Очень хорошая статья посвященная наследованию цвета у Биглей опубликована в журнале "Охотник на кроликов" (Rabbit Hunter Magazine) Дэном Лефгреном (Dan Lefgren, T-Times-V Beagles), одним из старейших биглеводов Америки. Приводится с разрешения автора. Написана в виде доброго и тёплого рассказа. К сожалению, только на английском - сейчас нет времени переводить. Может кто переведёт, а я помогу отредактировать?

A RAINBOW SPRING '06' by Dan Lefgren (an up date of an article previously published in Rabbit Hunter Magazine by the author)

The old houndsman eased into his lawn chair and settled back to enjoy the last rays of the spring sun. Ah! Finally, a chance to enjoy the spring warmth; a chance to pause life for a moment and escape the cares of a too long and busy day. This is sweet.

Sweet too quickly disappears when the violent thrashing of the lilac bush at the far end of the yard interrupts the spring spell. Now what! His ear is quickly focused on the thrashing bush; then his eyes capture the flashes of color close to the ground, at times the flashs obscured by grass long overdue for the first spring mowing. A smile of recognition. It is just another piece of spring. Beagle mom and pups playing hide and seek; run and chase among the long neglected landscaping and flower beds of some previous owner. What a bunch. So easy to spot each pup, for all six sport their own unique marked and colored suit. Their sire a fairly typical black blanketed tri-color and mom pretty much the same except for being more open marked and flashy, with her wide white collar and splashes of white along her sides and over her loin and rump. Yet, here were these six pups, now eight weeks, looking like they had fallen off a rainbow.

A quick string of loud yelps and more violent thrashing from the lilac bush forces the houndsman to his feet. Obviously a closer inspection of the ruckus is called for. The black blanketed male pup has trapped himself among the limbs and while in no danger, except that of being left behind by mom and mates, cries out for a rescue.

The old houndsman does what mom can not and quickly crosses the yard to pluck the youngster from his lilac prison; then hoists him skyward so that both he and pup might have a better view. He of the pup; the pup a wonderful aerial view of the ongoing mom and puppy Olympics. They watch. He smiling; the pup now wide eyed and squirming to be free, as the main mob frantically circle the yard with iris blossoms clutched in their teeth; the remainder crying full voice and tumbling after in hot pursuit.

Let's take a closer look at you young'en, while I've got you captured, and see what we can learn about you and also those young parents of yours. You've got some white on you. Just a bit more than dad; a bit less than mom. That's about the normal expectation from such a mating. You've got their black blanket and a bit of tan, the typical saddle pattern of the beagle. They all
have that, or at least all I've seen. Even those extremes with just a bit of white on feet and chest, that some might call black and tan, or those few we see with just a spot of color on an ear or back on the rump. Actually you're all what geneticists call double recessive for 'a^sa'. That's the recessive that gives the blanket, we'll use that ol' caret symbol to indicate the fancy superscript 'sa' for saddle, and besides this stuff is just between you and I and not a full academic presentation. A guy named Little back in the 50's (Little, C. C. (1957) The Inheritance of Coat Color in Dogs. Cornell Univ. Press) graded a bunch of beagles by the percent of white. By his scale you're about a 4 (like 40% white), your dad a 3 and mom around a 5. About 60 percent of beagles will be in that 30 to 50 percent white range while the rest will be split about equal on either side.

You've all got that white because you are all double recessive for 's^p'. That 'p' meaning piebald; the 's' for spotting. If you were dominant for self spotting or no white, an upper case 'S', you'd be marked like a German Shepard or some Airdale Terrier. They are a couple of breeds that also have the 'a^sa' recessive pairing for saddle pattern. Nope you're 's^p,s^p' just like all the rest of your breed no matter what their color. I see those tan hairs creeping up your leg and intermixed among the black. Your blanket is probably not going to be as big as dad's but a bit larger than mom wears. That black is dominant and the symbol is 'B'. Your parents both have it so you got that big 'B' from one of them or maybe both. We'll not know until you've sired your own bunch of rascals.

I see you've got some little ticks of tan on those white legs. Just a few, most won't notice, but that tells me you are dominant for ticking, a big 'T', just like both your parents. Virtually all beagles are, just some have more than others and it is more obvious if they have a lot of white and the ticks happen to be larger or more of them. Folks call them ticked hounds but actually you're virtually all ticked, it is just a matter of being able to see it and how dense it is. The more white the more obvious the ticking. The hound with no ticking, the double recessive, 'tt', they are the rarity. I doubt I've actually seen a half dozen that do not have any ticking on them anywhere. Yah, I know time to go. Don't jump. I'll set you down you generic plain label hound of the publics mind and TV ads. Oh, I'm just teasing you. Go play while I catch another of your mates.

A quick grab at the flash of white and brown movement behind a very damaged peony bush and a wide-eyed young lady is quickly lifted to the level of the houndsmen's eye. She briefly floats at eye level long enough for him to see the amber of her eyes and the red pencil eraser color of her nose before he cuddles her to the more secure position against his shirt. What a nice mahogany
colored blanket you have, missy. Some would call you a liver or red tri, others might say you're a Warfield Red as if all that have your color were descendents of that famous Field Champ grandson of Grays Linesman. Warfield Red died in 1957 and he's not in your pedigree so that mahogany blanket did not come from him.

The double recessive 'bb' that gives you your blanket is from another source and I don't know what particular old hounds passed it on up to you. One thing for certain each one of those little b's came from each of your parents. That tells me they both carry the recessive 'b', although both have black blankets making them dominant for 'B', the second of their pair at that locus must be the recessive 'b'. That makes them 'Bb'. The recesssive 'b' from each has come together to make you a chocolate drop. The books call that recessive color chocolate or brown even though the color can be almost a black-brown to a very light orange-red, and nose and eyes will vary in shade to go with the color shade of the blanket. I know, easy, we are burnin' daylight and you have much to do, so be about it and I'll take a longer look on another day.

OK! OK! Now who is this pair of heathens tugging on my shoelaces. Very small chance of hurting those ol' laces. Far too many pups have taken their best shots at them and devoured and shredded the ends long ago. OK, I'll sit down and you can scramble across my lap. Ouch! Time to take a snip off the end of those nails. Let me see your nose missy. You are a blue. You've got that slate colored nose and a lighter eye, your tan is also more toward the cream end of the color scale. That blue or dilution of the black that our eyes see is caused by those round black color granules being clumped together which leaves gaps along the hair follicle. It has the same affect on yellow. Many folks might not notice you're a blue and standing alone in a picture they might mark it off to differences of lighting or film. I've seen you standing next to a black nosed and blanketed hound, the slate color of your eye rims and nose tells me what you are. You've got a pair of recessives 'dd' that books call dilution and looks blue to our eyes. The dominant is 'D' and both your parents have that and then they each carry the recessive to make up the pair of genes at the dilution locus for them of 'Dd'. The dominant at the 'D' locus does not allow the color granules to clump. It's the recessives that gives you that slight look of blue to our eyes and you got one of the 'dd' recessive pair from each of your parents.

Hey! That's my shirt pocket. What's this grey ghost blanketed brother of yours doing? I've got him. I know what you are. Your nose is kind of slate and eraser along with some might even call blue eyes at this age. Those eyes will darken just as your blue sisters will as you both get older. That grey blanket, that others might call fawn or if worn by a Dobe they might call it Isabella or even silver if on a Weimariner, is genetically a lilac. I think that is the best term to use because it indicates the genetic blending of the chocolate (liver, red) and blue. That color actually tells me you are a dilution of the chocolate just as your blue sis is a dilution of the black. The genetics name for your color is lilac, regardless of the particular shade. They use that a lot to describe the same color in cats and other mammals. Imemtioned the clumping of the round black granules in each hair that makes a color look blue, well in you we've clumped the oval granules that make a blanket chocolate to make your blanket a lilac one. The symbols for the double recessive
genes of the lilac color are 'bbdd'. You got one 'b' and one 'd' from each of your parents. Now, get out of here. I've got to stand up. My old knees are locking up and you've added too many new rips to my favorite ol' shirt. It's been rescued from the trash twice and though my partner disagees, it, like me, still has a lot of mileage left.

Now I've got to catch and handle those last two. I try to follow the method of the packs and houndsmen of the past that required that if you pick one up from a litter, no matter what the reason or the age, then you have to do them all and for the same length of time. No playing favorites no matter what. Let mom and the pups sort out the pack order among themselves. They need to be socialized to humans but picking out a favorite pup and playing with it more than the rest can cause problems down the road. Always best if you pups sort out your own pack order and we masters and hunstman should honor it and not disrupt it. Helps keep rioting to a minimum and those fights where all gang up on one to push it down the order are less common.

Ah! Here's that opened marked male some might call a patch pup. Let's look at you. My what a lot of white. You're a pieblad grade 7 or maybe even 8. You've got a lot of ticking along with those few big patches, some with edges of tan. Your legs have tan ticks and the rest of you has little colored ticks, places quite dense. That shows me the extent of your blanket colors if you had a full blanket. The ticks will show up better as you get older. There are going to be a lot of folks call you a blue ticked hound. Let's see your nose and eyes to see if they'd be right. Yep, you're a true blue tick. You've got a slate colored nose and those kind of bluish looking eyes just like your blue sister. Many people see a ticked hound like you and immediately call you all blue ticks, but they are wrong in doing that. If you had the same black nose and dark eye of your black blanketed brother then you'd actually be a black ticked hound. Ol' Fred's last litter had one with an eraser colored nose and light orange ticks. She's a red tick or to be correct a chocolate or liver ticked hound. There isn't any reason that a lilac ticked hound might not also exist, but I've never seen one. Of course there would not be a place to mark the lilac color on the AKC registration form. Write it in the 'other' space and they will send it back. They really do not know how many possible
colors you guys come in. Most breeders will just mark blue and say so what. That is what I've always done, except once. They sent it back to me and told me to mark a box. I did. Who has time to fight city hall anyway. There are too many other things to do.

Marking the colors like that isn't right and it will confuse some one looking at your peds on down the road because they won't be able to figure out where that chocolate pup in their last litter came from. Heck, I've seen peds where both parents were marked as blue ticked but none of the pups colors were marked as blue. Genetically impossible, and it tells me the parents were or at least one was actually a black tick. All the offspring should have been blues from such a mating of two blues and some of them possibly blue ticked if they were open marked. Oh well, few things are perfect and AKC have had those colors messed up for decades. AKC will give you about 25 choices and many of those really just duplicates of the same genotype, only in a different order. The actual number of color choices and combinations is pretty easy to figure out and a lot of us have seen most of the 32 basic identifiable phenotypes. If we take into account the many different patterns, mixtures of hair colors and hair types that are also in the breed the numbers can get very very large. Many years from now having full DNA panels and the ability to read them might
change some of this, but it will not be quick in happening.

As I said before, virtually all beagles are ticked for it is the dominant, symbol 'T', it is just the amount, its density and the amount of white that lets us see it. When I look at old peds that say such and such hound was blue ticked, I'm always skeptical unless I've examined the hound myself. Far too many have wrongly marked the color without looking at the nose and eyes. Most of those so called blue ticks are actually black ticks. Makes trying to figure things out using old AKC peds tough at times. Always best to be cautious unless I've looked at those ol' hounds myself. Down you go. Where's the last one. She needs her share of time.

There you are sis. Just two colors. A tan and white, although many might call you red and white or if lighter maybe yellow and white, lemon and white or even cream and white. They are all just shades of yellow coat hair which is the genetic term for them. Those shades of yellow, we'll call it tan as most beaglers do, are due to polygenetic plus and minus modifiers of a bunch of
genes called rufus polygenes (plus darker, minus lighter). They effect the other colors also except of course the black and white, or at least our eyes can not see it's effect on the last two. Let's look at your nose and eye to see what color your blanket would have been if you did not have that recessive pair 'ee', that stops the primary color of your blanket from being expressed or extended. The 'E' locus is called the extension locus. The dominant is the one that allows the primary color to be extended or expressed. The recessive removes it. Your parents and all your littermates are dominant for 'E' as they all show one of the four primary colors (black, chocolate, blue or lilac) in their coat.

You have dark eyes and good black eye rims along with a very nice black nose. That means you'b be a black, white and tan if you did not have the 'ee' recessive pair. Shows your parents are carriers of the recessive and the genes at the extension locus for them are 'Ee'. You got one recessive from each to make your pair. If your nose had been slate or eraser, and your eyes a different color then that would have told me you actually were either chocolate, blue or maybe even lilac if your genes would have allowed your primary blanket to be expressed. Darn! You've got teeth like needles. Trying to amputate my mouse-clicking finger. Not fair. You might as well have taken a bite out of my voice box. Whoops! Suns about gone. Time to kennel you all up for the night.

Here puppies! Here puppies! With me! With me! Oh, sure under the porch. Mom why did you let them get back under there. I'll have to get the flashlight. Let's see. Which corner? There you are. Six pairs of eyes. Four pairs glowing green and the other two red. A Russian researcher back in the 30's figured that red eye glow was passed on as a recessive. What he failed to
discover is that the red reflection is directly connected to the 'bb' for chocolate. We know that now. This can be useful when trying to determine the color genes for an 'ee' (tan/white). Other recessives are rear dew claws and even some of the tail kinks that we see or hear about from time to time.

Your mom had rear dew claws, which I removed when she was 5 days old, so she has the recessive pair and half you pups have it also. Half having the characteristic is the normal expectation when a carrier, in this case your sire, is mated to a partner, your dam, that exhibts the particular recessive characteristic. Half you guys obviously have the same recessive pair as your dam although one of that pair came from each of your parents. The rest of you are all carriers of that same recessive gene and also the dominant which suppresses the expression in you. The dominant came from your sire the recessive from your dam as that is the only gene she could contribute to you pups.

Now if your parents were both just carriers and did not have rear dew claws, then I'd expect to see only one in four of you guys have the characteristic. All recessives are passed on like that, even if it is something bad like some forms of epilepsy, black hair folicle dysplasia or cherry eye or so many other things. Not all these problems are caused by genetics for you can get cherry eye or epilepsy from injury. Always best to be cautious before we call what a dog has as being genetic. There are other causes. Genetic is kind of the pop buzz word of the 90's used to explain all kinds of afflictions and some things like cherry eye, bad bites, open fontanels, etc do have other causes.

Ok, in you go. Kennel up. Good hounds. Sleep tight. I've learned a bunch about sire and dam and also each pup. I'm going to write the symbols down in the records of all of them. They all are 'a^sa, a^sa' at the 'A' locus so we'll skip putting that down and they are all piebald spotted so no need to mark the 's^p, s^p' for each although I will mark the grade (percent of white).

No albinos, extremely rare in dogs, so we'll also skip the 'C' locus. The 'C' being the locus that determines full or bright color expression rather than chinchilla, 'c^ch' and a couple call kinds of albino. (Some researchers and authors, Robinson for one, see little if any evidence that 'c^ch' actually exists in canines and variations of shade attributed to 'c^ch' are better explained by the plus and minus modifiers. There does not appear to be a sharp cutoff that allows us to actually identify in the phenotype of canines the dogs that are double recessive 'c^ch' and those that are dominant 'C'.) Actually there is only one albino recessive the other was improperly named long ago as a blue eyed albino recessive, 'c^b'. True albinos will not have any iris color in the eyes. The iris is clear and the red or pink is actually the blood vessels inside the eye. Virtually all beagles are dominant for 'C' as near as I can tell. I have seen pictures of a couple that may be exceptions but they are extremely rare. They might have had the double recessive 'c^b'. Doubtful, but not impossible.

No tail kinks to note, all bites are good. So no notes there. I use 'Dc' for rear dew claws, the recessive pair meaning they have the condition for those two characteristics. I also make note if they have hair whorls in the coat on shoulder or flanks and how many. A recessive 'wo' causes that also. The question mark indicates unknown and we may learn what the second gene is when we see the next generation bred from these pups. Here is the file for the parents and the rainbow litter.
Sire : Bb Dd Ee T? Dcdc Wo? Grade 3
Dam : Bb Dd Ee T? dcdc Wo? Grade 5
Blk mal : B? D? E? T? Dcdc Wo? Grade 4
Chco fem : bb D? E? T? dcdc Wo? Grade 4
Blu fem : B? dd E? T? dcdc Wo? Grade 5
Lilac mal : bb dd E? T? Dcdc Wo? Grade 3
BluTk mal : B? dd E? T? Dcdc Wo? Grade 7
TnWh fem : B? D? ee T? dcdc Wo? Grade 6

Well enough for now. Time to get some sleep. If you want more info about breeding plans, systems of mating, genetics, coffecient of inbreeding, etc. I'd try to find a book called Genetics for Dog Breeders by Roy Robinson, Second Edition, Copywrite 1990. Probably get a copy at your library or by using the interlibrary loan system. It is well worth a serious breeders time. Could have skipped some false starts and mistakes if we'd had the info in that book back in the 50's and 60's. I sure hope it is a rainbow spring in your backyard next year. If that rainbow shows up, you'll know how it got there.
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Oleg V
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