DNA Shows that KV55 Mummy Probably Not Akhenaten

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Lid geworden op: vr 23 jan 2009, 19:52

DNA Shows that KV55 Mummy Probably Not Akhenaten

Bericht door king tutankhamun » ma 04 okt 2010, 14:15

Abstract

The paper Ancestry and Pathology in King Tutankhamun's Family
by Hawass al. (Journal of American Medicine, 2010 - JAMA. 2010;303(7):638-647), states that the mummy in KV55 is “probably” Akhenaten – hereafter “the JAMA paper”. The media has accepted the attribution as affirmed fact, although the attribution has attracted considerable comment and debate with a number of writers questioning the forensic data. I believe, however, that the correct focus of dissent to the attribution should be the STR analysis which shows that the KV55 mummy is highly unlikely to be Akhenaten and that an alternative family tree is a better fit to the genetic findings of the Hawass study.

(As this article is longer than normal, if you are reading this abstract on the front page of the blog then please click here or on the article title to display the full article below this point.)


Background

In the JAMA paper, Hawass et al. outline what they see as the most likely 5-generational family tree including the identification of the KV55 mummy and KV35YL as full siblings. I accept the family tree with the exception of the sibling identification which I believe should be questioned and the identification of Akhenaten.

The Hawass theory hypothecates KV35YL (“the Younger Lady”) as a previously unknown wife of Akhenaten. As his sister, the lady was of royal birth. As she gave birth to a son, who presumably was the heir apparent, by convention she was probably a queen. We are therefore asked to hypothecate an unknown queen who avoided mention in any of the surviving reliefs, although Kiya, known to be despised by Nefertiti, was mentioned on a number of occasions.

The paper also identifies that the Younger Lady was killed, probably murdered, by a sharp blow to the face. Throughout history the murder of a queen has been a serious matter. It is hard to envisage circumstances in which this queen, rather than the more prominent Kiya or Nefertiti would be a murder victim, especially since her son, Tutankhamun, remained alive.

Age at Death of the KV55 Mummy

The age at death of the KV55 mummy has attracted considerable debate. The forensic assessment of the age at death based upon skeletal and dental analysis remains contentious. Since this paper identifies that the KV55 mummy is not Akhenaten on genetic grounds, I don’t discuss the forensic assessment at the age of death in detail. However, I note in passing that once the identification of KV55 as Akhenaten is set aside, the this forensic data ceases to be problematic – at least for now – as so little is known of Smenkhare (if indeed the KV55 mummy is even he).

Ankhesenamun?

History records only a single wife for Tutankhamun – Ankhesenamun – whose image and name is prominent on a number of items found in KV62, Tutankhamun’s tomb in the Valley of the Kings. It is tempting to indentify the two foetuses from that tomb as Ankhesenamun’s daughters. History is clear that Ankhesenamun was the [third] daughter of Akhenaten and Nefertiti. If the KV55 mummy is Akhenaten then as well as being the paternal grandfather of the foetuses (via Tutankhamun), then he must also be the maternal grandfather (via Ankhesenamun). However, this contradicts the genetic data. Examination of the STR analysis for locus D7S820 shows that Foetus 1 presented characteristics (alleles) 10 and 13 and Foetus 2 characteristics 6,15. Tutankhamun (the father), presented 10 and 15 so the other characteristics (alleles) must have been inherited from their mother (Ankhesenamun), who must therefore have presented characteristics 6 and 13.

Since one of each pair of alleles is inherited from the father and one from the mother, her father (Akhenaten) must have presented either 6 or 13. The KV55 mummy presents neither of these and is instead the diploid 15,15. The KV55 mummy cannot be the maternal grandfather of these mummies, ie it cannot be Akhenaten if Ankhesenamun is the mother of the foetuses.

This contradiction arises without consideration as to whether the KV21A mummy is Ankhesenamun or not, or indeed whether the KV21A mummy is the mother of the two foetuses (although it seems likely that she is the mother). The contradiction could be resolved by hypothecating that the foetuses had different mothers, or that their mother was not a daughter of Akhenaten and Nefertiti. In either case, this would require us to believe that Tutankhamun had another wife who went without mention in his tomb. As his tomb was found almost intact we must conclude that no mention of this other wife was recorded when the tomb was closed. While an additional wife is far from impossible, hypothecating an unknown wife (and probably queen) for Tutankhamun whose foetuses were buried with Tutankhamun but who herself was unmentioned must surely be problematic.

In short the Hawass theory requires the hypothecating of unknown queens for both Akhenaten and Tutankhamun. While not impossible, it casts serious doubt upon the Hawass theory that KV55 is Akhenaten.

Enter Smenkhare?

The contradiction in the D7S820 STR results, and the need to hypothecate either additional queen, can be resolved by instead identifying the KV55 mummy as an unknown prince. There is no way to name this prince but he would seem to fit what is known of Smenkhare and, for convenience at least, I’ll work with this attribution throughout the rest of this paper.

The mummy of Akhenaten would remain missing or unidentified. Indeed the question would remain whether either he or his wife, Nefertiti, was ever returned to the Valley of the Kings from Amarna, although science hopes they were and that their last resting place might be found. (Romantics might wish to find them interred together.)


Multi-inter-generational Allele Jumps

In Table 1 below I have repeated the STR findings for 6 loci reported in the JAMA paper: D13S317, D7S820, D2S1338, D21S11, D16S539 and D18S51. I have omitted the final two loci tested as the findings are materially incomplete in the latest generations so they add little to this discourse. (I have nonetheless checked that the reported alleles in those loci are consistent with my conclusions in this paper.) I have added my deduced assessment of the genetics of Akhenaten and Nefertiti (both mummies unknown) as I will explain later.


Afbeelding
[Author's note: if you are having trouble reading this table, my apologies. Please see the Typographical Note at the bottom of the paper which presents a bare copy without formatting. Alternatively click on the table to see the source image which is also much cleaner (and retains the formatting).]

For now, I would draw your attention again to the 13-characteristic in the D7S820 locus. This is present in the 1st generation (Thuya) and the 5th generation (Foetus 1). I have examined a number of papers to assess the frequency of the 13-allele:

Paper Incidence
Population Data on the STR Loci D7S820, D8S1179, and D12S391 in a Sample Population of Rio de Janeiro, Brazil (Gomes et al, Forensic Science Communications, January 2004 - Volume 5 - Number 1) 3.09%
Allelic Frequency Distribution of Four STR Loci (FGA, D7S820, D1S533, D9S304) in the Argentinean Population (Raimondi, E, Toscanini, U, Haas, E.) 2.81%
Analysis of the three STR loci (D16S539, D7S820, D13S317) in a population sample of Marmara region of Turkey (A. Hadi Çak›r et al, Journal of Cell and Molecular Biology 1: 25-30, 2002) 2.00%
Table 2 - References for the D7S820 allele distribution


This locus has been extensively studied. The most comprehensive analysis I have found was published by the Heinrich-Heine-Universität Düsseldorf which states an incidence for modern Egypt of 1.32%. (The university’s databank is best accessed via its front page.)

The possibility that the 13-allele has re-appeared spontaneously in the 5th generation (actually in the 4th generation as the mother [Ankhesenamun] must also have presented this allele) is therefore of the order of 1.3% - 2%.

There is a similar pattern with allele-35 at locus D21S11 Again this is allele is presented by Thuya but does not appear again until the 4th generation in the KV21A mummy, as well as in the 5th generation in Foetus 2. Heinrich-Heine-Universität Düsseldorf again has collected research into the distribution of this allele. In this case, this allele is undetectable in many populations but the incidence within the modern Egyptian population is shown as 1.1%, one of the highest incidences worldwide and a likely upper-bound for the incidence in the population of ancient Egypt. The chance of this allele being reintroduced into the 4th generation from the general population is therefore of the order of 1%.

In combination, the chance of both alleles being reintroduced from the general population is therefore of the order of 0.02%. Given the consanguinity of the royal family, it would be inappropriate to regard this as precise; however, it does give some sense that the re-introduction of both alleles from outside the royal family is extremely unlikely. (The incidence of these alleles within the ancient Egyptian nobility is unknown.) Put another way, there is a very high probability (about 99.98% ignoring the effects of consanguinity and the difficulty of relying on modern population statistics) of a secondary ancestral line from Thuya to the KV21A mummy. This is consistent with theories that Nefertiti was the grand-daughter of Thuya (potentially via Ay). (An alternative woud be an additional acestral line from one of Thuya's parents or grandparents which each successive remove in generations becoming less likely.)


Allele Jumps in the 3rd Generation

In addition to the allele jumps from Thuya to KV21A/foetuses examined in the previous section, allele 16 at the D13S317 locus, and allele 6 at the D7S820 locus jump from Amenhotep III to the foetuses. (Although the Younger Lady presents allele 6, there is no suggestion that she is an ancestor of the foetuses other than as their paternal grandmother via Tutankhamun who did not present this allele.) Referring again to the Heinrich-Heine-Universität databank, the frequencies of D13S17-16 and D7S820-6 in the general modern Egyptian population is immeasurable in a sample population (ie 0%) in both cases. Clearly the incidence within a population is non-zero: it is just immeasurably low in a study with a typical sample size. It may therefore be reasonable to state that the incidence of each allele is not expected to exceed 0.1% in the general population.

Again, the chance therefore that these rare alleles have re-appeared in the foetuses from the general population, rather than inherited via Amenhotep III (or Yuya in the case of D7S820) is extremely unlikely. That is, there is a 99.99%+ chance that the foetuses have inherited these alleles from these earlier generations rather than from an unknown progenitor from outside this familial group. In the case of D7S820-6, inheritance could have been from Amenhotep III or from Yuya via one of their children other than AmenhotepIII; however, neither Yuya nor Thuya presented D13S17-16 so there is a high (99%+) chance of a secondary ancestral pathway from Amenhotep III to the KV21A and Foetus 1.

Taken together with the allele jumps from Thuya's generation described in the previous section, then the probability of a secondary royal lineage for these foetuses is so high that we should regard it as proven, subject to our caveats about the paucity of DNA loci.

It is worth recapping at this point that if Ankhensenamun cannot be the mother of the LV62 foetuses if KV55 is her father. However, in order to resolve the secondary royal lineage we find ourselves drawing on historical evidence of Nefertiti and her daughters in explanation. The genetic evidence also favours a son of Amenhotep III as the foetuses' maternal grandfather. Taken together, the two lines of evidence make it exceedingly difficult to continue to attribute KV55 as Akhenaten as it directly contradicts the DNA evidence.


Caveat – Unkown Family Members

Before we reconstruct a family tree which can explain these additional ancestral pathways it is important to recognise my proposal is merely the simplest possible family tree which explains the findings of the STR analysis. It is possible, if unlikely, that these alleles were carried by other members of the earlier generations and passed via peripheral members of the royal family, one of whom was the mother or father of KV21A. It is difficult to reconcile this with what we know of the history of the royal family, although we cannot rule out the possibility entirely.

We should also remember that only a small number of loci were analysed and for this reason all conclusions are potentially invalid, that’s especially the case when inferring descent via an even smaller number of loci. This is a difficulty with the JAMA conclusions altogether that has been raised by other commentators. However, perhaps instead of worrying that we have only small amounts of DNA data to work with, given the antiquity (and in some case the poor state of preservation) of the mummies involve, we should perhaps consider ourselves fortunate to have as much data as the team did manage to resolve. It seems probable that the study of such ancient DNA, as the science evolves, might require different approaches to extracting mazimum value from limited DNA than is customary in contemporary forensic assessments.

For these reasons I have described my hypothesis merely as a theory. I regard nothing as proven.

Reconstructing the Amarnan Family Tree

We are therefore faced with 3 features not explained by the Hawass family tree:

* KV55 cannot be Akhenaten if Ankhesenamun (or one of her sisters) is the mother of the two foetuses found in tomb KV62.

* There is a very strong probability that Thuya is the ancestress of the foetuses (and KV21A) other than via Amenhotep III.

* There is a strong probability that Amenhotep III is the great-grandfather of the foetuses (and grandfather of KV21A) not via Tutankhamun.


It should be noted again that we are working with limited data on only 8 alleles, two of which have revealed little information about the foetuses and their mother(s). To some degree any family tree must be tentative; however, I do not believe that the putative family tree presented in the JAMA paper is a good fit for the findings, and certainly not the best fit. Moreover, I propose that a classic chronology does fit the data and we could envisage a family tree as follows in Figure 1. (My apologies, Word is a terrible word processor when it comes to constructing family trees.)
Afbeelding


If we identify the KV55 mummy as Smenkhare and assume that Akhenaten remains missing, we can add Akhenaten and Nefertiti into the family tree, while retaining KV55 (Smenkhare) as the father of Tutankhamun as shown by the Hawass team. It’s tempting to consider that KV21B and the Younger Lady are also daughters of Akhenaten and Nefertiti. While not essential for my revised theory, this would neatly fit historical facts. In considering whether the DNA data would fit the revised family tree, it seems appropriate to consider that KV21B and the Younger Lady may also have been daughters of Akhenaten and Nefertiti as this represents the tightest constraint. That is, if the DNA data fits an assumption that these two queens/princesses were also daughters of Akhenaten and Nefertiti then if they aren’t daughters the theory wouldn’t be invalidated.

Table 1, above, shows that it is possible to build a family tree with all these constraints and predict the DNA of Akhenaten and Nefertiti. In the JAMA paper, the Younger Lady is shown as a daughter of Amenhotep III and Queen Tiye. Table 1 shows that it is equally possible for her to be a daughter of Akhenaten ie a niece of Amenhotep III. I have written before that the JAMA paper ignored how consanguinity can change the probability of relationships and could wrongly conclude that a particular relationship was proven, when other relationships were possible.

(Note, if any of these constraints are removed, a greater range of possibilities exist for the DNA of Akhenaten and Nefertiti. Conversely, if their mummies are found and DNA extracted, then the findings might eliminate the Younger Lady and or KV21B from consideration as their daughters.)

Queen Meritaten?

Just as we cannot name the KV55 mummy, the identity of the Younger Lady must also remain unknown. However, it is tempting to consider that she is one of the daughters of Akhenaten and Nefertiti. Meketaten is known to have died in child birth, so Meritaten seems a strong candidate. It is known she was married to Smenkhare. Again, this is just conjecture, although it seems helpful - indeed essential - at all stages to check that the proposed family tree could fit known historical facts.

King Ay

It is also possible to consider that the Pharaoh Ay is the missing link between Yuya/Thuya and Nefertiti. It is possible within the context of the DNA data and I have shown this in the putative family tree, but it’s really impossible to do more than conjecture. Indeed, there is no reason from the DNA data to assume that Nefertiti’s ancestry from Yuya and Thuya was via her father rather than her mother. Nonetheless, if the DNA of Ay was tested, it would be possible to establish whether, within my theory, he could be the father of Nefertiti even without Nefertiti’s own mummy because we can predict Nefertiti’s DNA which such accuracy from her surrounding family. However, some commentators wonder whether Ay the son of Yuya and Thuya was the same Ay who succeeded Tutankhamun to the throne.


Mitochondrial DNA

The JAMA paper records that the team also established paternal lineage of the male mummies by sequencing 16 loci on the Y chromosome. Work has started on examining the hyper-variable regions of mitochondrial DNA (mtDNA) but no results have been published. It will be interesting what this reveals. In the Hawass theory, the Younger Lady is a daughter of Amenhotep III and Queen Tiye and therefore inherits mtDNA from Tiye and Thuya. In my theory, the Younger Lady could be a daughter of Nefertiti (although this is not central to my theory) and Nefertiti could be descended from Yuya and Thuya via her father and potentially therefore have different mtDNA - notwithstanding that we know nothing of her mother. While the Younger Lady and Tiye sharing the same mtDNA would not make any difference to my theory, if they have different mtDNA it would be damaging to the Hawass theory. Likewise, in my theory, if the Younger Lady is another daughter of Nefertiti, then she and the foetuses should share mtDNA which would not be explained by the Hawass theory within the context of the family tree shown in the JAMA paper.

An Alternative Theory on the Parentage of KV55

Some people have conjectured that Nefertiti might briefly have been the consort of Amenhotep III before marrying Akhenaten. I have therefore considered whether Amenhotep III and Nefertiti (as I have deduced her DNA) could be the parents on the KV55 mummy. I have reservations about whether this is chronologically possible, but it fits all but one of the loci in my analysis.

The difficulty lies with D7S820. As previously seen, the mother of the foetuses presented as 6,13, so she must have inherited one of this pair from each of her parents. As the son of Amenhotep III and Queen Tiye, Akhenaten could not have inherited allele 13 from either parent and therefore Nefertiti must have presented allele 13 at the D7S820 locus - and Akhenaten therefore allele 6. If the Younger Lady is another daughter of Akhenaten and Nefertiti, then likewise her alleles of 6,10 must have been inherited one from each parent. Therefore Nefertiti’s alleles at this locus must either have been 6,13 or 10,13. It would therefore be impossible for Amenhotep III and Nefertiti to be the parents of the KV55 mummy whose alleles are a duplicated 15. Therefore, if the Younger Lady is a daughter of Akhenaten and Nefertiti she cannot be the mother of the KV55 mummy (Smenkhare), although nothing can be deduced about whether or not she was the consort of Amenhotep III. However, if it was assumed, or proven, that the Younger Lady was not the daughter of Nefertiti then I could not rule out that Nefertiti was the mother of the KV55 mummy: she can be the mother of either but not both. Personally I am inclined to follow the JAMA paper in accepting Amenhotep III and Tiye as the parents of the KV55 mummy (Smenkhare) but the alternative cannot be ruled out on the basis of the published STR analysis. Again, mtDNA analysis may be helpful in excluding the possibility of some alternative theories like this one.



Conclusions

* Combining the DNA data with known historical facts shows that Akhenaten is probably not the KV55 mummy.

* It is tempting to consider the KV55 mummy (Tutankhamun’s father) alternatively as Smenkhare although this can be no more than conjecture on the basis of the available data.

* There is a very strong probability of a second line of descent from Yuya and Thuya to the KV62 foetuses.

* This secondary line of descent is consistent with the historical Nefertiti.

* There is a strong probability of second line of descent from Amenhotep III to the foetuses not via the KV55 mummy.

* This second line of descent is consistent with the historical Akhenaten.

* It is possible to construct a family tree along these lines which fully fits the STR analysis published in the JAMA paper and which assumes Nefertiti is a granddaughter of Yuya and Thuya.

* It is possible, but not essential, to accommodate the Younger Lady and KV21B mummies in this revised family tree as further daughters of Akhenaten and Nefertiti.


Afterword

This paper has been a real stretch for me. I could be wrong. If I am, I won’t mind anybody telling me so – if you explain why! At least I will have learned something new. One advantage of being an amateur with no reputation or credentials is that a bit of embarrassment will not harm me.

On the other hand, if you think I may be right, then I would appreciate you cross-linking and promoting this article. I don’t often ask, but in this case I would like the widest possible circulation and critique to determine whether my theory is viable. If you are competent to assess what I have written, then writing in support would be very much appreciated.

It is also possible that there is unpublished data which the authors of the JAMA report used to inform their conclusions and which invalidates the theory I have presented in this paper. If so, then I look forward to seeing it.

Although I disagree with one or two detailed findings of the JAMA paper, the true legacy of Dr Hawass and his colleagues shouldn’t be seen as the specific conclusions drawn but that they have opened an entirely new window on Egyptology through the assessment of mummy DNA. I have disagreed with one minor aspect of their paper, which in the paper itself rates only a "probably" in the paper itself, and would like to pay testimony to the standard of their work and thank them for a considerable advancement in our knoweldge of the late 18th Dynasty.

Typographical Note

I apologise for adding Table 1 as a graphic. I wrote the paper in Word and there is no easy way of converting tables from Word into HTML - that I know of. Since there were links in Table 2 I hand-wrote the HTML for that table but that would have been too onerous for Table 1. If you find it hard to view, click in it so that it is displayed alone in your browser. For me it is then easy to read. Subsequently I have managed to grab it as an HTML table sans formatting, so I have also presented this below. I'll edit the post to add the key highlighting tomorrow.

As my regyular readers know, I get word blind when I write something so it will take me a few days to iron out all of the typos. Where these are just obvious typos and don't change the sense of the paper, I'll just make them inline without comment. I will highlight any revisions to the substance.


D13S317 D7S820 D2S1338 D21S11 D16S539 D18S51
Thuya (KV46) 9,12 10,13 19,26 26,35 11,13 8,19
Yuya (KV46) 11,13 6,15 22,27 29,34 6,10 12,22
Tiye (KV35EL) 11,12 10,15 22,26 26,29 6,11 19,22
Amenhotep III (KV35) 10,16 6,15 16,27 25,34 8,13 16,22
Akhenaten (N/K) 12,16 6, {10,15} 16,26 25,

{26,29} 8,11 16,

{19,22}
Nefertiti (N/K) 10,

{9,11, 12,13} 13,

{6,10} 17,26 29,35 11,13 10,19
Smenkhare? (KV55) 10,12 15,15 16,26 29,34 11,13 16,19
Younger Lady (KV35YL) 10,12 6,10 16,26 25,29 8,11 16,19
Tutankhamun (KV62) 10,12 10,15 16,26 29,34 8,13 19,19
Ankhesenamun? (KV21A) 10,16 - ,26 ,35 8, 10
? (KV21B) 10 - 17,26 - 11,13 -
Foetus 1 (KV62) 12,16 10,13 16, 29, 8, ,19
Foetus 2 (KV62) 10, 6,15 ,26 29,35 8,13 10,19
Table 1 – Extended STR analysis interpolated to add Akhenaten and Nefertiti
XX alleles demonstrating KV55 mummy cannot be maternal grandfather of foetuses

XX alleles which have “jumped” generations in the Hawass theory


de link <greensmile>
http://www.kv64.info/2010/03/dna-shows-that-kv55-mummy-probably-not.html
the defender of her royal highness, the supreme queen of egypt, Ankhesenamun :)

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