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Red German Shepherds?

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Daylene Alford
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Red German Shepherds?

There seems to be something odd going on how white in Shepherds in inherited. If it wa straight recessive then you would expect it t affect red in other base coats. This may be happening in dogs that are "silver and black "silver sable" which are common in Shiloh Shepherds where white and regular dark co are bred together. However, if this is the cas there should also be solid red or yellow shep who are ee without the dilution that makes t white and this does NOT seem to be happen I'm not sure however if they are really not th if I haven't found them.

Thanks, Daylene

Daylene Alford
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There is something going on,

There is something going on, as you are correct in that ee in the breed should produce gold, cream and white. I have seen cream, but never gold. This is simlar in Tibetan Spaniels by the way, where IThere is something going on, as you are correct in that ee in the breed should produce gold, cream and white. I have seen cream, but never gold. This is simlar in Tibetan Spaniels by the way, where I have yet to see a gold/red ee Tibbie. It is possible that the 'e' allele in these breeds has been mostly bred out, and in the remaining dogs which have retained it, it might be closely linked with a c'e' (or c'ch') on the chromosome, and then customarily inherited together. Both breeds went for a great length of time where the all pups displaying the ee combination were culled from the breeding programs.

SOB

Daylene Alford
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I was wondering if perhaps

I was wondering if perhaps the dilution was linke to the recessive at extension (e). If this were the case, however, then we shouldn't be seeing the "black and silver" and "silver sable" that seems to occur when White Shepherds and "full colored" are bred together. It is possible that the "black an silver" and "silver sable" aren't related to the dilution but that seems a bit of a stretch to me. D you disagree?

Daylene Alford
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"Silver" is a reflection of

"Silver" is a reflection of diluted phaomelanin pigment. The genes that cause the dilution are not exactly identified, but c'ch' and c'e' are the ones commonly known . . . there could be others. Keep in mind that the 'black and silver' and 'silver sable' Shepards STILL have one 'e' allele, sitting recessively beside their E or E'm' allele on the extension locus. If that 'e' allele has accompanied with it a c'ch', then the phaomelanin in that coat will be lighter in shade (as C is only imcompletely dominant to c'e' or c'ch'). When I look at the phaomelanin color in most Shepards I see many that I believe are C/c'ch' or even c'ch'/c'ch', so many of those 'silver sable' or 'black and silver' pups from 'full colored' mixed with whites could easily be c'ch'/c'e'. Does that make sense? I'm thinking this through as we post.

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Daylene Alford
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Now let me see if I can

Now let me see if I can remember what I had said in response.

Yes, I understand you perfectly. Have you read this paper? http://jhered.oxfordjournals.org/content... It seems that the C locus is not responsible for the cream coat color.

I don't think its a incomplete dominant. It seems to me that it affects the red portions on a black based dog to the same extent that it does the red coat of an ee dog. Can you show me an example?

I'm beginning to think that it may be a simple dominant linked to e. That would explain the "black and silver" and "silver sable" pups that result from a white x full color breeding, how full color pups can happen from a ee white x black and silver pairing, and the lack of a full color red. If it was a incomplete dominant these full color pups should not be possible. http://www.siriusdog.com/riddle-of-white... The genetics on this page are out of date but the two example pairings are interesting.

spanielorbust
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I just thought I would post a

I just thought I would post a photo of a Shepherd that looks to show enough red in his phaeomelanin coloring to be considered C/C.

IMAGE(http://www.royalair.org/falcostand%20copy.png)

Most Shepherds I see have much lighter phaeomelanin, indicating to me that in most 'full color' Shepherds there is already an amount of dilution going on (see a lot of dogs that look to be c'ch'/c'ch').

SOB

spanielorbust
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I'm aware that the

I'm aware that the 'chinchilla' or proposed 'C' locus has been proven to NOT be what is at play in dogs, but as we've been given no new name for how to talk about this, I still refer to the locus as Little did.

I also love this photo as it shows the full range of phaeomelanin that can be displayed by e/e dogs.

IMAGE(http://retrieverman.files.wordpress.com/2011/01/golden-color-range1.jpg)

Little's explanation of a recessive inheritance pattern with incomplete dominance of C, c'e', and c'ch' works well for many breeds. Where it is eventually identified, and which letters to be used can be figured at another time.

From the paper.

"In all the families studied, cream coat color fit an autosomal recessive inheritance pattern among the dogs with e/e MC1R genotype. This e/e genotype alone does not cause cream in all dogs of all breeds, and so we assumed that some other gene, which varied in some breeds, must interact to cause cream instead of yellow or red."

I agree that whatever the name, it does affect the red portions on a black based dog just as it would affect a dog that was ee in genotype.

In my mind the silver sable and black and silver pups from a full color x white Shepherd pairing are most easily explained with the idea of recessive inheritance and incomplete dominance, and linkage of c'e' or even rarely c'ch' with the e from the white parent.

IMAGE(http://www.siriusdog.com/images/articles/512/2-Shiloh-litters---color-st.jpg)

In this image above, looking at the Gem x Quest litter, neither Gem or Quest represent full coloring on the C locus. They are C/c'e' or C/'ch' or c'ch'/c'ch' or it is possible Gem is c'ch'/c'e' - as his phaomelanin is so light. As they have a white pup I would suggest Gem IS c'ch'/c'e' and Quest IS C/c'e', but with incomplete dominance those are always guesses. The color of the pups in the litter make complete sense, with only one white pup produced because of the recessive inheritance pattern of white, requiring both parents to carry e.

Elf x Scout also makes sense, as Scout very likely has no e allele, and therefore would not be able to throw white pups. Elf has a lot of coloring for a white Shepherd - maybe c'ch/c'ch - BUT she is also whitened from the underside up. Urajiro markings? Are Grizzle or Domino at play in Shepherds that you know of?

I'd love to see a better photo of Scout, as the color of his phaeomelanin is not clear.

(About Grizzle/Domino)- http://jhered.oxfordjournals.org/content...

SOB

Daylene Alford
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Let me see if my

Let me see if my understanding is correct. All the alleles are recessive to C and may be incompletely dominant or co-dominant with each other?

I agree that whatever the name, it does affect the red portions on a black based dog just as it would affect a dog that was ee in genotype.

This wouldn't fit for a recessive. If all the dilution alleles are recessive to C and linked to e then in theory should be no expression on a Ee dog.
It doesn't fit for an incomplete dominant either as the red portions of the dog should be less diluted in a Ee dog then in an ee dog and there are dogs who are Ee and the red areas are very "white".

If, as we are theorizing, the mutation is linked to e then how likely is it that there really is more than one allele?

In horses you often see as much variation in palominos (caused by one allele Ccr). These two horses are examples of that variation. The first is not common but not rare either the second is the only example I've ever seen that dark (probably with a "sooty" modifier) but they are both palomino.

IMAGE(http://equine.colorgenetics.info/equine_gallery/d/227-4/Thunder.jpg?g2_GALLERYSID=0dacc6874e3888abd7b781d4b537b06b)

IMAGE(http://equine.colorgenetics.info/equine_gallery/d/917-3/Bodi5.jpg?g2_GALLERYSID=0dacc6874e3888abd7b781d4b537b06b)

Grizzle and Domino are supposed to only occur in Saluki and Afghan Hounds and must be atat or perhaps ata (a isn't present in Saluki or Afghans) most likely what we're seeing is a Ee or EE vs EM.

Please don't hot link images from other sites. (unless of course you actually own the site or the photos.)

spanielorbust
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Lots of confusion here, so

Lots of confusion here, so we'll have to start clarifying that we are speaking about the same things. :)I have a feeling that when I am speaking to dominance hierarchy at a locus, you are taking that to mean dominance of one locus over another.

Let me see if my understanding is correct. All the alleles are recessive to C and may be incompletely dominant or co-dominant with each other?

Yes. From the study you linked earlier in the thread.

The second gene in these breeds must have multiple alleles, only one of which causes phaeomelanin pigment to be diluted or pale.

Please keep in mind that the first 'gene' they are speaking to is the e/e allele pair at the extension locus. It is the most recessive pairing of alleles there.

The second gene is the one we are calling the C gene. We do know it is not the same as the one in other animals studied. My understanding is that the letter 'I' might be the candidate name for this gene, but that hasn't been decided yet, so C is continued with in conversations.

There are two alleles at the C locus of all dogs whether or not an 'e' is there.

All the alleles at the C locus are recessive to the semi-dominant 'C' allele there. Those alleles, as Little postulated them, would be C, c'ch', c'e', and when I google I find there is now a fourth one postulated called c'p' (for platinum).

This wouldn't fit for a recessive. If all the dilution alleles are recessive to C and linked to e then in theory should be no expression on a Ee dog

Could you please explain your thinking here. I don't get why you would think this way? How would E/e affect expression of the alleles on the C locus? The linkage of a c'ch' or c'e' to the 'e' allele that is in this breed would mean that E/e dogs would also be ?/c'e'. Incomplete dominance at C would then indicated the the phaeomelanin on these E/e and ?/c'e' dogs would be light.

The only dilution alleles recessive to the dominant C allele, that have been mentioned, are the c'ch', c'e', and now I've brought in the c'p'. I don't quite understand what you mean here by ALL the dilution alleles? There are many loci with alleles that dilute/lighten pigment. Their heirarchy of dominance within their allele set will be independent of each other.

Looking JUST at the C locus, and the hierarchy of alleles there, a dog that is C/C would be darker than one that is C/c'ch'. C/c'e' dogs would be even lighter. c'ch'/c'ch' might look the same as c/c'e' which is why it is hard to say, in the mid range shades, what pair of alleles might be present. The darkest dogs would be C/C at that locus. The lightest combination would be c'p'/c'p'.

Other alleles at other loci, like the Extension locus, have their own hierarchy which is separate from/not affected by the alleles at the C locus.

Keep in mind that if you look at other breeds, an E/e dog can also be C/C. Cavalier Spaniels are a good example as the black/tans are, in fact, E/E or E/e and C/C (C/C is set in the breed). . . but in Shepherds it appears that the dogs with 'e' don't have C. That is an indication that e and a lighter c allele are linked in Shepherds. Whatever founders brought 'e' into the breed had light phaeomelanin. I understand a White Shepherd was in the foundation.

It doesn't fit for an incomplete dominant either as the red portions of the dog should be less diluted in a Ee dog then in an ee dog and there are dogs who are Ee and the red areas are very "white".

I don't understand this conclusion either. Dogs that are E/E or E/e and c'ch'/c'ch' will have light phaeomelanin. If they are E/e and c'e'/c'e' their phaeomelanin might even look white. Dogs that are E/E or E/e and C/C will have darker phaeomelanin. Why should the red portions be 'less diluted'?

Sharing eumalanin and phaeomelanin pigment in a coat does have some effect, though. If you study dogs with both eumelanin and phaeomelanin in a pattern on their coat, you will find that often phaomelanin closest to the black is darker in shade, and then fades out as it moves further away from the eumelanin influence.

Keep in mind that if you look at other breeds, an E/e dog can also be C/C. Cavalier Spaniels are a good example as the black/tans are, in fact, E/E or E/e and C/C (C/C is set in the breed). . . but in Shepherds it appears that the dogs with 'e' don't have C. That is an indication that e and a lighter c allele are linked in Shepherds. Whatever founders brought 'e' into the breed had light phaeomelanin.

Grizzle and Domino are supposed to only occur in Saluki and Afghan Hounds and must be atat or perhaps ata (a isn't present in Saluki or Afghans) most likely what we're seeing is a Ee or EE vs EM.

I believe in Aug 2010 an E'G' allele was named, putting grizzle/domino at the E locus. It has only been found/studied in Salukis and Afghans. That doesn't mean it can't have migrated to other breeds at one point, which is why I was asking. Can't find if we aren't looking.:)

Urajiro pattern would be more my guess anyway, for the underside lightening on Elf. I've seen Urajiro on many, many breeds.

Please don't hot link images from other sites. (unless of course you actually own the site or the photos.)

.

I thought images not copyrighted were fine. How about linking images found on Google images. Is that allowed? (I don't know what 'hot link' means).

SOB

Daylene Alford
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I'll answer the last first

I'll answer the last first because it is easy lol.

Images that are not copyrighted are fine but unless an image has been openly add to the public domain it is, under US copyright law, considered copyrighted by default by whomever took the photo or made the image. Most images on google are copyrighted and google always has a link back to the original site. Hotlinking is using the [img] tags. Feel free to hotlink images that I have loaded to the galleries on this site and also wikipedia wikimedia or other sites that are openly public domain because when uploading a photo to those sites the owner agrees it put it in the public domain. Otherwise, please just add a link back to the site your referencing.

I think what may be throwing us off is that I have been looking at Shepherds specifically, trying to understand why there are no full red pups. I'm making alot of assumptions that would all get thrown out the window if a full red pup could be located.

I'm assuming that there is a dilution linked to e (we're calling it C locus for lack of a better term). I'm assuming the linkage is very tight with no known crossover rate ie no full color red pups). If this is the case all Ee dogs would also have a C allele. If all the C dilution alleles are recessive to C then all the dogs from a full color x white mating should be full color. If there was another allele (maybe cch) that was linked to E in some dogs then I guess that would explain some things. Then you could have a EE cc (I'm not accounting for the individual alleles) that could be black and silver. This could be disproved, however, if a black and silver x white ever produced a full color pup. I wonder that the chances of two different C alleles being linked to two different extension alleles actually is? :rofl

I believe in Aug 2010 an E'G' allele was named, putting grizzle/domino at the E locus. It has only been found/studied in Salukis and Afghans. That doesn't mean it can't have migrated to other breeds at one point, which is why I was asking. Can't find if we aren't looking.

What I meant was a dog must be EG? atat to express grizzle. And yes I agree with you on looking there are some huskies like this one IMAGE(http://upload.wikimedia.org/wikipedia/commons/thumb/1/11/Syberian_brazowoczerwony_pl.jpg/656px-Syberian_brazowoczerwony_pl.jpg) that I would love to see tested. If fact if you ever come across a dog that has been tested positive for EG I'd love to know.

What is the Urajiro pattern? I'm not at all familiar with that one.

spanielorbust
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Thanks for the photo

Thanks for the photo explanation. I am Canadian but our copyright is probably similar.

Easiest to hardest :)

The husky shows above shows a domino pattern. There have been huskies genotyped, and apparently this pattern is from a different genes influence than the one influencing Afghans and Salukis. This is about Domino in Huskies (nice photo explanations). - http://nicholescritters.homestead.com/ma...

A Urajiro explanation and phots can be found here. - http://dogs.lokatt.se/urajiro.htm

I'm assuming that there is a dilution linked to e (we're calling it C locus for lack of a better term). I'm assuming the linkage is very tight with no known crossover rate ie no full color red pups). If this is the case all Ee dogs would also have a C allele. If all the C dilution alleles are recessive to C then all the dogs from a full color x white mating should be full color.

No, not so. If C is incompletely dominant at its locus then it allows the other allele that is paired with it to show some effect.

E/e dogs could then very well be C/c'e' at the C locus (or c'ch'/c'e' etc), and therefore have lighter phaeomelanin color.

I have permission to post these photos.

This girl is E'm'/e (genotyped by DNA). She has a Cavalier dam and a Tibetan Spaniel sire. Cavaliers are set at C, and e/e Tibetan Spaniels are always almost white, so probably c'e'/c'e'.

She, therefore, has a dominant C at her C locus, paired, probably with a c'e'. (She is C/c'e')

Look how light her phaeomelanin is. (She also has a white/cream spotted sister that is e/e).

IMAGE(http://i1106.photobucket.com/albums/h363/japers-18/Missieonboatforupload.jpg)

If there was another allele (maybe cch) that was linked to E in some dogs then I guess that would explain some things.

We know that E and c'ch' are found together in some dogs. I would suggest that is what is found in most 'full color' German Shepherds.

Then you could have a EE cc (I'm not accounting for the individual alleles) that could be black and silver.

I don't get this. How can you not account for individual alleles in any type of genotypic explanation? What do you mean by EE cc then?

Black and Silver Shepherds are very probably E/?(E or e), and c'ch'/c'e'or c'e'/c'e'.

Every dog, whether there is linkage involved or not, will have two alleles at E and two alleles at C. Those alleles have to be taken into consideration if trying to figure out the colors of phaeomelanin or eumelanin that will be displayed.

This could be disproved, however, if a black and silver x white ever produced a full color pup.

Please explain what is being proved/disproved. I can't understand.

I wonder that the chances of two different C alleles being linked to two different extension alleles actually is?

It might be that you and I understand linkage quite differently. I am of the idea that in many different dogs E can be found with a different C allele. Linked or not would be hard to determine.

There will be dogs that are E/E and c'e'/c'e', just as there will be dogs that are E/E and C/c'ch'. E is not always found with the same C allele, not even within a single breed. e is also not always found with the same C allele. Lots of examples of this all through dogdom

We KNOW ruby Cavaliers are e/e and show very vibrant phaeomelanin, (as do their E/E an E/e counterparts - the black/tans) - so C/C.

We KNOW Maltese are e/e and show very white phaeomelanin - so c'e'/c'e' (maybe c'p')

SOB

Daylene Alford
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Thanks for posting those

Thanks for posting those links.

No, not so. If C is incompletely dominant at its locus then it allows the other allele that is paired with it to show some effect.

I thought we had established that all the other alleles at C were recessive to C? If some are incompletly dominant to C then I get to think things through again :rofl

We know that E and c'ch' are found together in some dogs. I would suggest that is what is found in most 'full color' German Shepherds.

Ok I'm starting to understand.

don't get this. How can you not account for individual alleles in any type of genotypic explanation? What do you mean by EE cc then? Black and Silver Shepherds are very probably E/?(E or e), and c'ch'/c'e'or c'e'/c'e'.

This is what I meant. That a black and silver would have to have two of the recessive alleles at C.

Please explain what is being proved/disproved. I can't understand.

If we assume black and silver is E? and cece or E? and cchce and a white is ee cece or ee cchce then they could never have a full color pup because neither parent has a C allele to pass.

It might be that you and I understand linkage quite differently. I am of the idea that in many different dogs E can be linked to a different C allele.

By linked I mean that the linked genes travel together. If you have hypothetical dog that is Ee Cce and the e is linked to ce then the ce will only be inherited along with e. This dogs offspring will never get E and ce together from this parent (unless a crossover occurs). In horses we get this with extension and kit. With this case in horses, there is thought to be a crossover rate of between 3-7%.

I think I'm going to give this a rest for a while. My brain is hurting. Thanks for being so patient with me.

spanielorbust
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Quote:I thought we had

I thought we had established that all the other alleles at C were recessive to C? If some are incompletly dominant to C then I get to think things through again

All the other C locus alleles are recessive to C there. C has incomplete dominance to c'ch' and c'e' (or extra alleles if there are any). c'ch' has incomplete dominance to c'e' (or extra alleles if there are any).

Being recessive does not mean that they cannot show effect. If the allele they are paired with has incomplete dominance (as the C allele does on that locus) then some effect of the less dominant (or recessive) allele will still show.

This is what I meant. That a black and silver would have to have two of the recessive alleles at C

Yes, we agree there. That also indicates the the E in Shepherds can be found with both C and c'ch', as there are Black and Red Shepherds, and also the usual 'full color' Black and Tan, which I believe are E/? and c'ch'/c'ch'.

If we assume black and silver is E? and cece or E? and cchce and a white is ee cece or ee cchce then they could never have a full color pup because neither parent has a C allele to pass

But they will have pups that are E/e and c'ch'/c'ch', which is what I believe the majority of 'full color' Shepherds are. They can also have E/e and c'ch'/c'e', which will have some color in the phaeomelanin. In Shepherds the 'full color' name is a misnomer . . . most have light colored phaeomelanin, much lighter than a Ruby Cavalier, for instance, which indicates phaeomelanin is commonly diluted in the majority of the population. When I google German Shepherd 'full color' images, most of what I see have diluted phaeomelanin. My reference point, of course, are Ruby Cavaliers and Red Cockers, as those breeds are where I got into canine genetics from.

By linked I mean that the linked genes travel together. If you have hypothetical dog that is Ee Cce and the e is linked to ce then the ce will only be inherited along with e. This dogs offspring will never get E and ce together from this parent

Yup, that is what I mean too. With your example, though, the e is linked with the c'e' and ONE parent will give that. The other E parent will give their C allele as well . . . let's say it is c'ch'. Then a pup will have E/e and c'ch'/c'e'. With c'ch' showing incomplete dominance and allowing some affect from the c'e', the E/e pup will have extremely light phaeomelanin coloring (perhaps as light as silver).

Will be glad to continue at another time.:)

SOB