by Peter Neville
Clinical Professor, Dept of Veterinary Medicine, University of Miyazaki, Japan
Adjunct Professor, Dept of Animal Sciences, The Ohio State University, USA
Centre of Applied Pet Ethology (COAPE)
PO Box 6, FORTROSE,
Ross-shire, IV10 8WB, UK
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Studies of weaning in mammals have often concentrated on feeding behaviour and the energetics of changes in nutrition, from feeding on milk to more energy dense solid food provided by the mother. The critical behavioural changes that occur around the time of weaning are often glossed over as the acquisition of ’independence’ for the young, implying an increasing drive for initiation in social encounters and greater self-determination along with competence in the discovery of food. However, the acquisition of survival skills and social abilities is governed by the mother, notably manipulating the withdrawal of milk and directing access to solid foods in teaching her offspring new forms of behaviour. Cats, while sometimes social, are nonetheless destined to be solitary and non-cooperative when hunting and so kittens must develop prey handling and hunting skills quickly in order to become nutritionally self sufficient by about 14 weeks of age, and certainly by the end of the juvenile phase at 18 – 20 weeks of age when they usually leave their mother or are driven out from her home range so they then don’t compete with her for its resources.
Maternal Manipulation Of Feeding Behaviour: The Effects of Frustration
The behavioural and emotional effects of frustration in terms of reinforcement of behaviour patterns have been summarised into stages. Stage 1 considers the continuous reinforcement of the approach behaviour of kittens towards their mother during the first 3 weeks or so of life. Motivated to seek their mother’s teats through instinctive rooting reflexes, the kittens’ or puppies’ approach behaviours towards her are always rewarded by the provision of milk at the teat or warmth and comfort in her fur. Later, behavioural responses become reinforced through the process of classical conditioning in that many unconditioned signals of sound, scent, vibration, and temperature etc of the arrival or presence of the mother become paired with approach behaviours and reinforced by the same rewards. The mother simply makes herself available to her young in the nest usually leaving them only to feed and eliminate.
At the onset of weaning, Stage 2 of a reinforcement sequence of new behaviour occurs. At 3-4 weeks of age, the kittens and puppies have now become capable of more varied and coordinated behaviour in their efforts to obtain food from their mother. With their increasing size and complexity, their energy demands increase rapidly and she cannot produce milk in sufficient volumes to sustain their growth, body temperature and increasing levels of activity. The young are forced to become active rather than passive in their efforts to derive food from their mother. Their approaches become increasingly vigorous, which is uncomfortable for her as their milk teeth become painful to endure for long periods at the teat.
The mother now needs to leave the nest more to feed in order to produce the greater volume of milk required to sustain a rapidly growing litter and her return is increasingly marked by greater activity as the young demand to be fed. On presentation of the same stimuli associated with the mother’s arrival at, or presence in, the nest, the young repeat their earlier approach behaviours but find that access to the teats is now no longer always guaranteed. For the first time, the young start to experience rejection by their mother, finding that she moves away from them after short periods of feeding and, in the process of standing up to do so, may even cause them to drop painfully to the ground, or she may drag them a short distance out of the nest, as they desperately try to maintain their hold. Persistent young may even be firmly nosed away from the teats and all such rejection has the immediate effect of causing frustration, this is likely to cause a kitten or puppy to increase its vigour in attempting to gain the desired reward of food, of which there is insufficient now to sustain the whole litter.
Motivation for change in their feeding and social approach behaviour is now at its peak and the mother will use their hunger to shape their responses into new forms, the desirable elements of which will be reinforced by the provision of food and social contact. Their approach behaviour meets increasingly with ’non-reward’, as their mother may not always allow them access to her teats, or curtails their sucking before rolling away or moving off satiates them. This causes frustration and an invigorated repetition of their previous approach behaviours. But their frustration at failing to gain access soon causes them to experiment with different approaches in their effort to derive food. At the same time, the mother cat starts to introduce new signals of a particular call and visual presentation and movement of stunned rodent targets, indicating their availability as a different, solid and more energy-dense food source. She will also bat dead rodents with her forepaws to move them in front of her litter in order to stimulate their instinctive interest in small moving targets (Stage 3 of the sequence). In ethological terms, their investigative instinctive interest in the new food sources can be viewed as improved behaviours that become conditioned to the new signals given by the mother. In this transitional stage, the young still approach the teats but only receive sporadic milk feeds and only for as long as an adequate daily intake of food must be assured for them by their mother. In times of short supply of prey, this process will be delayed and she will continue to allow them to suck until more solid food can be provided. (Breeding seasons have usually evolved to coincide with seasonal highs in prey availability). The greater rewards of more energy-dense food for the new bigger, better, faster responses to the signals of the mother’s arrival at the nest ensure that the kittens’ new approach behaviours are rapidly reinforced.
The development of these new behaviours thus represents a crucial stage of behavioural shaping at weaning, but the process is clearly not without risk. Having become firmly conditioned in a Pavlovian manner to expecting food for approaching the teat as a neonate and in the transitional stage of development, and then exposed to the frustration effects of non-reward engineered by the mother for the same approach behaviour at the onset of the socialisation stage, the young may now begin to anticipate the prospect of non-reward. They may avoid rather than approach their mother when she signals her arrival. At the same time, they retain a strong motivation to approach her as a result of previously reinforced success at gaining food and so they could be left in a state of approach/avoidance conflict as a direct result of having been rewarded and denied reward for the same behaviour. It is at this stage, defined as Stage 4, that the mother must relieve the state of conflict in her kittens by clarifying the signals of reward and non-reward, and so encourage them to persist with the new behaviours that lead to the rewards of more energy-dense food that their growing bodies demand. To achieve this, she must rapidly decrease the frequency of reward for approaching to feed at the teat and consistently offer signals of non-reward such as standing up, rolling over and walking away, in response to such approaches. At the same time she must ensure that signals of the prospect of the provision of prey occur at sufficient frequency to become consistently associated with her arrival or demonstration of those signals.
The resolution of conflict at Stage 4 of the weaning process through clear signalling decreases the intensity of the approach/avoidance conflict of the kittens’ view of their mother. The new desired approach behaviour is reinforced by the primary reward of internal emotional relief resulting from the prospect of new but predictable feeding rewards, and secondarily, by the feeling of success at the acquisition of those rewards. This is, therefore, a two reward learning process, such processes being particularly resistant to extinction and therefore facilitating the firm establishment of social and food acquisition skills as the young approach a more independent phase of life. The behaviour changes engineered by the mother during the process of nutritional weaning reinforce the responses of the growing kitten to its emerging innate motor patterns of behaviour of prey chasing, handling and killing that will ensure its survival as a self-catering solitary hunter.
Signals of non-reward in response to instinctive rooting and sucking behaviours are given by mothers of both cats and dogs at Stage 2 of the weaning process, but mothers of wild canids start to provide food by regurgitating semi-digested stomach contents. Thus they reinforce following and socially interactive, food-soliciting behaviour in their puppies, and so help them develop social communication skills. Queens on the other hand, direct their kittens towards dead or stunned prey that they drop nearby, usually accompanied by a specific call. The queen is clearly changing the signal of food availability by directing her kittens towards new solid food and away from her body, even though the other signals of her presence that used to announce the prospect of sucking are still in evidence. If the prey is still alive but the kittens show little or no interest in it, she may then kill the prey and represent it to them for eating, or then offer her teats for sucking to ensure that they continue to be fed. But with repeated offerings of prey, the kittens soon begin to show more interest, practise their instinctive prey chasing and handling behaviours and then quickly go on to consume their prey once it is dead. This occurs as the kittens learn to recognise the dead prey as a source of food. Ultimately, they do not have to communicate with their mother to gain this potential food source, and respond with other instinctive behaviours to the movement of stunned prey or in starting to ingest dead prey. The mother cat may encourage physical investigation of the stunned or dead prey by moving it with her paws, but the kitten’s responses are nonetheless instinctive rather than coordinated learned behaviours and are not mediated through new forms of direct social contact with her.
Whether wild, feral or domestic, from the end of the weaning period at about 6-8 weeks of age, kittens are increasingly refining their independent prey detection and stalking behaviour through individual excursions away from their mother. They practise on moving inanimate objects such as wind-driven leaves and then on live, moving but largely inedible insects and worms and, finally move on to pursuing rodents and birds. The reinforcement of their hunting behaviour occurs not through the rewards of satiation of appetite in consuming their prey but in successful capture of the moving targets that initiate their predatory behaviour sequences. Hence the mother has no role in reinforcing or ensuring the success of their hunting behaviour once her kittens start to practise away from the nest. It is a process fraught with risk, and most kittens born to feral or wildcat mothers do not survive to reproductive age, despite the precision of their evolution as hunters and development of predatory skills from a very early age. Most kittens born to pet cat mothers in our homes however, are destined to live a nutritionally subsidised and protected life and will not have to hunt to survive. Hence the majority do indeed survive into adulthood and become pets themselves, even if in the western world the price is to be sterilised for ease of management for their owners and to help prevent what would otherwise be a massive population explosion under such nurtured conditions!
This article is a revised version of scientific papers by the author to focus on kitten development and has been produced especially for the society free of charge. Input is also taken from course notes from the renowned National Award Level Diploma offered by COAPE in ‘Practical and Theoretical Aspects of Companion Animal behaviour and Training’. Details of this and COAPE’s many other courses in both feline and canine behaviour and behaviour therapy, such as Think Cat (1 and 2) and You and Your Cat can be found atwww.coape.org/courses.html
Note from OCA Club Secretary: Please note that this article may not be reproduced or copied without the written or e-mailed permission of the author, although he’s a generous kind of guy and is usually very happy to give such permission for the purposes of improving feline welfare and understanding, so please do contact him first on firstname.lastname@example.org mention the Oriental Cat Association.