Elven Hair and Eye Color

[MithLuin]

Okay, so....this is a ridiculously pointless thread, but I just kinda had to.

Tolkien once answered a (hypothetical) technical science question about his invented world by saying that you would just have to go there and study the biology for yourself.

Clearly, that's not really an option.

So, any speculation about the genetics of elvish inheritance patterns is simply that - speculation. However, he did give us examples of elves intermarrying with humans and producing viable fertile offspring, so there are enough similarities there to draw some conclusions. Like, elves have the same number of chromosomes (23 pairs) as humans, and that *some* of the genetics must work the same way.

Not all - obviously there are some key differences, like seemingly perpetual youth, near-immortality, prevalence of telepathic abilities, etc.

But the inheritance patterns for hair and eye color probably work the same way ours do. Only problem is that that turns out to be a rather involved issue in itself - we know what genes control eye color, and what pigments are responsible, and yet we're still not exactly sure how it works. Oh, and at least 16 genes are involved.

So...we usually tend to oversimplify it in human families, too, when we try to explain the inheritance pattern.

There are a few things to note about elvish traits that differ from human traits. One, grey eyes seem to be exceedingly common in the family trees we know the most about. And so it is safe to assume that the frequency of whatever allele is responsible for that is much higher in elvish populations than in human populations, though it's still likely recessive, as it is for humans. (It's a variation on blue eyes. Sorta. It's the difference between a clear blue sky and a grey sky promising rain.)

The other elvish trait is being born with silver hair. We typically see white and silvery grey hair in the elderly, but elves seem to have this as a natural hair color that is less grey and more like a variation of blond.


And so....if we oversimplify/fudge human genetics, we can maybe get at what this looks like.

Eye color first - remember, this is NOT RIGHT, even for humans.

First, we are ignoring albinism as a separate issue, though of course that also affects your eye color. Likewise not attempting to explain anything complicated, like dichromism or the darker ring around the iris some people have, etc. Just some basic eye color - brown, green, blue and hazel

Let's use three genes to explain what is going on.
First, you either have brown eyes, or you don't. The allele for brown is dominant.
B = brown, b= not brown
The way simple dominance works, if you have BB, you will have the brown eyes, and if you have bb, you will not have brown eyes...but if you have Bb, it's just like having BB and your eyes will be exactly as dark brown as if you had two dominant alleles. You are a carrier for not-brown, though, so your kids may have other eye colors.

Next, we worry about how dark the eye is. We know there are variations in shades of eye color. You could, for instance, have dark blue or light blue eyes. Let's...over-simplify this, and look at a single incomplete dominant gene to see what that looks like.
D = dark, d = dilute
If you have DD, you will have the dark eye color. If you have dd, you will have a much lighter eye color ('washed out'). And...if you have Dd, you will have something in between - not dark, not pale, just regular old eye color. [Is this really what is happening? No, not really.]

Third, we have to account for the difference between green and blue eyes. Green eyes are dominant, but *only* appear when the brown coloration is absent. So, this could be seen as green being dominant to blue, but recessive when compared to brown. [But they are separate genes, so not really.]
G = green, g = blue

So, what does this look like?

Eye Color
Genotype : Phenotype
BBDDGG : Dark brown
BBDdGG : Light brown
BBddGG : Hazel
BBDDgg : Dark brown
BBDdgg : Light brown
BBddgg : Amber
bbDDGG : Green
bbDdGG : Green
bbddGG : Green
bbDDgg : Dark blue
bbDdgg : Light blue
bbddgg : Grey

Note that no combinations with Bb or Gg are shown, because as simple dominant traits, they would be the same as BB or GG, respectively. Also, since the dilution gene is set up to be affecting the brown eye color gene, it should have no effect on the shade of green of the eye.

So, all the grey-eyed elves would (perforce) be bbddgg, and thus if they marry other grey-eyed elves, they'll have grey-eyed kids, too. But if they marry elves with other eye colors, the grey is likely to get lost, as that is a recessive trait.

 

[MithLuin]

That was really just the warm-up to see if anyone wants to talk about elvish hair color. Because of course we do!

Similar rules - oversimplification makes life easy, and I'm not offering to do a Punnett Square any larger than a tri-hybrid cross.

And so....only three genes to play with again. But the rules are a little trickier because we have to incorporate silver hair.

First - is the elf dark-haired or blond-haired?
B dark (brown) hair, dominant
b light (blond) hair, recessive
We will treat this as a simple dominant trait.

Next, we account for the silver vs black hair pigment. This one is trickier. We will treat this as another 'dilution' gene.
D dark (black) hair, dominant
d light (silver) hair, recessive
DD will give you the darker version of whatever color it is, and dd will give you the lightest version, with Dd being somewhere in the middle. But unlike for eye color, this will have an effect on *all* colors, to give us the most shades to work with.

Finally, just like in humans, the gene for red hair is separate, and recessive, and only shows up if dark (brown/black) hair isn't masking it. Unlike in humans, we aren't necessarily linking it to the same typical red hair traits (freckles, pale skin, profuse bleeding, fiery tempers....oh, wait).
R non-red hair, dominant
r red hair, recessive

Genotype : Phenotype
BBDDRR : Black
BBDdRR : Dark Brown
BBddRR : Brown
BBDDrr : Black
BBDdrr : Dark Brown
BBddrr : Reddish-brown
bbDDRR : Blond
bbDdRR : Silver-blond
bbddRR : Silver
bbDDrr : Dark Red
bbDdrr : Copper Red
bbddrr : Strawberry Blond (pink hair was requested, so we get this 😉 )

Keep in mind that BB is the same as Bb, but does make a difference in what your offspring can look like. (Same for Rr being equally non-red compared to RR).

And now, some 'known' genotypes, based on what we know of their family tree and their phenotypes.
Name : Genotype : Phenotype
Finwë : BxDDRx : Black
Míriel : bbddRx : Silver
Feanor : BbDdRr : Dark Brown
Nerdanel : Bbddrr : Reddish-brown
Mahtan: bbDdrr : Copper Red
Galadriel: bbDdRx : Silver-blond

'x' means that the genotype is unknown, because while the elf clearly has the dominant allele, we do not know for sure if the recessive allele is present or if the elf is homozygous for the dominant trait. Note that we know that either Finwë or Míriel (or possibly both) are carriers of red hair. But we do not know for sure which one it would be, so we have to stick with the 'x' there. In Fëanor's case, we *know* he is a carrier, so we can safely mark him as Rr.

Fëanor and Nerdanel can have children with the following hair colors (based on this):
BBDdRr: Dark Brown
BbDdRr: Dark Brown
BBddRr: Brown
BbddRr: Brown
BBddrr: Reddish-brown
Bbddrr: Reddish-brown
bbDdRr: Silver-blond
bbddRr: Silver
bbDdrr: Copper Red
bbddrr: Strawberry Blond

So, this method does not account for the dark red hair on one of the twins, but does give a variety of options for Celegorm: silver, silver-blond, or brown. It is also cheating, as the silver and blond should be co-dominant in Galadriel's hair, with her having a mix of silver strands and blond strands. So, not perfect, but it makes sense as a starting point to me. If people would like to work with other methods, I'd be happy to adapt. Just...nothing too convoluted. If we wind up trying to account for five different genes controlling elvish hair color, I'm out!

 

[MithLuin]

Alternative plan to account for silver hair.

Everything above is the same, EXCEPT that silver is an alternate allele of the first brown/blond gene; and both silver and blond are recessive to dark hair, but co-dominant with one another. So, for that first pair, an elf could have:
BB = black
Bb = black
Bs = black
bb = blond
ss = silver
bs = silver-blond

And yes, this is mostly bs 😛 Because for the silver-blond trait to be rare, it would still have to only show up under a particular dilution (otherwise, every time a blond elf and a silver-haired elf married, you'd have Galadriel's unique hair). Let's go with the idea that you can't dilute silver, just blond, so to get true silver-blond, you need DD for the blond to show up; otherwise, it gets washed out/overwhelmed by the silver?

So...
BBDDRR = black
BbDDRR = black
BsDDRR = black
bbDDRR = golden blond
ssDDRR = silver
bsDDRR = silver-blond (Galadriel)

BBDdRR = dark brown
BbDdRR = dark brown
BsDdRR = dark brown
bbDdRR = blond
ssDdRR = silver
bsDdRR = silver

BBddRR = brown
BbddRR = brown
BsddRR = brown
bbddRR = pale blond
ssddRR = silver
bsddRR = silver

This might allow for a greater variety of red-hair types, if we need that, too. I haven't really thought about how the silver allele and the red allele would interact, just that the silver does not dilute, so it would behave differently from the blond. So...what color would ssrr give?

BBDDrr = black
BbDDrr = black
BsDDrr = black
bbDDrr = dark red
ssDDrr = dark red?
bsDDrr = dark red

BBDdrr = dark brown
BbDdrr = dark brown
BsDdrr = dark brown
bbDdrr = copper red
ssDdrr = dark red?
bsDdrr = copper red

BBddrr = reddish-brown (auburn)
Bbddrr = reddish-brown (auburn)
Bsddrr = reddish-brown (auburn)
bbddrr = strawberry blond
ssddrr = dark red?
bsddrr = red-orange


So, to redo some of our Noldor:

Name : Genotype : Phenotype
Finwë : BxDDRx : Black
Míriel : s***Rx : Silver
Feanor : BsD*Rr : Dark Brown
Nerdanel : Bbddrr : Reddish-brown
Mahtan: bbDdrr : Copper Red
Galadriel: sbDDRx : Silver-blond

Giving Fëanor and Nerdanel potential children with:
BBDdRr - Dark brown
BBDdrr - Dark brown
BBddRr - Brown
BBddrr - Reddish-brown
BsDdRr - Dark brown
BsDdrr - Dark brown
BsddRr - Brown
Bsddrr - Reddish-brown
BbDdRr - Dark brown
BbDdrr - Dark brown
BbddRr - Brown
Bbddrr - Reddish-brown
sbDdRr - Silver
sbDdrr - Copper red
sbddRr - Silver
sbddrr - Red-orange

So, dark brown, brown, reddish-brown, silver, copper red, red-orange....but still not dark red (because they can't have ss or DD the way I currently have it set up). I think I would have to play around a bit with what genotype I give Míriel and Nerdanel for that to work. But see how introducing a single new allele opens up the options significantly!

 

[Haerangil]

Very Nice. Rationalized Feanor shouzld have dark brown hair... but according to the text he had "...raven-dark hair"... Tolkien was no biologist and didn#t seem to have cared mucxh about inheritance patterns..

 

[MithLuin]

To be fair, Tolkien seems to have done quite well with keeping inheritance patterns correct in his lineages. For instance, he has red hair skip a generation, but he does not do the same with black hair. In humans, red hair is recessive, so it can do that, but black hair is dominant, so it can not. I have fewer issues with working out the genetics of elvish hair color than I do with working out, say, the genetics of inherited magical ability in Harry Potter. Because the author seemed to be under the impression that any trait that ran in families must be dominant, but then created a bunch of characters with non-magical parents. Also, squibs. It can be worked out, but it's frustrating. Whereas blond hair in hobbit families looks very reasonable compared to blond hair in human families.

The 'dilution' factor I'm using that says Fëanor's hair is dark brown rather than black is one of the over-simplified details. In other words, it makes some things work, but is not a perfect match to reality and is mostly made-up, not biological. So, I would not be overly critical of Tolkien based on this! If Finwë had black hair, it is quite reasonable that Fëanor did, too. I just have to find a way to make it work while accounting for whatever genetic markers Míriel would have had with her (made up) silver hair. The 'dark brown' allows for more variety in Fëanor's kids, but I think I can still get the variety I need and give him black hair. I think the 'silver' in the red is allowing for change over time, but I haven't really accounted for that either.

And so....

Name : Genotype : Phenotype
Finwë : BxDDRx : Black (x can be any possible allele)
Míriel : s*D*Rx : Silver (either sb or ss; but if sb, then Dd)
Fëanor : BsDDRr : Black
Nerdanel : Bsddrr : Reddish-brown
Mahtan: sbDdrr : Copper Red
Galadriel: sbDDRx : Silver-blond

Fëanor and Nerdanel's kids:
BBDdRr - dark brown
BBDdrr - dark brown
BsDdRr - dark brown
BsDdrr - dark brown
ssDdRr - silver
ssDdrr - dark red

Well, close! I don't get Maedhros' copper red this way, but despite the constraints, I still get a variety of dark-haired, light-haired, and red kids out of this cross. The constraints are that I cannot give any of their kids DD - Nerdanel *must* be dd to show the red highlights. So, to get the dark red, I need an ss in the kids, requiring that both Fëanor and Nerdanel be Bs. But then there is no way to get the copper red, because I can't have bs or bb in any of the kids. So, the issue isn't the dark brown vs black (not really, anyway). The issue is how do I mesh the red and silver inheritance patterns in just the right way to make Mahtan and Míriel be grandparents to both of the twins?

 

[Faelivrin]

I had posted to the Feanorians thread but deleted that. Before I read these posts, here's my earlier reply:

Nerdanel has simply brown hair, not red. Red is recessive, because it's capable of skipping a generation. She had to inherit a dominant not-red gene from somebody who isn't Mahtan, = her mother.
Feanor is a carrier for red. If silver is also simple recessive, he's also a carrier for that recessive silver. If Nerdanel also carries silver, then yes, either Celegorm or Maglor could be silver.*
If silver is a simple dominant, then Feanor inherited whatever not-silver allele Miriel had, and can't pass on silver at all. But this is quite unlikely because Galadriel's hair is mostly gold with a sprinkling of silver, not the inverse, and none of her brothers nor Orodreth, nor Finduilas are silver. And Celebrian's name suggests that she's probably silver, which would mean silver skipped a generation much like red does.

And that online genealogy calculator is not even close to correct.

*I don't think we should have two elf characters, both with silver hair, named Celegorm and Celeborn. That's confusing, even if they hate each other.



EDIT: Now responding to newer posts:

I started first with the simple dominant/recessive model and could not account for both Finarfin and Fingolfin having the same black-haired father and blond mother. Turns out that dominant/recessive doesn't describe anything in European hair except redheads.

I've read some descriptions of hair heredity here: http://genetics.thetech.org/ask/ask39 and on some other pages of that website. European* dark/light hair is a really complicated (and poorly understood) interaction of many genes, not one or two. I've been going with a simplified model with only 10 on/off eumelanin genes per person, 5 from each parent. As long as silver is a simple recessive, and Finwe and Turin are allowed to be very dark brown instead of absolutely black-haired, it all works out fine.

Possibly Luthien's shadow-black hair is a simple dominant that overrides all genes inherited from mere Incarnates, until some kid doesn't inherited it at all.

*Blond and red hair in Melanesia are both simple recessives on a totally separate gene, but if Elves have something like that, then Earendil can't be blond.

Here's what I figured:

italics are guesses or deductions
rr = red, +r = red carrier, ++ = no red genes
ss = silver, +s = silver carrier, ++ = no silver genes
? = unknown gene (there are a lot of these)

# = number of on eumelanin genes (or range of possible #s) (these are rarely precise)
10 = really black
9 = brown that's almost black
6-8 = brown (or chestnut or auburn)
5 = light or golden brown (or copper)
0-4 = blond or gold (or ginger or strawberry blond)



Ingwë: +?, +?, 0-4 gold
Indis' mother: ??, ??, 0-9
Indis' father: ??, ??, 0-9
Mahtan: rr, +?, 6-7 auburn
Mahtan’s wife: +?, ??, 0-10 not red
Míriel: +?, ss, 4-10 silver
Finwë: ??, +?, 9 black
Indis: +?, +?, 0-4 gold
Olwë: +?, ss, 0-9 silver
Olwe’s wife: ??, ?s, 0-9
Elmo: ?
Elmo’s wife: ?
Thingol: +?, ss, 5-10 silver
Melian: ??, +?, 10 deep black
Nerdanel: +r, +?, 5-8 brown
Fëanor: +r, +s, 9-10 black
Findis: ??, ??, 4-9
Fingolfin: ??, +?, 5-9 dark
Anairë: ?
Írimë: ??, ??, 4-9
Finarfin: +?, +?, 4 dark gold
Eärwen: +?, ss, 0-4 silver
Galadhon: ??, ?s, 0-10
Galadhon’s wife: ??, ?s, 0-10
Maedhros: rr, +?, 5-7 copper or auburn
Maglor: ??, ??, 4-10
Celegorm: ??, ??, 4-10
Caranthir: ??, +?, 5-10 dark
Curufin: ??, +?, 5-10 dark
Curufin’s wife: ?
Fëanor’s twins: rr, +?, 7-10 dark auburn or chestnut or black (started with enough genes suppressed to give a 5-6 copper or light auburn phenotype)
Fingon: ??, +?, 5-10 dark
Turgon: ??, ??, 0-9
Elenwë: ??, ??, 0-9
Aredhel: ??, +?, 5-10 dark
Eöl: ??, ??, 4-10
Argon: ?
Finrod: +?, +s, 0-4 gold
Angrod +?, +s, 0-4 gold
Edhellos: ??, ??, 0-9
Aegnor: +?, +s, 0-4 gold
Galadriel: +?, +s, 3-4 gold with silver speckles
Celeborn: +?, ss, 0-10 silver
Galathil: ?
Galathil’s wife: ?
Beren's ancestors and sister: not silver
Beren: +?, ++, 5 golden brown
Lúthien: ??, +?, 10 deep black
Celebrimbor: ?
Baragund: ?
Morwen's mother: +?, ++, 0-4 blond
Galdor: ??, ++, 0-9
Hareth: ??, ++, 5-9 dark
Belegund: ?
Rian's mother: +?, ++, 0-4 blond
Maeglin: ??, +?, 9-10 black
Orodreth: +?, +?, 0-4 gold
Oro’s wife: ??, ??, 0-9
Nimloth: ?
Dior: ??, +?, 5-10 dark (or magic black)
Morwen: ??, ++, 5-6 dark (brown)
Húrin: +?, ++, 3-4 blond (gold)
Huor: ??, ++, 0-9
Rian: ??, ++, 5-9 dark
Túrin: ??, ++, 8-9 very dark
Lalaeth: +?, ++, 0-4 blond
Nienor: +?, ++, 0-4 blond
Tuor: +?, ++, 0-4 blond
Idril: +?, +?, 0-4 gold
Eärendil: +?, +?, 3-4 gold
Finduilas: +?, +?, 0-4 gold
Gil-galad: ?
Elwing: ??, ??, 5-10 dark (or magic black)
Eluréd: ?
Elurín: ?
Elros: ??, ??, 0-9
Elrond: ??, +?, 9 very dark (or magic black)
Celebrían: ?
Elrond’s twins: ??, +?, 5-10 dark (or magic black)
Arwen: ??, +?, 5-10 dark (or magic black)
Aragorn: ??, +(+), 5-10 dark
Eldarion: ? probably not silver


Where I seem to be stuck is calculating the probability that a person with X eumelanin genes passes on A of them. It's really complicated and hard. (If a bag contains 10 marbles, X black and Y yellow, and you pull out 5 without putting any back, what's the probability you draw A black and B yellow marbles?) There are 252 possible combinations per "genome" and... that's as far as I've gotten.

 

[Haerangil]

:-O

 

[MithLuin]

Yeah, it gets really complicated - hence my reliance on a single dilution factor to take care of some of that shading, because, blerg, nobody wants to do that Punnett square! But if there are 10 on/off genes that each act as a simple dominant, then the possible gene combinations are MUCH higher for the darker colors. Because someone who has most of those genes turned 'off' is pure recessive for those, and will definitely be passing that on to their kids. Same with a pure dominant. It's if they're heterozygous that they have a 50/50 shot of passing each one on...and it's really hard to determine if someone is heterozygous or not! And then you have to combine the pale-haired parent with the other parent and just...such a mess.

(But as for how you determine the probability without doing any math...the answer is 'construct a Punnett square' - once you nail down the parental alleles. 😉 )

 

[Faelivrin]

because, blerg, nobody wants to do that Punnett square!

Well, I do*, but I can't make a 10-dimensional Punnett square without ... software that calculates things like that. (I want to figure out these probabilities so I can narrow down those broad eumelanin number ranges on my list, or at least say "blond is a 0.4% chance for So-and-so, they probably aren't blond".)

*Until I get bored by this math.

But if there are 10 on/off genes that each act as a simple dominant, then the possible gene combinations are MUCH higher for the darker colors.

The European eumelanin genes aren't dominant/recessive, they're additive/codominant. So it isn't quite that difficult to get blonds. But I started out thinking 0-5 are blond and 6-10 are dark, but that didn't allow me to get Finarfin and Fingolfin as brothers. With 5 = brown and 9 = black-ish, it is possible, and as a bonus Feanor still gets to be black-haired.

 

[MithLuin]

Okay, I'll bite.

It's virtually impossible to identify genotype from phenotype when working with 10 different genes and only a small fragment of a family tree. So this is all massive guess work. I'm also a bit confused whether you mean 10 genes or 5 genes with 10 alleles. I...can do the 5 gene version easier, obviously (though it's nowhere near *easy*), but then might have trouble translating it to your 1-10 scale.

Say there are 5 genes for the pigment. If you get the dominant allele, the gene is 'on', and you get pigment. If you get only recessive alleles, the gene is 'off', and there's no pigment. To see the end result, you add up all the genes that cause pigment, and see how much you get total. (For the curious, human height works like this, too. But instead of 'pigment,' it's 'growth spurts'.)

I will label the 5 genes with the letters V thru Z (we can use Q-U if you want 10 genes, but I'm hoping to avoid that unpleasantness). Each person has two alleles for each one, one for each chromosome. One from mom, one from dad. And so....

VVWWXXYYZZ = all 5 pigment genes firing = really black hair

And all of his/her children are getting black hair, too, because this person is passing on:
VWXYZ

Conversely, an individual with
vvwwxxyyzz = 0 pigment genes turned on, so none of that pigment; red and silver hair colors are visible, or bleach blond (Malfoy blond)

And all of this person's children will be getting
vwxyz ... meaning that all of the pigment will be determined by the other parent.

But those are the extremes. Most people have some, but not all, of these pigment genes turned 'on.' There's no way of telling the difference of which pigment genes someone has, so that part is not predictable in finding the interactions. Also, since these aren't disease genes, it's not a very important question to answer. We just want to know if the inheritance works.

5 = black
4 = dark brown
3 = brown
2 = blond (or dark red, or silver?)
1 = pale blond (or copper red, or silver?)
0 = platinum blond (or strawberry blond, or silver)

Our goal here is to identify the phenotype of Fëanor and Nerdanel, and see if the model will work to predict the correct children outcomes. They are a good starting point for a lot of reasons, but this conversation *did* start on the Fëanorean thread, so.

The text says that Fëanor has 'raven' dark hair. That sounds like a lot of pigment! So, probably he has a dominant allele for each gene. However his mom has silver hair. For the silver to show up, she would have to be missing some of these pigment genes. So, say, she has AT MOST two dominant alleles, and the rest were pure recessive. That leaves these types of possibilities for Fëanor:
VVWWXxYyZz
VvWwXxYYZZ
VvWwXXYyZz
VvWwXxYyZz
etc
[one copy of the dominant allele of each gene, with 2, 1 or 0 genes being homozygous dominant and the rest being heterozygous.]
And yes, I realize that it will be a lot harder to make this work if he has all the dominant alleles, but it's worst for the ones he's homozygous for. So maybe we decide silver is the zero value....

Nerdanel has a red-haired father, a non-red-haired mother, and brown hair herself (with visible red highlights).
So, she has about 3 dominant alleles, but at lease one (and probably two) of her markers is pure recessive. Her father has AT MOST two dominant alleles to pass on; the rest of his contribution has to be recessive.
So, some possibilities for Nerdanel include:
VVWwXxyyzz
VVWWXxyyzz
VvWwXxyyzz
Etc
[Dominant of three genes present. 0, 1 or 2 of these are homozygous dominant. Two genes are homozygous recessive. Up to three genes could be heterozygous dominant.]


So, let's do the EASIEST cross of these two as an example, to see if our system of pigment assignments is working.
Fëanor: VVWWXxYyZz
Nerdanel: VVWWXxyyzz

With these variations, ALL of their children would have VVWW, so we don't need to cross those. That just leaves the remaining three (like I said, tri-hybrid cross is my limit 😛 )

All possible offspring:
XXYyZz = black hair
XxYyZz = black hair
xxYyZz = dark brown hair
XXyyZz = dark brown hair
XxyyZz = dark brown hair
xxyyZz = brown hair
XXYyzz = dark brown hair
XxYyzz = dark brown hair
xxYyzz = brown hair
XXyyzz = brown hair
Xxyyzz = brown hair
xxyyzz = blond/dark red/silver hair

As you can see from this cross, we ran into a problem with the parents being homozygous dominant for two of the pigment genes. They should only have a 1/16 (or is it 1/32?) chance of having a red-haired kid from this, and 3 of their 7 children are red-haired. Too much pigment!

So, let's dial Míriel back to pure recessive and redo this.

We now know the alleles for Fëanor and his parents:
Finwë: V*W*X*Y*Z* (his family with Indis fills in some of these blanks, but let's ignore that for now)
Míriel: vvwwxxyyzz
Fëanor: VvWwXxYyZz

Fëanor is now pure heterozygous, so he can pass along ANY combination of pigment alleles.
Nerdanel still has three (3) dominant alleles expressed, and up to (2) of those could be homozygous. But...we don't want her to be homozygous dominant for two genes; it won't work. So, let's make her homozygous for only (1):
VVWwXxyyzz

And now to calculate their children (*deep sigh*) I'm gonna need more music....

 

[MithLuin]

Fëanor: VvWwXxYyZz

Fëanor can pass on the following combinations of alleles in his gametes (2^5=32 combinations):

VWXYZ
VWXYz
VWXyZ
VWxYZ
VwXYZ
vWXYZ
VWXyz
VWxYz
VwXYz
vWXYz
VWxyZ
VwXyZ
vWXyZ
VwxYZ
vWxYZ
vwXYZ
VWxyz
VwxYz
vwXYz
VwXyz
vWXyz
vWxYz
VwxyZ
vwxYZ
vWxyZ
vwXyZ
vwxyZ
vwxYz
vwXyz
vWxyz
Vwxyz
vwxyz

Nerdanel = VVWwXxyyzz

Nerdanel has CONSIDERABLY fewer possibilities, because she is homozygous for three (3) of the 5 traits. So, the possible combinations of alleles that can be found in her gametes exclude anything with v, Y or Z, leaving only 2^2 = 4 possibilities:

VWXYZ
VWXYz
VWXyZ
VWxYZ
VwXYZ
vWXYZ
VWXyz
VWxYz
VwXYz
vWXYz
VWxyZ
VwXyZ
vWXyZ
VwxYZ
vWxYZ
vwXYZ
VWxyz
VwxYz
vwXYz
VwXyz
vWXyz
vWxYz
VwxyZ
vwxYZ
vWxyZ
vwXyZ
vwxyZ
vwxYz
vwXyz
vWxyz
Vwxyz
vwxyz

So, to cross these two, we'll get a total of 32x4 = 128 possibilities
And that is happening in Excel, or not at all. brb

 

[Faelivrin]

I mean 5 genes, diploid (so 10 gene copies) with possible 2 alleles each. Neither allele is dominant or negative. You just add up the number of M's between 0 and 10.

So a 6 would be MMMMMMmmmm (brown)
And a 3 would be MMMmmmmmmm (blond)

A 6 has 252 possible different combinations of 5 genes they could pass on to each child. I've been trying to calculate how many out of 252 contain 2 M, how many contain 3 M, etc. I should really go to bed though.

I'll have to look more closesly at how you're treating silver, it's very different from the assumption I used.

 

[MithLuin]

Ugh, yuck. That's like taking my 'dilution factor' and ramping it up to...5. Do not like 🙁 [I'm not suggesting you're wrong; I'm suggesting it would be tedious to figure anything out that way!]

Well, I did the cross above anyway, and determined that while you can get copper red out of it, you only have a 1/64 chance of doing so (1/8 chance of getting the darker red). While that isn't an impossible way to get Fëanor and Nerdanel's family, it's not the most likely way, either. So, if I kept them that way I set it up (on/off switches), then you'd have to go with no homozygous dominant as the most likely way of getting the desired outcome.



So, Fëanor would be VvWwXxYyZz and Nerdanel would be VvWwXxyyzz

Their family would be set; next step would be to figure out how blond and silver work using other families. But not necessary if this isn't the underlying assumption.



You can't really do anything with genetics without establishing dominance in the inheritance pattern. I think you will find it easier (not easy, just less-likely-to-pull-your-hair-out) if you treat each gene discretely, rather than trying to combine them from the start. You are positing (2) alleles for each gene, with incomplete dominance. So, MM = 2 units of pigment, Mm = 1 unit of pigment, and mm = no pigment. But times 5, so it's M1, M2, M3, M4, and M5. Which does create a spectrum of 0-10 possible pigmentation outcomes. And the advantage is that it's unlikely that a child would have darker hair than both parents, but not impossible, which fits real life examples well enough. However, if there's no attempt to assign a specific pigmentation to each number, then....such an elaborate system isn't necessary. At the very least, dial it back to 3 genes!

M = 6 = black = MMMMMM
M = 5 = dark brown = MMMMMm
M = 4 = brown = MMMMmm (or MmMmMM, etc)
M = 3 = golden blond (or dark red) = MMMmmm (or MmMmMm)
M = 2 = blond (or copper red) = MMmmmm (or MmMmmm, etc)
M = 1 = pale blond (or strawberry blond) = Mmmmmm
M = 0 = platinum blond (or pink, or silver) = mmmmmm

Or, if that's getting complicated, represent the three genes with three different letters.
So, MMNNOO = black
and mmnnoo = platinum blond
and all the other combinations are based on the number of capital letters present. While it's not impossible to do this for five incomplete dominant genes, I'm going to keep balking at that until someone shows me the need to go there.


Red overrides this pigment as soon as you get into the 'blond' shades, which makes sense, as that's how it works in humans. And I can understand the concern about 'pink', but since the only family where we see red hair is in Mahtan's descendants, and Nerdanel marries the dark-haired Fëanor, this isn't a risk (Well, okay, maybe it was a 1/256 risk or something, but it didn't happen, so we're fine).

But what about silver? Silver isn't likely to be *more* pigment added *on top of* the blond, cancelling it out...is it? So what's the interaction between silver and blond? I consider this part of the point of this exercise, to work out what that relationship is and still make the family trees work. But...obviously, there's no way of testing this out in real life, so it's all guess work. I figure we take the handful of example family trees with silver, and if the proposed solution works for them, good enough. I'm by no means set on keeping the ones I've put forward so far, since I haven't even investigated the families with silver hair in any detail yet. So far, nothing I've said works perfectly; I was trying to do it *without* resorting to 7 different genes to keep track of!

 

[Faelivrin]

LOL yes it is probably more work. Although, so far I haven't found calculations tedious. I'm frustrated because I haven't figured out what function(s) to use to calculate the probabilities.

But I want to read over your model in more detail.

I did assign colors to numbers, like so:

10 MMMMM|MMMMM = deep black
9 MMMMM|MMMMm = “black” (very dark brown)
8 MMMMM|MMMmm = dark brown (with rr = chestnut)
7 MMMMM|MMmmm = brown (with rr = dark auburn)
6 MMMMM|Mmmmm = brown (with rr = light auburn)
5 MMMMM|mmmmm = light brown (with rr = copper)
4 MMMMm|mmmmm = dark/ash blond (with rr = ginger)
3 MMMmm|mmmmm = deep gold (with rr = ginger)
2 MMmmm|mmmmm = blond/gold (with rr = ginger)
1 Mmmmm|mmmmm = light blond (with rr = strawberry blond)
0 mmmmm|mmmmm = platinum blond (with rr = strawberry blond)

I made silver a simple recessive with no impact on non-silver hair color, but I like your idea that it's tied to dilution in Elves. Being a carrier for silver can't be enough to dilute, or all of Galadriel's brothers would have hair like hers, but it must interact with other genes to determine whether the eumelanin is diluted. So Galadriel (definitely a carrier for silver in my model) and the twins (potentially carriers) have that effect, without being silver.

I'm also beginning to suspect that the inheritance patterns make more sense if silver can only drown out 5-6 eumelanins, but can only reduce 7-10 to a lighter brown instead. Miriel and Thingol have to have at least 4-5 (well, Miriel does; Melian could have a simple dominant super-black that overrides everything, though I haven't tried to incorporate that.)

 

[MithLuin]

Hmmm, okay, I can work with that. I think. Maybe 😛

Fëanor would be easy to type:
VVWWXXYYZz = 9 of 10 possible pigment alleles = dark brown/blackish

Finwë would be this also, so make him
VVWWXXYYZz (Obviously, his one recessive allele could be different from his son's - he has one, it doesn't matter which for now).

Leaving Míriel as
V*W*X*Y*** - a lot of unknowns, but she has at most one pure recessive, leaving at least 4 pigment alleles, minimum (as you said).

Using this model, Nerdanel's 'brown with red highlights' hair makes her likely to be 6-7 range on pigments, right? So, some possibilities for her include:
VVWWXXyyzz
VVWWXXYyzz
VvWwXxYyZZ
VvWWXXYyzz
vvWWXXYYzz
etc
Consindering she's got at least one kid at the 5 dominant allele range, and a husband who is passing on at least 4 dominant alleles, there's only so many possibilities for her that allow her to be the mother of Maedhros. Namely,
Fëanor: VWXYz
Nerdanel: vwxyZ or Vwxyz or vWxyz or vwXyz or vwxYz
to give Maedhros: VvWwXxYyZz or VVWwXxYyzz or VvWWXxYyzz or VvWwXXYyzz or VvWwXxYYzz
So, that makes Nerdanel a confirmed 6 (no more alleles possible for her), and she can only be homozygous dominant for ONE of those. Leaving her final options:
VVWwXxYyZz
VvWWXxYyZz
VvWwXXYyZz
VvWwXxYYZz
VvWwXxYyZZ

Since she has a grand total of one grandson, and he's non-descript dark-haired, there's no need to differentiate between these possibilities. Presumably, most of her dominant alleles are from her mother, and most of her recessive are from her father, but we only needed him to have at least one dominant, and he probably had 5 (his nickname, 'Rusco' = fox indicates his hair color, so even though foxes do vary in shade a bit, it's clear that what is intended is the same copper-color Maedhros has).

So, I'll cross Nerdanel (VVWwXxYyZz) with Fëanor (VVWWXXYYZz) to see what the probabilities are for their children, and see if that is close enough.
Fëanor produces two possible alleles: VWXYZ or VWXYz
Nerdanel produces 2^4 =16 possible alleles:
VWXYZ
VWXYz
VWXyZ
VWxYZ
VwXYZ
VWXyz
VWxyZ
VwxYZ
VWxYz
VwXyZ
VwXYz
VWxyz
VwXyz
VwxYz
VwxyZ
Vwxyz

Giving 32 possible combinations:



10 (1) = 1/32 = 3%
9 (5) = 5/32 = 16%
8 (10) = 10/32 = 31%
7 (10) = 10/32 = 31%
6 (5) = 5/32 = 16%
5 (1) = 1/32 = 3%
4 (0)
3 (0)
2 (0)
1 (0)
0 (0)

This distribution says that this model gives us only a 3% chance of Maedhros' hair color if Fëanor has 9 of 10 dominant alleles, and Nerdanel has 6 of 10, but only one homozygous. There's a 16% chance of light auburn, though, which is plenty to account for the twins. Also, there's a 3% chance of getting a pure black 10/10 from this, too, so Caranthir can be as dark as we like (even if he's more likely to be a 9/10), and Celegorm can be no paler than a light brown. So, it does work, though only just. If we gave Nerdanel light brown hair (a 5 instead of a 6), it would work even better. No matter what, they can only have children one shade lighter than Nerdanel.

 

[MithLuin]

Okay, I've been thinking about the purpose of the 0-10 system. And while certainly using real human genetics gets us A+ for accuracy, I struggle to see the value of applying such a detailed selection of color swatches to this project. But...then it occurred to me. We can use this to pick out the wig colors we intend to use, and then give that to costume people so they have something to work with beyond pictures of actors with their regular (probably too short) hair.

And so, I've been doing some 'research' 😉

Welcome to wigisfashion, a great place to get discount lace-front wigs if a) your head isn't too big and b) you don't mind waiting for them to ship from China. 😀
https://www.wigisfashion.com/

So, here's their products as an illustration of the pigment distribution.

"Super" gene - jet black

10 - natural black

9 - black mixed brown

8 - dark brown

(or auburn brown)

7 - brown

(or brownish orange)

6 - mixed brown

(or reddish brown)

5 - golden brown

(or mixed reddish brown)

4 - golden blond

(or red mixed orange)

3 - ash blond

(or light orangish brown)

2 - blond

(or orangish brown)

1 - light blond

0 - white platinum blond

 

[Faelivrin]

Great! Although we'll need a brighter gold color for some Elves, I think. And something special for Galadriel, of course. And we can't forget beards, and a blue beard for Dwalin. 😉

I'd allow Maedhros and Mahtan to be as dark as 7 ("auburn" is the specific color indicated by "russa", according to some linguistics in Vinyar Tengwar) even without the added possibility of the silver gene being a diluter. Which inevitably brings back the blond possibility for Celegorm, though much more speculatively (I'd prefer light brown). I also set Feanor at a 10, because "raven-dark" sounds darker than "black", though I acknowledge that the genetics make 9 more plausible. And, more plausibly, I gave Maeglin 10.

Twins' colors: they started the same but one "grew darker". How dark? Copper to dark auburn, or auburn to chestnut? Or all the way to black, 9-10? They kept calling each other Ambarussa, but I don't know whether that proves Amros stayed perceivably red. What do you think?

Do you think the names Celebrian and Nimloth suggest those women have silver hair, or is it a reference to something else (not necessarily appearance)?

Where did Tolkien specify Celebrimbor's color? I didn't have any data for him.

And please tell me, how did you calculate those %?? I want to do that too.

 

[MithLuin]

I added the graphic of the Punnett Square I used to calculate that. Once you know the possible parental gametes, it's easy. You just combine them, and in this case, count the number of combinations that give you a particular shade out of the total.

Yeah, none of this takes silver into account. It's really just black/blond/brown and red right now.

Russandol means 'copper-top' and Rusca means 'fox'. So, whatever hair color we pick for Mahtan and Maedhros, I'd like for it to be fairly light and coppery.

For additional shades of wigs, feel free to search through this for what we need:
https://www.arda-wigs.com/blogs/news/shop-by-color


I'll try another pairing...I guess Finwë and Indis is the obvious choice.

 

[Faelivrin]

Oh man! I just realized that by naming all the eumelanin genes M or m, I forgot that parental alleles can only pair with the corresponding allele from the other parent. (I'm ignoring recombination and crossovers and other shenanigans.) You're doing it the correct way with VvWwXxYyZz. Yeah, there must be far fewer than 252 possible combos per gamete. It's not picking 5 marbles out of a bag of 10, it's flipping 5 independent coins.

OK, back to the drawing board. Hopefully this time it'll be less impossible to calculate. 🙂 (And looking at your Punnett square, I see it's much simpler with somebody who's mostly homozygous.)

EDIT: Well, that was a lot easier. A 5-dimensional Punnett square only took 5 minutes or so. 🙂 The worst was only out of 32, not 252.


EDIT2: Is the name of that wig site just a coindence? I think there can be some variation in silver hair, what Arda Wigs calls 'pure white', 'snow white', and 'silver'.

For blond Elves I think 'yellow blond' should be an option, and maybe even 'banana'. For Galadriel, 'fairy blond' might have the right variegation, but the yellow strands are much too light.

For "pink" hair (silver + red, if silver doesn't override red), 'custard', 'light peach', and 'Serah pink' are semi-close, but they don't have a proper beige.

For Dwalin, I think 'dusty blue' or even 'cerulean' or 'periwinkle' would be reasonable, but not the brighter blues. I don't picture Dwalin himself having blackish hair, but 'steel blue', 'Marianas', and 'raven' look like reasonable blue shades as well. Maybe Longbeards have a blue hair allele but not a red hair allele, and Firebeards have the inverse. Er, but then if you combined blue with blond you'd get green, so it should not combine with other colors the way red does [in Eruchin].


EDIT 3: It's easier/more plausible to explain Maedhros by dilution/gene deactivation since it happened to Amrod too, instead of trying to get a 5 with such low probability. But I'd also say Nerdanel could be as light as 5, at least in my model 5 still counts as brown.

 

[MithLuin]

I see it's much simpler with somebody who's mostly homozygous.

Yep, the worst would be all heterozygous, which yields the most variety of results. It's called a pentahybrid cross, and I am *NOT* doing one...though I'm sure I could find the results somewhere if need be and skip that messy step. There would be 32x32 boxes for a total of 1024 outcomes. That's a huge NOPE from me!

You will not be surprised to learn that Arda sells wigs to the anime cosplay community. You can often find them in the dealers room at anime conventions. I've never asked them where the name came from, but I would imagine it's from Tolkien. They do get credit for having the full rainbow, though!

WigIsFashion is a better quality product, with more natural hair colors. But Arda wigs fit better heads and ship from Chicago. So.

The two of these are the first two companies to be mentioned if you go online and say 'Where can I buy an elf wig?'

EDIT 3: It's easier/more plausible to explain Maedhros by dilution/gene deactivation since it happened to Amrod too, instead of trying to get a 5 with such low probability. But I'd also say Nerdanel could be as light as 5, at least in my model 5 still counts as brown.

Yes, I think we either have to bump Nerdanel down to a 5, or allow the silver to impact the shade of red in some way. But, technically, Maedhros could be a 5, and the twins could be 6 or 7, and no one has broken probability. Strained credulity in a minor way, perhaps 😉.

This exercise demonstrated to me that Fëanor is definitely a 9 (not a 10), and that Nerdanel is a 5-6, no higher. Do share any other crosses you do, to see if that tightens up this system or sheds light on the silver issue.


So, Indis.

She's obviously blond, but 'how pale?' is an interesting question.

I'm okay starting with giving her (4) alleles for pigment, putting her at golden blond range. The story of her and Finwë deciding to get married involves the light of the Trees in her hair...and both the light and her hair are described as 'golden' in two different places. So...golden blond it is! (For now)

Finwë has the dark hair of the Noldo, so I am giving him (9) alleles.

Finwë's genotype is thus: VVWWXXYYZz, and like Fëanor, he can form two variations of gamete: either VWXYZ or VWXYz.

Since Indis will need to have both dark haired and light haired children, I think we should make her as heterozygous as possible for maximum variation in results.
So Indis' genotype is VvWwXxYyzz
The 16 possible gametes she can have are:
VWXYz
VWXyz
VWxYz
VwXYz
vWXYz
VWxyz
VwxYz
vwXYz
VwXyz
vWxYz
vWXyz
Vwxyz
vWxyz
vwXyz
vwxYz
vwxyz


10 - (0)
9 - (1) blackish (3%)
8 - (5) dark brown (16%)
7 - (10) brown (31%)
6 - (10) brown (31%)
5 - (5) light brown (16%)
4 - (1) ash blond (3%)
3 - (0)
2 - (0)
1 - (0)
0 - (0)

Hmmm, I see that using your scheme, golden blond should be at (3). I thought ash blond looked lighter than golden blond, but I'm no artist...so...I can flip that.

Finwë, at 9, cannot have any children with less than (4) pigment alleles. So, making Indis lighter haired will increase the probability of them having a child at (4), but no matter how light she is, the child will not be any lighter. For the sake of differentiation on this project, we wanted to make Findis blond haired and Irimë dark haired, so we are portraying half of Indis' children as being blond. So, I should definitely bump her down to a (3), I think.

So, Finwë is still VVWWXXYYZz, but now Indis is vvWwXxYyzz (So, I'll remove all of her gametes with 'V' alleles from the cross)



Now, the result is:
10 - (0)
9 - (0)
8 - (1) 6% dark brown
7 - (4) 25% brown
6 - (6) 38% brown
5 - (4) 25% light brown
4 - (1) 6% ash blond
3 - (0)
2 - (0)
1 - (0)
0 - (0)

I'm not sure that doubling the chance of a blond kid is worth losing the darkest hair color (9), but without bringing silver into this, those are our options.