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Daenerys: Very Inbred But Not Very Targaryen
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Vox has a post up, This comprehensive Targaryen family tree explains Game of Thrones’ most complicated dynasty. I don’t really know if the family tree explains much, but it’s interesting. It does highlight two important dynamics which are probably important.

Screenshot 2016-06-14 22.51.18 The Targaryen’s are notoriously inbred. The rationale for this is that their affinity with dragons is a heritable trait, and that is the basis of their power (or it was before all the dragons died, at least until Daenerys brought them back). If they mixed their genes with outsiders presumably they would dilute that ability. Of course that doesn’t take into account selection for the trait, which would diminish any dilution through outbreeding.

As you can see Daenerys’ parents and grandparents were full siblings. Using a pedigree method her Wright’s coefficient of inbreeding would be 0.375. For a point of comparison, the child of parents who were full siblings, but whose own parents were unrelated, would have a coefficient of inbreeding of 0.25. The terminal individual in the genealogy of the Spanish Habsburgs, often cited as a case study of how inbreeding can lead to the extinction of a lineage through sterility and imbecility, had a Wright’s coefficient of inbreeding of 0.254. Something really doesn’t add up in terms of the viability of the offspring of the Mad King and his sister-wife (to get a sense of what multi-generation inbreeding might do, please see the Colt family). But then, it’s fantasy, and genetics is one area where George R. R. Martin’s gritty verisimilitude gives way to flights of fancy.

a-knight-of-the-seven-kingdoms-jpg A second aspect of the generation which Daenerys is a member of is that they are not very Valyrian in terms of their ancestry. More precisely, they are at most ~1/8th Valyrian.

This might surprise some, as the incestuous and closed-off nature of the Targaryen’s plays a huge role in the backdrop of George R. R. Martin’s novels. But incestuous marriage is actually a resurrected custom of the previous two generations before Daenery. Her great-grandfather, Aegon Vth, “Egg” from the novellas, was the result of two generations of outmarriage (so he was outbred). This pattern of outmarriage was not a conscious shift in the mores of the Targaryen royal house, but happenstance. Aegon’s father and grandfather both came to the throne because other senior claimants died. They were not expected to become kings. Aegon’s mother was a Dayne, while his paternal grandmother was a Martell. He himself married Betha Blackwood (a family of First Men background which still worships the old gods). Therefore, Daenery’s two grandparents were both 1/8th Valyrian, her parents were 1/8th Valyrian, and she is 1/8th Valyrian (at least that’s the expected value).

 
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  1. Brett says:

    If the Targaryens (and before them all of the Valyrian nobility doing the “brother-sister” thing) practiced really ruthless infanticide for generations on any offspring with obvious physical or mental defects either at birth or in early childhood, would that weed out some of the bad consequences of the inbreeding over time? It seems like something they would do, especially since they’re also polygamous (or were polygamous) and have no lack of children.

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  2. would that weed out some of the bad consequences of the inbreeding over time?

    possibly. this works with plants (load purging). but it doesn’t seem to work too well in metazoans. one of the issues is if you select really strongly u start to reduce effective population a lot and that introduces drift.

    also, they weren’t sequencing. so they would get a lot of obviously ill infants, but a lot of deleterious traits show up an older ages. for example, several of the fritzl kid’s have some health issues which are treatable with modern medicine

    https://en.wikipedia.org/wiki/Fritzl_case#Elisabeth_Fritzl.27s_confinement

    kerstin’s only showed up in her teen years.

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    • Replies: @Anonymous
    >possibly

    If you haven't, check out Helen Dean King's successful, multi-year work on extreme inbreeding, from early 20th century. Her lines of nearly-homozygous rats are ancestral to approximately half of modern lab rats. The entire journal is accessible on Google Books:

    https://books.google.com/books?id=5WDXAAAAMAAJ


    She started getting very good results by around sixteenth generation. Obviously, doing inbreeding correctly requires progeny maximization and strict arbitration as to who goes to make the next round, for load purging to have its strongest effect. It's more-or-less doable on rats, but of course humans are a whole different story.
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  3. The need for inbreeding is not controverted by the lack of aversion of genetic introgression if you imagine that GRRM is genetics-literate, but not a genetics sperg. He designed the genetics of this plot mechanic back in the 80′s, with classic, high-school genetics literacy. He made the genetics of dragon riding simple and Mendelian, rather than complex and polygenic.

    The dragon gene is X-linked and codominant.

    Single-DrX individuals can more easily tame and ride dragons, and Double-DrX individuals (always women) have, in addition, a facility for dragon egg hatching.

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    • Replies: @Razib Khan
    The dragon gene is X-linked and codominant.


    codominance means a mix of traits
    https://en.wikipedia.org/wiki/Dominance_(genetics)#/media/File:Co-dominance_Rhododendron.jpg
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  4. @AnonymousCoward
    The need for inbreeding is not controverted by the lack of aversion of genetic introgression if you imagine that GRRM is genetics-literate, but not a genetics sperg. He designed the genetics of this plot mechanic back in the 80's, with classic, high-school genetics literacy. He made the genetics of dragon riding simple and Mendelian, rather than complex and polygenic.

    The dragon gene is X-linked and codominant.

    Single-DrX individuals can more easily tame and ride dragons, and Double-DrX individuals (always women) have, in addition, a facility for dragon egg hatching.

    The dragon gene is X-linked and codominant.

    codominance means a mix of traits

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    • Replies: @AnonymousCoward
    If you see trait: dragonrider and trait: not-dragonrider as *not* both being traits, then you'd not use the term co-dominant, that's true. If so, you're right, you'd use the term incomplete dominant. It's subjective, but you're right that the latter is probably clearer.
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  5. @Razib Khan
    The dragon gene is X-linked and codominant.


    codominance means a mix of traits
    https://en.wikipedia.org/wiki/Dominance_(genetics)#/media/File:Co-dominance_Rhododendron.jpg

    If you see trait: dragonrider and trait: not-dragonrider as *not* both being traits, then you’d not use the term co-dominant, that’s true. If so, you’re right, you’d use the term incomplete dominant. It’s subjective, but you’re right that the latter is probably clearer.

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  6. As you can see Daenerys’ parents and grandparents were full siblings. Using a pedigree method her Wright’s coefficient of inbreeding would be 0.375. For a point of comparison, the child of parents who were full siblings, but whose own parents were unrelated, would have a coefficient of inbreeding of 0.25. The terminal individual in the genealogy of the Spanish Habsburgs, often cited as a case study of how inbreeding can lead to the extinction of a lineage through sterility and imbecility, had a Wright’s coefficient of inbreeding of 0.254. Something really doesn’t add up in terms of the viability of the offspring of the Mad King and his sister-wife (to get a sense of what multi-generation inbreeding might do, please see the Colt family).

    It’s happened before. Amenhotep I’s parents and grandparents were also full siblings. His father, Ahmose I, died young (in his 30s) but appears to have been an exceptionally capable ruler. Amenhotep I himself married his own full sister, and unsurprisingly wasn’t able to have viable children with her. He also seems to have been a strong ruler, though. So in this case, sterility did become an issue but imbecility didn’t, and the 0.375 inbreeding coefficient was achieved.

    (It’s also possible that imbecility was an issue and Egyptian histories carefully covered it up; they are known to do that kind of thing. But the history I read ( https://www.amazon.com/Rise-Fall-Ancient-Egypt/dp/0553384902/ ) didn’t even mention this as a theory.)

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  7. bob sykes says:

    She’s also fire-proof. Is that also linked to the dragon-master gene?

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    • Replies: @Shaikorth
    GRRM has said that has a magical explanation:

    The thing with Dany and the dragons, that was just a one-time magical event, very special and unique. The Targaryans can tolerate a bit more heat than most ordinary people, they like really hot baths and things like that, but that doesn't mean they're totally immune to fire, no. Dragons, on the other hand, are pretty much immune to fire.
     
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  8. Bran Vras says:

    Not so fast. Consider what Viserys declared in the very first Dany chapter.

    For centuries the Targaryens had married brother to sister, since Aegon the Conqueror had taken his sisters to bride. The line must be kept pure, Viserys had told her a thousand times; theirs was the kingsblood, the golden blood of old Valyria, the blood of the dragon. Dragons did not mate with the beasts of the field, and Targaryens did not mingle their blood with that of lesser men. Yet now Viserys schemed to sell her to a stranger, a barbarian.

    Surely Viserys knew that his only greatgrandmother was a Blackwood. So why did he insist on blood purity? GRRM knows what he is doing by having Blackwoods, Daynes etc in the genealogical tree.

    There might be something clever at work. I can’t tell what for sure. But in the World of Ice and Fire, I noticed quite a few Targaryen girls unaccounted for. The look of the Daynes hints at It’s Valyrian blood (most likely Targaryen). Some speculate as much for the Dondarrions. It’s perfectly possible that the blood of the dragons made its way to the Blackwoods. After all, Bloodraven is a royal bastard with a Blackwood mother, and he was at the peak of his power when Aegon V married Betha Blackwood.

    The wilder speculation I entertain is that the true blood of the Dragon is composed of a Y chromosome and its female counterpart. Every king on the Iron Throne up to Aerys was descended from father to son from Aegon the Conqueror. Perhaps there is something to be found on the matrilineal line passing through Betha Blackwood (the dragon taming requires only one Targaryen parent though). GRRM is not adverse to playing subtle genealogical games (see Harry the Heir in Feast for Crows).

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    • Replies: @Razib Khan
    Y chromosome and its female counterpart.

    huh? female counterpart?
    , @CupOfCanada
    The Dayne point is interesting.

    So I'd just point out that Targaryens, and even Valyrians, weren't necessarily the only dragonriders:

    http://awoiaf.westeros.org/index.php/Dragonseed

    Many seem explainable as Targaryen or Valyrian bastards, but Nettles does not seem to have Targaryen ancestry.

    I kind of assume that the trait is linked to if not the same as the trait for warging. That trait seems to be pretty prevalent among Blackwoods as well (hence Brynden Rivers), and perhaps First Men in general (the Daynes?).

    The Daynes having Targaryen features is a very good and interesting point though. It certainly helps explain Danaerys' appearance.
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  9. Shaikorth says:
    @bob sykes
    She's also fire-proof. Is that also linked to the dragon-master gene?

    GRRM has said that has a magical explanation:

    The thing with Dany and the dragons, that was just a one-time magical event, very special and unique. The Targaryans can tolerate a bit more heat than most ordinary people, they like really hot baths and things like that, but that doesn’t mean they’re totally immune to fire, no. Dragons, on the other hand, are pretty much immune to fire.

    Read More
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  10. @Bran Vras
    Not so fast. Consider what Viserys declared in the very first Dany chapter.

    For centuries the Targaryens had married brother to sister, since Aegon the Conqueror had taken his sisters to bride. The line must be kept pure, Viserys had told her a thousand times; theirs was the kingsblood, the golden blood of old Valyria, the blood of the dragon. Dragons did not mate with the beasts of the field, and Targaryens did not mingle their blood with that of lesser men. Yet now Viserys schemed to sell her to a stranger, a barbarian.

    Surely Viserys knew that his only greatgrandmother was a Blackwood. So why did he insist on blood purity? GRRM knows what he is doing by having Blackwoods, Daynes etc in the genealogical tree.

    There might be something clever at work. I can't tell what for sure. But in the World of Ice and Fire, I noticed quite a few Targaryen girls unaccounted for. The look of the Daynes hints at It's Valyrian blood (most likely Targaryen). Some speculate as much for the Dondarrions. It's perfectly possible that the blood of the dragons made its way to the Blackwoods. After all, Bloodraven is a royal bastard with a Blackwood mother, and he was at the peak of his power when Aegon V married Betha Blackwood.

    The wilder speculation I entertain is that the true blood of the Dragon is composed of a Y chromosome and its female counterpart. Every king on the Iron Throne up to Aerys was descended from father to son from Aegon the Conqueror. Perhaps there is something to be found on the matrilineal line passing through Betha Blackwood (the dragon taming requires only one Targaryen parent though). GRRM is not adverse to playing subtle genealogical games (see Harry the Heir in Feast for Crows).

    Y chromosome and its female counterpart.

    huh? female counterpart?

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  11. Anonymous says: • Disclaimer

    No, she not “fire-proof”, she can resist lot more heat than normal people, stop with this nonsense.

    Dragon bending is not associate with X genes, look away back at targaryen family tree, you see that Aegon I mother is a Velaryon, valyrian house know for not to be dragonriders.

    Also, sorry for any english writing and grammar mistake.

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  12. The Spanish Habsburgs were not that isolated. They married cousins, they also married other European Royalty, which is how they came to rule Spain. Maximilian married Mary of Burgundy. Their son Philip the Fair married Juana (la Loca) daughter of Ferdinand and Isabella, and became the king of Spain Jura uxorious. Their son Charles V was Holy Roman Emperor, King of Spain, and Duke of Burgundy. As such, he ruled Spain, the Low Countries, Western France, Most of Italy, Germany, Austria, the New World, and the Philippines. When Charles retired, he gave the HRE to his brother Ferdinand and Spain, the Low Countries, and the Spanish Colonies to his son Philip.

    It is true that the descendants of Philip were inbred, but the inbreeding was with the Austrian branch of the family. There were, and still are, plenty of Austrian Habsburgs. The reason the Habsurgs lost control of the Spanish throne is that when Charles II of Spain died childless, he willed the kingdom to a French Bourbon one of Louis XIV’s grandchildren. There was a war, the War of the Spanish Succession, the result of which was that the Bourbons kept the Spanish Throne, but that the Bourbon heir had to renounce his claim to the French throne.

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  13. These many Razib posts made me buy a five book box set of GRRM’s books. I hope they won’t disappoint. Probably they won’t, because I won’t have time to read them anyway.

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  14. @Bran Vras
    Not so fast. Consider what Viserys declared in the very first Dany chapter.

    For centuries the Targaryens had married brother to sister, since Aegon the Conqueror had taken his sisters to bride. The line must be kept pure, Viserys had told her a thousand times; theirs was the kingsblood, the golden blood of old Valyria, the blood of the dragon. Dragons did not mate with the beasts of the field, and Targaryens did not mingle their blood with that of lesser men. Yet now Viserys schemed to sell her to a stranger, a barbarian.

    Surely Viserys knew that his only greatgrandmother was a Blackwood. So why did he insist on blood purity? GRRM knows what he is doing by having Blackwoods, Daynes etc in the genealogical tree.

    There might be something clever at work. I can't tell what for sure. But in the World of Ice and Fire, I noticed quite a few Targaryen girls unaccounted for. The look of the Daynes hints at It's Valyrian blood (most likely Targaryen). Some speculate as much for the Dondarrions. It's perfectly possible that the blood of the dragons made its way to the Blackwoods. After all, Bloodraven is a royal bastard with a Blackwood mother, and he was at the peak of his power when Aegon V married Betha Blackwood.

    The wilder speculation I entertain is that the true blood of the Dragon is composed of a Y chromosome and its female counterpart. Every king on the Iron Throne up to Aerys was descended from father to son from Aegon the Conqueror. Perhaps there is something to be found on the matrilineal line passing through Betha Blackwood (the dragon taming requires only one Targaryen parent though). GRRM is not adverse to playing subtle genealogical games (see Harry the Heir in Feast for Crows).

    The Dayne point is interesting.

    So I’d just point out that Targaryens, and even Valyrians, weren’t necessarily the only dragonriders:

    http://awoiaf.westeros.org/index.php/Dragonseed

    Many seem explainable as Targaryen or Valyrian bastards, but Nettles does not seem to have Targaryen ancestry.

    I kind of assume that the trait is linked to if not the same as the trait for warging. That trait seems to be pretty prevalent among Blackwoods as well (hence Brynden Rivers), and perhaps First Men in general (the Daynes?).

    The Daynes having Targaryen features is a very good and interesting point though. It certainly helps explain Danaerys’ appearance.

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  15. ohwilleke says: • Website

    “Therefore, Daenery’s two grandparents were both 1/8th Valyrian, her parents were 1/8th Valyrian, and she is 1/8th Valyrian (at least that’s the expected value).”

    From a true GOT nerd like you with your genetics expertise, we expect confidence intervals, not just mean expected values. ;)

    Also, how inbred are pedigreed dogs in early generations?

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  16. grmbl says:

    The actress who plays Daenerys on TV does look inbred, in a Euro blueblood kind of way.

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  17. sprfls says:

    Another question is exactly how much “blood of the First Men” exists in the north. ASOIAF admixture calculator would be cool. ;-)

    Warging, greenseeing, etc. obviously exist in much larger numbers in the north. I believe these traits pre-date even the First Men and instead relate to the Children or other archaics living at the time. So there must have been a mixing event between these populations, and that some of that archaic ancestry exists in the present — but pretty much only in north mediated by people with “First Men” ancestry.

    In addition to warging and greenseeing, Hodor having “giant’s blood” is another allusion.

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  18. Anonymous says: • Website • Disclaimer
    @Razib Khan
    would that weed out some of the bad consequences of the inbreeding over time?

    possibly. this works with plants (load purging). but it doesn't seem to work too well in metazoans. one of the issues is if you select really strongly u start to reduce effective population a lot and that introduces drift.

    also, they weren't sequencing. so they would get a lot of obviously ill infants, but a lot of deleterious traits show up an older ages. for example, several of the fritzl kid's have some health issues which are treatable with modern medicine

    https://en.wikipedia.org/wiki/Fritzl_case#Elisabeth_Fritzl.27s_confinement

    kerstin's only showed up in her teen years.

    >possibly

    If you haven’t, check out Helen Dean King’s successful, multi-year work on extreme inbreeding, from early 20th century. Her lines of nearly-homozygous rats are ancestral to approximately half of modern lab rats. The entire journal is accessible on Google Books:

    https://books.google.com/books?id=5WDXAAAAMAAJ

    She started getting very good results by around sixteenth generation. Obviously, doing inbreeding correctly requires progeny maximization and strict arbitration as to who goes to make the next round, for load purging to have its strongest effect. It’s more-or-less doable on rats, but of course humans are a whole different story.

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  19. This is not only common but expected in royal lineages!

    For the Spanish Hapsburgs to escape their Teutonic Austro-Hungarian roots, going to their close French aristocracy (de Borbon) created a way out of the medical health connundrum with a big assist from the disintegration of their former Empires and the World War 1 and 2 gig with a greater benificiary being the UK with the Battenberg/Mountbatten foresight of Queen Victoria.

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