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Studies on Temperament Inheritance

Written and presented by ©Libbie Kerr, October, 1997.  

The study of temperament inheritance goes to Adam and Eve. Was it in her nature to want more knowledge? Or in his nature to take a bite right along with her? Were we genetically set up for this scenario or was the environment influencing us? Interesting questions pondered throughout the history of humankind... and one that we are just now beginning to actually “sink our teeth into”.

Temperament has long been my primary interest in Bengal cats: the idea that one can influence temperament with breeding (nature) and influence the individual with the way it is raised (nurture). Science in the last few years has come up with cloning, DNA splicing, and all sorts of knowledge and information that reveal part of the answers and leave us with more questions. While a lot of information credits our genes with who and what we are, there is an equal amount of information pointing to the way the individual was raised (nurtured) has an equal play on how the individual develops.

Daniel Estep offers a clear definition in his “Ontogeny of Behaviors”.

One way to conceptualize how genes influence behavior is to think of genes setting the limits on how much of a given behavior occurs or that genes set the relative thresholds for the occurrence of a behavior.” (p 24)

Translation: it merely sets the framework from which we work. He uses as his example the bloodhound which is born with a set genotype that effects his scent-tracking ability. This individual animal can do no better than the physical limitations to the sensory and nervous system that direct the behavior. However, it is influenced by the neonatal environment, nutrition, training, and experiences received.

This relationship of genes and environment is noted in several different references that appear in the bibliography (see attached) . Robert Sapolsy wrote in the October, 1997 issue of Discover magazine:

“A gene, a stretch of DNA, does not produce a behavior. A gene does not produce an emotion, or even a fleeting thought. It produces a protein. Each gene is a specific DNA sequence that codes for a specific protein. Some of these proteins certainly have lots to do with behavior and feelings and thought; proteins include some hormones (which carry messages between cells) and neurotransmitters (which carry messages between nerve cells); theory also include receptors that receive hormonal and neurotransmitter messages, the enzymes that synthesize and degrade those messengers, many of the intracellular messengers triggered by those hormones, and so on. All those proteins are vital for a brain to do its business. But only very rarely do things like hormones and neurotransmitters cause a behavior to happen. Instead they produce tendencies to respond to the environment in certain ways.” (p 42-43)

So again, we are told that genes produce the tendency, not a given response but a tendency to respond in a certain way given a set of circumstances.

I will include here only a little more DNA and chromosome information for it can get very complex and interwoven, which, by the way, is physically descriptive of what DNA looks like.

A chromosome is made of DNA, a long string of it and we use letters to code for genetic information. We used to think that Gene 1 would cause a certain response and then Gene two would take over... but while Gene 1 might specify the construction of insulin in the pancreas, Gene 2 might give your eye color and Gene 3, active in neurons, “might make you more sensitive to aggression-provoking stimuli in the environment.” (p 44 Sapolsky)

Again, pointing to the fact that genes give a tendency toward a certain behavior.

There are several other sources who support the idea of genes giving a tendency to a behavior. It is this tendency that we are looking at in our inheritance studies. So, we look at the environment and see the response.

“The biological factors that genes code for in the nervous system typically don’t determine behavior. Instead they affect how you respond to often very subtle influences in the environment.” (p 43, Sapolsky)

But why should the cat fancy be interested in this documentation? The cat fancy at large (not just Bengal cats) has long had anecdotal information on genetic inheritance of temperament... and now with the pressure of limiting breeding both at the federal and state level, laws and ordinances being passed, and a general move against breeding due to animal overpopulation, finds the idea of temperament inheritance and proper socialization a major reason for pedigreed cats. The Bengal cat opened the door for the cat fancy by putting temperament in the forefront on the standard. Leading the way to a new and more complete understanding of companion animal breeding.

“Temperament must be unchallenging. Any sign of definite challenge shall disqualify. Cat may exhibit fear, seek to flee, or generally complain aloud, but may not threaten to harm. Bengals should be confident, alert, curious, and friendly cats.” (TICA Standard, ACFA Standard)

This shows the following:

  1. The fore sight of the standard to see the future of pedigreed cats. Temperament.

  2. That temperament was accepted as being affected by selection of breeding pairs.

  3. Recognition of breeder responsibility to select for temperament in breeding.

Now why, you might ask, is the future of cats in general linked to temperament? Because it is one of the strongest reasons for breeding pedigreed cats that are mainly used as companion animals. The ancestry of pedigreed cats is known and more can be known by simple follow through.

“Behavioral traits should be considered heavily in breeding decisions. It is a disservice to the public to breed animals that look good but behave poorly. Breeders who are specifically breeding animals for companion purposes have an ethical obligation to breed and place animals with stable temperaments and with low risks of aggression, fears, and other behavior problems.” (Daniel Estep, 27)

“The three major species of companion animals - horses, cats , and dogs - are kept because people are attracted to their behavior. Yet in breeding these animals, behavior receives considerably less attention than morphological characteristics.” (Benjamin Hart, 232)

James Serpell in his book, The Domestic Dog, offers the following on dog breeders, which could easily read “cat breeders”. p. 62

“There is a need to further document the genetics of canine behavior. Dogs are usually bred by people who have no training in either behavior or genetics despite having a wealth of hands-on practical experience. If canine behavior is to be modified appropriately, then breeders will need to be able to understand the consequences of their actions, both in term of breeding as well as husbandry and training. They will need to be able to make sense of new behaviors evidence and, increasingly, they will want to know the extent to which behavior can be predicted early in life and, if so, how early and by what means.”

All of these are telling breeders to be aware of temperament. ... luckily for us this is demanded by our standard.

But with the Bengal we have the added dynamic of the non-domestic and the domestic.... so what is domestication as seen by behaviorists?

“The biology of domestication. Domestication is the result of two interwoven processes, one biological, the other cultural. The biological process resembles natural evolution in that the parent animals become reproductive isolated from the wild population and form a small founder group, or deme, that will at first be very inbred, and which will then undergo a process of genetic drift. Over successive generations the domestic “species” will multiply in numbers and will be genetically changed by natural selection in response to factors in the new, human environment. (p. 15, James Serpel)

Domestic animals appear to possess few behavioral patterns that are not found in the wild ancestors. But domestication does imply genetic control over behavioral as well as physiological and morphological characteristics. By domestication we have accentuated some traits and reduced or eliminated others rather than create completely new traits. Thus, domestication has affected behavior quantitatively rather than qualitatively.” Benjamin Hart, p. 228

With the Bengal cat we are taking the Asian leopard cat genes and mixing them with domestic. This combination creates a dynamic that is in part genetic and in part environmental and in large part unknown.... so our Bengal cat breeding programs are a perfect place for the collection and understanding of temperament. We have two very different observational behaviors in our root species:

The Asian leopard cat:

  1. Shy of human contact.

  2. Seeks to flee.

  3. Avoids confrontation.

Bengal cats

  1. Confident

  2. Curious

  3. Friendly cats

  4. Alert

So how do we get from one point to the other?

James Serpel in his book The Domestic Dog speaks of changing the

“perceptual world. this means that, whereas a degree of perception combined with quick reactions to stress are essential for the survival of an animal in the wild, the opposite characteristics of docility, lack of fear and tolerance of stress are the requirements for domestication.” (p. 15)

The very mechanism that allow the Asian leopard cat to survive in the wild are what we do not want in the domestic Bengal cat.

The area of inheritance in feline behavior is very new... but I will go over the different studies that have possible relevance to the inheritance in cats.

1. Study of sire. There is a study (still in progress) at Cornell on the effect of the friendliness of the sire on the kittens. Sires were selected because you can have several litters of kittens from the same sire at the same time thus allowing the information to be collected much more rapidly,. What I find interesting about this study is that the conclusion that sires have strong impact has come from a fairly small sampling. 

Daniel Estep, p. 23

“The ... paternal effect on the friendliness of domestic cats toward humans. Cats in two different colonies were rated on their willingness to approach and make contact with humans. Examination of breeding records showed that friendly males were more likely to have offspring that were friendly. What led these researchers to suspect genetic influences was that the offspring never had contact with their fathers and there was no relationship between the mother’s friendliness and offspring friendliness.”

Michael Mendl and Robert Harcourt: pg 49

“Evidence supporting an effect due to genetic variation comes from the work of Turner et al. (1986). Using observer ratings of kitten ‘friendliness', Turner and his coworkers demonstrated that one factor which helped explain the variation in ‘friendliness’ scores was kitten paternity. Kittens of different fathers differed significantly in their ‘friendliness’ scores. Since the kittens never saw their fathers it is likely that genetic factors mediate this effect.”

Halliday and Snowdon,

...demonstrate that offspring from a particular male are reliably different from those of another particular male: variability of the trait ‘friendliness towards human’ is at least partly explained in terms of who the fathers were.”

One would conclude that the sire has a stronger influence on behavior than the dam... but I have not found that to be true in a test case.. Anecdote: (that is your warning)

 I was curious about how long it would take (generation wise) to breed shyness out. I had a  female three generations from the Asian leopard cat, very shy, could not even be handled without exhibiting extreme fear, bred her to an extremely personable male. I got one female kitten from her that I placed with a very friendly outgoing, interactive later generation Bengal cat female. The kitten was treated the same as all the others in the adoptive mom's litter and, until 5 weeks of age, I noticed no difference in behavior.

At 5 weeks, the kitten would watch as the others ran to me, at six weeks she ran to hide. When she was old enough I bred her to an extremely sweet, male that produces wonderful temperaments.... and I brought the offspring to a show. The offspring did very well, not showing signs of fear.  The reason I share this with you is that this is from the female side.

2. Study of the Dam. There are several studies done on the nurturing of the kittens by the mother, which has a marked impact, but the problem is discerning how much is genetic and how much is environmental. The influence of the mother falls under the area of socialization that I will present at a later date. There is one study that clearly shows a behavior characteristic inherited through the mother. (Bonnie Beaver, p. 183. )

A few cats, in addition to using the regular forms of drinking, have been observed to drink by dipping a paw into the water and then sucking moisture from it. If this occurs when fresh water is being added, it may represent a play behavior. However, this behavior has been seen when fresh water was not a factor. In one colony a female cat displayed this behaviorism and later so did a female kitten, a daughter of one of the first cat’s littermates. Paw dipping, may, therefore, be a genetic trait because the kitten’s mother was not a dipper and observational learning could not have occurred.”

Although paw dipping does not equate ‘friendlier to human contact’ it is one of the few comments that connects behavior from the mother, all the other studies have environment and socialization base. One can begin to see the problem of working on female genetic inheritance when the line is not so clearly drawn as with the males who are not around their offspring and you can have the influence of several different litters at one time to observe.

3. Breed characteristics. There are several studies on breed characteristics and an excellent book by Michele Lowell, Your Purebred Kitten in which she goes through each breed and evaluates them. This has long been assumed and judges make comments on the nature of cats all the time according to the breed. Mendl and Harcourt, p. 48.

“... a survey amongst cat show judges found that these judges, with their wealth of experience of many cats and many different breeds, reported remarkably similar and distinctive breed characteristics. Siamese cats, for example, were reported to be the most outgoing with strangers and the most demanding of attention and affection. Their vocalization were often described as similar to talking. On the other hand, the Russian Blue was general reported to be shy and withdrawn. A variety of other breeds were described in this way, most in agreement with the popular literature. ... despite their (the judges) characterization of different breeds, (they) were emphatic that within breeds there are large individual differences.”

There are many more studies done on breed characteristics of dogs than on cats. Dogs have long been seen as trainable, because of their dominant (Alpha) dog structure compared to cats who have a very loose social structure. Because dogs have been bred over the years to specifically fulfill certain tasks for their owners they have a clearer breed characteristic. This goes also for the physical... there is a much greater difference between a Great Dane and a Pug than between a Persian and a Siamese. Though the difference is evident, the amount of difference is much less. Behavioral studies show that certain traits are sought in a dog used as a guide dog and say one used for hunting or racing. I will not go into detail on these studies, and there are many (see bibliography) and concentrate instead on why cats are seen as less “trainable”.

Bonnie Beaver, p. 92

“Social animals species have fairly well-defined dominance rankings to minimize agonistic behavior between individuals. A threat display by the dominant animal leads to submissive displays by the subordinate. Because the cat is not highly social, there are distinct differences in its social orderings. In groups of cats, such as in homes or colonies, the pattern of social order is unique. One male assumes relative dominance based on territorial ownership... Aside from this dominant male, other cats do not differ in rank, which indicates the lack of a complete or stable hierarchy.”

What this means to us is that... in order to train a cat you have to use very different methods than with dogs... Cats will not submit , but they can be bribed!!!

This information is relevant to why cats have not been studied as much or for as long as dogs. They were always seen as a more passive social animal. Because of this, a majority of the remaining studies sited will have to do with other species and the conclusions we can draw are indirect.

4. Hair Color Genetics. This is a fairly complex concept that comes from some observation and from conjecture. Melanin is what colors the hair... and behaviorist surmise: (Bonnie Beaver, 102)

“Epinephrine has the same metabolic pathway as the pigment melanin, and the same precursor is need for synthesis of both. Genetic manipulation of coat color could then be useful for breeding in or out certain behavioral characteristics such as fear and aggression.”

Helmut Hemmer, Domestication

“The coat color of a mammal is related to the basic level of its activity, its reaction intensity and its environmental appreciation. The reason for this is probably to be found in the fact that up to a certain stage the pigments that determine color - the melanins - and the catcholamine group of neurotransmitters that are to a large extent the basis of the information processing system share a common biochemical synthesis pathway. Selection of certain coat colors can produce a behavioral change with a corresponding change in the stress system either towards attenuated behavior and increased tolerance or in the opposite direction. Combinations of the alleles of single color genes that deviate from the corresponding wild-type increase or alter their effect on behavior. It follows that the strategy of selecting and combining certain coat color types can produce direct effects on domestication.”

Clyde Keeler in the USA has done studies on laboratory rats, ranch mink, and foxes. I will summarize some of his studies which are often observational. He was visiting a fox farm. (M W Fox, The Wild Canids, 399)

All the foxes near me were ambers with an occasional glacier. Farther away I could recognize a pearl or a silver and in the far distance a few platinums. Red foxes never show themselves like this, although they sometimes find their way into the range - nobody knows how.”

From this observation , Keeler began a lifelong work in coat color and behavior genetics. “Quantitative determinations of behavior, size development and biochemical parameters in ranch foxes were the most significant of Keeler’s studies” The colors are as follows: (Fox, 123)

The animals used were wild red foxes, silver foxes (that is the non-agouti black animals with a silver allele): and pearl, a breed color that combines the non-agouti allele with a dilution allele (blue), amber which carries the brown allele of the piebald allele and for the non-agouti (black) allele of the agouti locus. The offspring were bred further and the behavior of the wild-colored animals (ones in which the agouti allele of the agouti locus was expressed) and the bland animals (with the non-agouti allele) was compared. The non-agouti black rats proved to be less timid and less aggressive and showed more confidence in new situations by exploring them more quickly. Albino rats in these experiments proved to less ready to react to olfactory stimuli than others.”

Keeler also studied rats. (Helmut Henner)

Keeler first crossed brown rats with albino laboratory rats that , in addition to the albino allele, were homozygotes for the Black locus in addition to the two former alleles, and glacier, animals in which a white allele is add to those for the amber coloring.”

This is a very elaborate study that measure adrenal gland response, size, and many other areas statistically. In essence, the amber fox behaved with the least amount of fear, and is also the largest of the fox in size. The more heterozygous for color the animal was, the “less fear it exhibited”.

Keeler went on to study mink and found that color made a difference in size and in behavior. Again, “the heavier, animals (pastel mink, ie. light brown) had smaller adrenal glands and were characteristically less active and aggressive, that is, they behaved more tamely.” (Hemmer)

Zebras have been studied, Hemmer, (p. 127) sites a study done by Eckehard Eich and Elisabeth Reichert.

“The activity values of the ‘whites’ coincide with those of domestic horses and donkeys, while the values of the normal-colored zebras differ distinctly from them. ‘White” (albino) zebras are less shy of observer in the enclosure than others.”

There are many more studies than I can even begin to note here... however, it should be pointed out that this is being studied and is not a known fact. It could very well be that hair color does indeed follow the same pathways and neurons but I go back to the article in Discover magazine:

“...By now, I hope, we’ve gotten past ‘genes determine behavior’ to ‘genes modulate how one responds to environment.’” (p. 46.)

It would be a gross mistake to make broad range assumptions on behavior and color alone...

most differences in behavioral traits are influenced by many gene, not just one or a few genes which means that inheritance will likely be complex.” p.27, Victoria Voith.

Fox bred for friendliness. Helmut Hemmer, p. 161-162. Chapter on New Domestications:

“In the preceding chapters, three principles of domestication were elaborated. (1) Wild animals seem to be more suitable for domestication the easier they are to breed successfully in captivity in crowded conditions. (2) Individuals from a species to be domesticated seem to be more suitable the smaller their relative brain size within that species. (3) The selection and combination of certain coat color types can elicit direct domestication effects. With the knowledge of these basic principles, it must now be possible to undertake purposeful selective and combinatory breeding for new domestications which enable the transition period between the wild and the domestic animal to be traversed within a few animal generations, in contrast to the domestication of animals of past millennia that proceeded slowly over hundreds of years and involved a great deal of chance. A further possible approach is to select rigorously for the typical domestic behavioral syndrome, where the fundamental suitability for domestication seems to exist according to the first principle but no differences in the populations or individuals corresponding to the other two principles can be defined.

An excellent example of this approach is provided by a study on breeding a fox that resembled the dog in its behavior, undertaken by K.K.Belyaev and L.N. Trut. This experiment was carried out with silver foxes on a fur farm belonging to t the Soviet Academy of Sciences near Novosibirsk. Young foxes aged between 1 1/2 and 2 months were selected according to the criteria of their tolerance of hand-feeding, their reaction to be handled by humans, and their response to being called. In this way, after 15 years of constant selection, foxes were finally bred that came when they were called, tolerated being petted and picked up by humans, wagged their tails in greeting and barked on seeing humans; in brief, they behaved in practically the same way as dogs.”

The study goes on to say that the foxes changed their reproductive cycles to resemble the domestic dog and the hair color of some of the fox changed to one of a piebald. Black and white fox. (there is a photo which I had, and sorry I misplaced!)

5. Brain size. In wolves, the brain size is decidedly larger than the brain size of the domestic dog. (Michael Fox, Integrative Development of Brain and Behavior in the Dog.) This is also supposed to be true of cats, though I did not find a direct reference. What bearing this has on the inheritance of temperament is unknown, but it is supposed (again by Fox) that the domestic animal does not have the need for all of the survival brain area, and because it is not needed, it is somehow lost. It would be interesting to measure when this takes place in the development of the Bengal breed. If indeed the Asian leopard cat has a larger brain than the domestic, at what point does this change take place?

6. Domestic Behavior. I am going to touch on one final area. (I am beginning to see this as a book in the making!) The final area is one of: what we consider domestic behavior.? What is our ideal cat temperament? Well, obviously this is going to be individual in our decisions and in our descriptions... however, there are some common themes. Interactively or neoteny “juvenile behavior”.

James Serpell, p. 43.

The concordance of morphological and behavioral features found in the domestic dog is best explained by the evolutionary mechanism of heterochrony. Neoteny, in particular, may have played an important role in creating an organism suitable for domestication. Gould stated that’...general juvenilization of behavior permits a more gregarious society.’ Selection for juvenile behavioral characteristics would not only make for a more tractable animal, with juvenile care-soliciting behavior and lack of species recognition, but serendipitously would also retard the onset of dispersing motivations and adult inflexible motor patterns resulting in a tameable and trainable companion. “

Discover magazine, p.43, speaks of novelty seeking gene...

The same theme continues in other behaviors as well. The exciting (made-of-protein) receptor that apparently has to do with novelty-seeking behavior doesn’t actually make you seek novelty. It makes you more pleasurably excited than folks without that receptor variance get when you happen to encounter a novel environment.,”

Conclusion (temporary conclusion ... I do plan to go forward with this and talk about environment, sensitive periods in development, and optimal care giving to bring out the best in the individual animal.)

After reading many books, articles and attending seminars on feline behavior, it is evident that we are just beginning to understand the interplay of environment and genetics. As breeders and cat lovers we can go a long way in learning more about these animals that permit us to be a part of their lives by observing their behavior on their terms.

Leyhausen, says:

“As I see it, the question what is the normal behavior of an animal species in a particular situation that is “natural” to it may be posed in three ways.

  1. Which behavior patterns occur at all in the situation? ...

  2. What can I expect with (what degree of) probability?...

  3. How does the animal behave when everything goes smoothly? ...”

These questions give us some basis for our observations. I am working on collecting data on behavior using a quantitative approach in collecting data. That of handling kittens. Anyone wishing to contact me about this please do so... it is a work in the very beginning stages.

I will conclude with one final quote, substituting cat in for child.:

“It’s not just what happens to you that counts - it’s what you think happens to you,” says Kagan. “And it is inordinately difficult to figure out what a (child) cat is thinking. Until we devise ways to measure what is in a (child’s) cat’s head, we’re not going to understand the (child’s) cat’s environment. “

Karen Wright, Babies, Bonds and Brains. p.51

BIBLIOGRAPHY

Beaver, Bonnie: Feline Behavior: A Guide for Veterinarians. WB Saunders Company. Philadelphia.1992.

Fox, Michael: Integrative Development of Brain and Behavior in the Dog. University of Chicago Press. Chicago. 1971.

Fox, Michael: The Wild Canids: Their Systematic, Behavioral , Ecology, Evolution. Reinhold Co. NY.

Halliday,TR, Snowdon, Charles, editors: Animal Behavior. Bailliere Tindall, London, 1986.

Hart, Benjamin: The Behavior of Domestic Animals. WH Freeman and Co. NY. 1985.

Hemmer, Helmut: Domestication: the Decline of Environmental Appreciation. Cambridge University Press, NY 1983.

Leyhausen, Paul: Cat Behavior: The Predatory and Social behavior of Domestic and Wild Cats. Garland, STPM Press. NY.

Overall, Karen: Clinical Behavior Medicine for Small Animals. Mosby. NY. 1997.

Sapolsky, Robert: A Gene for Nothing. Discover. October, 1997. Vol. 18 No.10 p.40-46.

Sepell, James: The Domestic Dog, it’s evolution, behavior, and interactions with people. Great Britain, Cambridge University Press, 1995.

Thielig, Sidrid and Harald: Smart Cats: How to Understand and Train Them. Sterling Publishing Co. NY. 1994.

Turner, Dennis, Bateson, Patrick, editors: Mendl, Michael: “Individuality and the Domestic Cat.” in The Domestic Cat.: the biology of its behavior. Cambridge University Press, NY 1988.

Voith, Victoria and Borchelt, Peter, editors: Estep, Daniel “Ontogeny of Behaviors”, in Companion Animal Behavior.

Wright, Karen.: Babies, Bonds and Brains. Discover. October, 1997. Vol. 18. No.10. p.75-78.

 

SEMINARS

Ohio State University: Feline Behavior and Behavior Problems. 1995.
Cornell University. Feline Behavior, The Genetics of Temperament. 1996.
Ohio State University: Feline Behavior, Behavior Modification, 1997.

©Libbie Kerr

Terminology “Shared and Compared” edited by Libbie Kerr

“Tabby cat!” he said for the third time, “My cat is not a tabby cat it is a leopard!”

This comes up quite often, one need only flip open a magazine and see ads for “snow leopards”, “leopard spotted”, exotic, wild ... you name it, it is advertised somewhere. But what is the truth, at least as far as what is known in the cat fancy? All cats are tabbies. (see Bulletin, September issue) As to whether the pattern is a different gene is a good one... is the Asian leopard cat pattern gene an entirely new gene that is not described in the tabby literature? What can we hypothesize in the interplay of known domestic and unknown non-domestic genes? To begin... let’s go over the term “tabby”. (note: the drawings of the cats included in this article are very crude adaptations of the cats used in the book The Book of the Cat, who is not responsible for any resemblance the skins may have to the living....please note no cats were harmed in the making of this article) 

TABBY PATTERN:

DOMESTIC

The mackerel tabby pattern seen on the far left here, is commonly found on the domestic cats of today.   The broken mackerel, or spotted pattern is produced when the straight lines of the mackerel pattern are interrupted and the background coloring comes through. This pattern is seen on the domestic cats of the Egyptian mau and Ocicat.    The flow of the spotting remains extremely vertical and though the lines are not the stripes of mackerel they are aligned in the same manner.  

There are three readily accepted patterns of the tabby:

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agouti (Abyssinian or ticked coat)

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mackerel (striped)

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classic (blotched, marbled, bull’s-eye)

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Question as to whether independent gene:
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spotted

Spotted can come as:

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broken mackerel, vertically aligned

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broken classic, follows the bull’s-eye

DOMESTIC

On the left is the pattern referred to in the cat fancy as the classic tabby pattern.  Notice the bull's-eye effect of the pattern in which the swirls create a circular motion.   The pattern to the right is called a spotted or broken classic.  Again, you see that the spotting is actually an interrupted classic pattern.  The effect, if the spots are connected is to create a bull's-eye pattern.   Though many of the cats with this pattern also have rosetting, it is from the domestic rather than the non-domestic genes.  

These two are seen as being only a modification of the mackerel and classic genes and not an independent gene.

The difference the Asian leopard spotting throws into the mix:

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horizontality

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spotting on tail

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straight lines coming back on forehead instead of tabby M

Asian Leopard Cat Patterns

The drawing to the left shows the patterning on the Asian leopard cat.  Notice the strong movement to the horizontal and to the diagonal rather than to the vertical.  This movement is found in  the small  non-domestic cats: ocelot, margay, fishing cat, to name a few.     This is the pattern flow that the Bengal cat is working to duplicate.  Notice also the spotting found on the feet and tail, extremely hard to replicate. 

 

What follows are the arguments against there being a separate spotting gene... and this is before the Asian leopard cat spotting factor was thrown into the genetic mix

The Book of the Cat, p. 32

Tabby cats are so universal that it is probably no great surprise to learn that they are the basic type of cat - the “wild” type, from which all others evolved... Some breeders have suggested that there is a further mutation of the tabby gene, responsible for the spotted pattern seen in the such breeds as the Egyptian Mau, spotted Oriental Shorthair, and Ocicat. The existence of spotted patterns in various wild cat species some support to this suggestion, but most geneticists feel that in the domestic cat the pattern is due to a breaking up of the mackerel tabby’s stripes. The fact that there are many gradations, from mackerel through half-spotted to fully spotted, backs up this view. In fact, breeders of spotted tabbies must continuously select for well-defined spots, or mackerel tabbies result. Barring of the chest and legs persists in any event, and it is also difficult to maintain unbroken striping in mackerel tabbies. All this suggests that polygenes are involved.

In Robinson’s book, Genetics for Cat Breeders p. 151

The last form of tabby is spotted. The genetics of this tabby is unknown in the sense that it could be due to a distinct mutation or it may merely be a variant of the mackerel. The striping of the mackerel has a decided tendency to break up into short bars or spots. It is fairly easy to take advantage of the tendency and to develop a spotted tabby. Some really remarkable and beautiful spotted cats have been produced. The fact that these cats can be bred is consistent with either of the above explanations. However, the latter one is adopted because it is not politic to assume the existence of mutant genes unless these have been well founded. The almost continuous variation from mackerel to clearly spotted is indicative that the difference between the two is polygenic. Breeders would be wise to breed only from those spotted cats which come closest to their ideal animal.

Article, Cats September 1997 by Barbara French

Spotted. This tabby has been the subject of debate among students of coat color genetics. For a time, some believed the spotted coat resulted from a unique gene. Others have theorized more convincingly that the spotted tabby results from a mackerel or classic tabby pattern breaking up and forming spots instead of stripes. Other genes may come into play, but the general consensus is that the spotted tabby is a variation of the existing patterns, not a new pattern. In support of this theory, experts note some litters of spotted cats contain both spotted and striped kittens in very similar patterns.

A close visual exam often will reveal that a spotted cat actually is a broken mackerel or a broken classic. Broken mackerels tend to have smaller spots in more regular lines, while broken classics have larger randomly placed spots or classic “bull’s-eyes” on the sides.

How does this information apply to Bengal cats? For our purposes let us set aside the argument of whether the genes at work are separate, are mutations, or are polygenic in their effect on the patterning. These are all terms that are necessary to know to understood what the Bengal cat pattern is now and what it can become.

The characteristic one notices from the Asian leopard cat influence is that of horizontal patterning... versus the more vertical mackerel type patterning and the more classic type flow that follows the bull’s-eye pattern. In the Bengal cat the introduction of the spotting pattern from the Asian leopard cat has a different influence on the pattern... a horizontal flow or diagonal flow that is more directly tied to the Asian leopard cat pattern.

Is this a new gene? This question cannot be answered completely. What can be said is that there are polygenes that are influencing the patterning in ways other than what has been known in the past. If these are completely new genes separate from the domestic or just the genes influencing each other completely or incompletely is unknown. What can be seen is that there are patterns that are emerging that show polygenes at work, or partial dominance:

1. Incomplete marble and spotted or is it partial dominance of genes? This pattern can have strong outlines of the typical butterfly pattern over the shoulders, a strong vertical line immediately past the front leg and then spotting after the bar. Several of these have “stumped the judges” as the debate began of: Is it spotted? Is it marble? This pattern usually has large rosettes sometimes in the position of where the bull’s-eye would have been.

2. There is the diagonal pattern which has what might be a combination of the spotting from the Asian leopard cat, completely horizontal, and the mackerel vertical... a compromise is reached between the influence of the two and the diagonal is the result? Or is it a continuation of the diagonal shoulder markings on the Asian leopard cat?

3. The chaining that is appearing has a unique look that is not found in the domestic and seems to be effected by the rosetting whether coming from the marble or spotted horizontality. Is this the beginning or horizonally striped cats? Is this horizontal striping going to be seen as a fault like, spots running together according to the standard?

So the Bengal cat has patterns that are reminiscent of the: ocelot, clouded leopard, cheetah, etc.. In looking at the pattern only, not color, not rosette so much as pattern flow... look at your cats. Sketch them out on the blank Bengal cat hides and think about pattern only.... Study the pattern and ask yourself:

NON-DOMESTIC PREFERRED PATTERN ON THE BENGAL CAT 

The two pattern on the left show the difference in patterning from the domestic vertical movement to the horizontal patterning on the non-domestic.  Since the idea is to replicate the look of a non-domestic, selection should be based on those cats that take the breed closer to the ideal. There can be many variations of these patterns and there are! The  images show the pattern flow… the shape and type of pattern can vary a great deal within the pattern flow. I hypothesize that this is caused from a pattern interruption gene or polygenes. It will need further study to tell for sure what is happening. These genes are in the present gene pool and do not need to be brought in by further hybridization. Selection is the key… There are some Bengal cats that are 12 and 13 generations removed from the Asian leopard cat that suddenly have this horizontal pattern. As the saying goes: “Recessives are forever!” In the case of horizontal flow, that's a good thing!

 

Is the pattern:

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Mackerel?

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Classic?

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Diagonal?

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Horizontal?

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A combination?

 

What creates the “wild” appearance in the pattern... play “connect the dots” and link spots together... or flow with the marble pattern... is it truly a marble or more classic in its pattern? Marble implies a completely different appearance in pattern from the classic bull’s-eye... Are the rosettes created shaped in a mackerel pattern or classic pattern? Perhaps the term used for the “ideal” is the term horizontality. A pattern has this influence or not... this influence appears to be the one most closely linked to the Asian leopard cat genes.

So, in answer to the first statement... It is evident that the Asian leopard cat gives a different pattern from either the broken mackerel or the broken classic... Leaving us with the questions: Is this a different gene for tabby pattern and how do we work with it in our breeding selection?

Vocabulary words: source: The Book of the Cat

modifiers: polygenes that change the effect of major genes.

mutation: a change in a gene - due to environmental influences or a ‘mistake’ during replication - that results in an abrupt change is hereditary characteristics between two generations.

partial dominance: The occurrence of a pair of alleles where neither is fully dominant to the other, so that heterozygous individuals show mid-way characteristics.

polygenes (polygenic) Groups of genes, small in individual effect, that act together to produce bodily characteristics. See also modifiers.

The new Bengal cat words for the day:

horizontality: indicating the pattern flow that is reminiscent of the Asian leopard cat pattern. This flow can appear as connected spots in a chain, elongated spots following the horizontal, lines that flow horizontally and combinations. Also seen as a very strong

diagonal flow to the pattern... flows shoulder to hock (see drawing).

What it the pattern of your cat?  

Published in the Bengal Bulletin copyrighted by libbie kerr

 

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