The veterinarian can use pheromone therapy without pharmacological support or in combination with psychoactive drugs. The F3 facial fraction of pheromones can be used when adopting a kitten, painting the rooms of a house, moving home, going on holiday in a flat or hotel, introducing a new animal, when a baby is born, or when a guest arrives; a pheromone diffuser is placed in the room most frequently used by the kitten/adult cat. The product should be left in place day and night. The rooms can be ventilated even several times a day because the pheromones remain in the environment in any case. During hospitalisation of a kitten/adult cat in a clinic or the subsequent period of confinement in its owner’s home, synthetic pheromones can be administered by diffuser or spray to reduce the patient’s stress and restore feeding and exploratory behaviours.

To facilitate a clinical examination, pheromones can be sprayed on the examination table (previously washed with soap and water) about 20 minutes before the arrival of the patient (Fig. 3 [10]). This can be done as the veterinarian completes the preceding clinical examination. Furthermore, pheromones can be sprayed into a carry box before every journey in a car. The product must be sprayed into the carry box (one or two puffs) in open air (in order to facilitate evaporation of the alcohol solvent) about 20 minutes before the animal is put into the box. It is also advisable to spray the pheromones into the car and places where the carry box will be put. On returning home, the carry box should be washed with water and neutral soap and left in the house, open and available to the cat. By doing this, the cat will not associate it with “a terrible experience” and will not transform car journeys into “a rodeo".

Following the appearance of signs such as urinary marking, increased vertical scratching, loss of appetite, reduced desire to play and decreased interest in interactions, the choice of treatment depends heavily on the diagnosis (nosographic, functional and contextual) made during the behavioural examination. Pheromones cannot be considered a panacea: the disappearance of the patient’s signs presupposes the introduction of appropriate cognitive-relational therapy.

Summary of the use of synthetic pheromones in the cat. 

Suggested readings

1. Arpaillange C. et Mège C., (2000), “Texte de conferences”, Scuola di Specializzazione in Patologia del Comportamento del cane e del gatto, Tolosa; Bradshaw J. W. S. (1996), “Il comportamento del gatto”, Edagricole, Bologna;
2. Colangeli R., Fassola F., Giussani S. “L’uso dei feromoni nella terapia comportamentale del gatto”. Rivista di Zootecnia e Veterinaria, febbraio 2003: 35 – 64;
3. Dehasse J., (2001), “L’educazione del gatto”, Alberto Perdisa Editore, Bologna;
4. Galimberti U., (2000), “Dizionario di psicologia”, UTET, Torino;
5. Pageat P., (1998), “ Pathologie du comportement du chien “, 2 éditions, Edition du Point Vétérinaire, Maison Alfort Cedex ;
6. GECAF, (2001), Cours de base de GECAF ;
7. Giffroy J. M. (2000), “L’éthogramme du cheval et l’éthogramme du chat”, Scuola di Specializzazione in Patologia del Comportamento del cane e del gatto, Tolosa;
8. Leyhausen P., (1994), “Il comportamento dei gatti”, Ethologica 1, Adelphi edizioni, Milano;
9. Mège C., (2000), “Troubles de la territorialitè chez le chat”, Scuola di Specializzazione in Patologia del Comportamento del cane e del gatto, Tolosa;
10. Pageat P. (1997), “La communication chimique dans l’univers des carnivores domestiques“, Le Point Vétérinaire, vol. 28, n° 181;
11. Pageat P. (1999), “Les phéromones d’attachement “, Pre – congrès Mondialvet, Lyon;
12. Pageat P. (2002), “Il comportamento del gatto in condizione di sovrappopolazione “, 44° Congresso Nazionale SCIVAC, Milano, maggio 2002;
13. ZOOPSY, (2001), “ Le comportement du chien et du chat dans la pratique quotidienne “, Module prévention et éducation, Ecole Nationale Vétérinaire de Lyon ;
14. ZOOPSY, (2001), “L’anxiété “, 3 – 5 octobre, Poitiers.