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The mystery of Lactase Persistence (LP) in Europeans

It comes with farming, The first evidence of animal-related diseases is Tuberculosis identified in remains of 7,000 BC Greek early farmers. Farming populations gradually build up resistance, by means of an intensive selection process (so their population drops, goes up again, drops, etc.), and after some time they are immune but still carry the germs. The biggest killer of all, btw., is still today the flu (via poultry, pigs, but also cattle), but most Europeans now survive it. Native Americans didn't. And by 6,000 BC, HGs were still quite isolated, and met East Mediterranean early farmers that already had gone through 1-2 millenniums of genetically building up resistance.
I heard that few times but I don't really buy it. Domesticated animals, especially at the beginning of farming, were exactly the same as their wild cousins. HG had contact with them all the time and if unlucky they caught infections from them. They were actually infested with parasites all the time; hookworms, brainworms, flatworm, etc. Wild boar was always the favorite food of HG of central europe. I'm sure they didn't need to wait till Neolithic to catch the flu.

Europe had "American" moments with the plague, which seems to be endemic around the Himalaya, with the Antonine Plague (probably a henceforth unknown strain of smallpox or measles), most likely also when Malaria was imported into the Mediterranean from Sub-Saharan Africa by migrants (the Anopheles mosquito was probably already around, but it only starts transmitting after it has bitten an infected person).
There is a nice study in German on diseases they identified in Slovakian and East German bronze-age grave fields: Lepra, Syphhillis, Borrelliose / Meningitis, 50% with chronical respiratory infections (possibly also due to the flu), and around 25% with symptoms of arsenic poisoning from working with arsenic bronze, or, especially the women, carrying arsenic bonze necklaces and earrings. Quite a lot of Vitamin B and C shortage as well (actually the first problem encountered by early farmers when they switch to a cereal-based diet, the second one is caries).
I agree, no matter what continent people live on, from time to time a nasty sickness comes and can kills most of the population.
 
I heard that few times but I don't really buy it. Domesticated animals, especially at the beginning of farming, were exactly the same as their wild cousins. HG had contact with them all the time and if unlucky they caught infections from them. They were actually infested with parasites all the time; hookworms, brainworms, flatworm, etc. Wild boar was always the favorite food of HG of central europe. I'm sure they didn't need to wait till Neolithic to catch the flu.

Influenza is actually a highly interesting case. It spreads from mammals by aerosols (sneezing), so infection risks from domesticized (living) animals are much higher than from (dead) hunting prey. Mammal viruses can survive outside the body, but on skin at 20°C for only five minutes. Thus, the risk of a south European HG of catching the flu from hunted boar (anyway only the third common prey, far behind Aurochs and deer) is minimal. An Anatolian pig or poultry farmer, OTOH, should have had quite an exposure, especially as poultry feces are also infective and may stay so for several days. Another issue here is that swine is susceptive to both bird and mammal flu, which makes it the perfect vessel for turning one variant into the other one. As such, by the time of spreading to the Central Mediterranean early farmers should already have had quite some infection history.

Survival of mammal influenza viruses outside the body depends on four factors:
  1. Temperature: The cooler, the better. At 0°, the virus can survive up to 30 day, in ice permanently. It is killed at around 60°C.
  2. Humidity: The dryer, the better. In aerosols at 21°C, it can survive over one hour below 40% humidity, less than half an hour at 70% humidity.
  3. UV radiation: Though I haven't read details, higher levels of UV radiation also appear to reduce survival times.
  4. Medium/ surface: In sweetwater (22°) it can survive more than 4 days, 2-4 days on smooth surfaces (plastics, steel), 8-12 hours on textiles or paper, and only 5 minutes on skin (that may also be temperature-related, though).
For these reasons, flu epidemics mostly occur during late autumn/ winter in moderate to cold climate zones. Vitamin D, production of which is stimulated by sunlight, seems to protect against influenza, and may be another factor for seasonal variation in infection.

The main virus reservoir appear to be birds, especially waterfowl and gulls, from where it may have spread to other fowl (chicken, turkey etc.). Alternatively, deer and horses are considered (German research from 1980 found antibodies against a specific strain in wild deer, which predate corresponding human and swine infections later that year). Seals and bats appear to have similar recombination abilities as swine/boar. Aside from the aforementioned, relevant secondary hosts include whales, reindeer, sheep, goat, camels, dogs/ wolves, bears, ferrets / polecats, marten, minks, raccoons, possibly also squirrels.
http://www.nytimes.com/2012/07/31/s...o-seals-is-studied-for-human-threat.html?_r=0
As such, I agree that north Eurasian HGs most likely had caught the flu, and developed some immunity from it, already before the entrance of early farmers. Climate is one factor here, the other one is the reliance on seal, deer, reindeer and wild horses as hunting prey. Moreover, North Eurasian HGs may have picked up flu viruses from seal-infected brackish water (Baltic Sea!), or from ice (igloo construction etc.). Domestication of wolves/ dogs as hunting or traction animals should also be considered. In fact, north Eurasian HGs' exposition, immunity and transmission potential should have been substantially higher than that of EEFs.

The human defence against influenza is based on various antibodies that are produced in plasma cells, but also included in the mother milk to provide the infant with basic protection as long as its own immune system is still building up. In 2012, a specific antibody (FI16) that effectively blocks the cell entry of all the 16 influenza strains currently known has been isolated from a human blood donor. Another such antibody, CR6261, protective against 50% of influenza strains including the "Spanish Flu", was in 2010 successfully tested on ferrets. The distribution of the European polecat, from which ferrets were domesticated, may give an indication where that antibody originated genetically (see Wikipedia map below). I tend to say that we can exclude EEF and steppe populations from the list. No detail is given on the individuals from whom the antibodies were extracted, but both finds were reported by a Dutch research team. Details are included in the following report - maybe someone who is more into the details of genetics than I am can extract further information (responsible SNPs, etc.) from it.
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2596486/#pone.0003942.s001
411px-Mustela_putorius_distribution.svg.png


Now, let's get back to the OT. Lactose tolerance enables adult humans to take up antibodies from other mammals' milk. For influenza, cow milk is not an issue, but reindeer, horse and goat milk could be relevant. Nevertheless, drinking cow milk could enhance protection against other, cow based diseases such as pox and measles.
Moreover, mother milk, and afterwards (to the extent it can be digested) cow milk include the highest concentration of sialic acids found in human food. Sialic acids have multiple functions in the human body, including stimulating brain growth and supporting neural transmission. They are also making up the outer part of cell membranes. Many infective organisms, including influenza viruses, but also Botulinum, Tetanus and Cholera toxins, bond themselves to those sialic acids in order to be able to release the toxine and virus RNA, respectively, into the host cell. Influenza viruses later on release a special enzyme, neuramidase, to cleave that bond in order for the replicated viruses being released and allowed to spread further.
The mechanism is used by a number of popular flu medicaments such as Tamiflu, which essentially flood the body with synthetic sialic acids. This makes it more difficult for influenza viruses to identify cells - instead they bond to the free-floating sialic acids, and can be neutralised by lymphocytes. Moreover, the presence of free-floating sialic acids is inhibiting release of fresh viruses from the cells they are bond to. For its comparatively high sialic acid content, intake of fresh milk has essentially the same effect. One litre of fresh cow milk contains 220-300 mg of sialic acids, the bio-active content of Tamiflu (which is re-synthesized in the liver) is around 60 mg per capsule. The benefit isn't restricted to fresh milk - certain milk products, e.g. paneer, also include substantial amounts of sialic acids, while the content decreases substantially in yoghurt, cheese and especially butter (the sialic acids appear to concentrate in the whey).
Note that not all of the 43 types of sialic acids are beneficiary. One variant, N-Glycolylneuraminic acid (Neu5GC), has been demonstrated to cause immune response and promote cancer cell growth, though most humans appear to have developed antibodies which allow for a limited intake and digestion (150 mg/day on a single occasion, no further tests done due to ethical / health considerations). Neu5GC is especially found in red meat, eggs, and goat milk and cheese, but also present in cow milk (1% of total sialic acid content). Thus, technically, flu protection comparable to one Tamiflu capsule can also be obtained by eating 450g of pork, 750g of beef fat, a kilo of lean beef, or 800g chicken meat per day. Aside from the fact that the first three possibilities should significantly increase cancer risk, consuming a quarter litre of cow milk daily might be the most easily achievable way for EEF to enhance their flu resistance.
http://en.wikipedia.org/wiki/Sialic_acids
http://www.ijabpt.com/pdf/57048-V.M.Biju (1).pdf
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC218710/


Bottom lines & conclusions:
  1. For climatic and nutritional reasons (seal, reindeer), North Eurasian populations are likely to have been earliest and most intensively affected by influenza;
  2. The resulting process of genetic adaptation may among others have promoted lactose tolerance, which substantially increases influenza resistance. Rather than cow-herders, reindeer herders could have been the first group that effectively benefitted from the new genetic trait. Given that reindeer are also affected by small pox, it is probably no coincidence that pox resistance (CCR5-32 allele, see my previous post) has the highest prevalence in Finnish and adjacent Russian populations.
  3. Lactose tolerance became widespread when cattle-based farming reached northern Europe with the Funnelbeaker culture.
  4. Influenza prevalence in already reasonably resistant (pre-) Funnelbeaker populations should have promoted substantial influenza epidemics in less-resistant cultures further south, especially during the epi-Rössen and epi-Lengyel (Schöningen) phase (4,100-3,800 BC) when the contact intensified. These should primarily have been equestrian / maritime strains (H7N7, H3N3, H3N8) yet mostly unknown to central European farmers. The latter may of course have "paid back" with swine flu (H1N1, H3N2), but since the H1N1 strain also occurs with deer, Funnelbeaker people might have been already prepared.
 
Influenza is actually a highly interesting case. It spreads from mammals by aerosols (sneezing), so infection risks from domesticized (living) animals are much higher than from (dead) hunting prey. Mammal viruses can survive outside the body, but on skin at 20°C for only five minutes. Thus, the risk of a south European HG of catching the flu from hunted boar (anyway only the third common prey, far behind Aurochs and deer) is minimal. An Anatolian pig or poultry farmer, OTOH, should have had quite an exposure, especially as poultry feces are also infective and may stay so for several days. Another issue here is that swine is susceptive to both bird and mammal flu, which makes it the perfect vessel for turning one variant into the other one. As such, by the time of spreading to the Central Mediterranean early farmers should already have had quite some infection history.
There is no doubt that catching flu from killed animals is minimal, however it still can occur occasionally from bodily fluids during butchering. Giving thousands of years of hunting wild boar and wild birds and catching occasional flu gave HG some immunity to this pathogen.

For these reasons, flu epidemics mostly occur during late autumn/ winter in moderate to cold climate zones. Vitamin D, production of which is stimulated by sunlight, seems to protect against influenza, and may be another factor for seasonal variation in infection.
Since taking vitamin D supplements I can attest to fewer flu cases and if infected having much milder symptoms.

As such, I agree that north Eurasian HGs most likely had caught the flu, and developed some immunity from it, already before the entrance of early farmers. Climate is one factor here, the other one is the reliance on seal, deer, reindeer and wild horses as hunting prey. Moreover, North Eurasian HGs may have picked up flu viruses from seal-infected brackish water (Baltic Sea!), or from ice (igloo construction etc.). Domestication of wolves/ dogs as hunting or traction animals should also be considered. In fact, north Eurasian HGs' exposition, immunity and transmission potential should have been substantially higher than that of EEFs.
My take on this would be that EEF could have brought intensity and faster mutation of pathogens due to much higher population density and closeness and interaction of various domesticated animals. Although these new strains affected them and surrounding HGs equally, HGs might have not been exposed to all of them (living far away), later paying higher consequences being attacked by completely (for them) new strains, unlike EEF.



  1. For climatic and nutritional reasons (seal, reindeer), North Eurasian populations are likely to have been earliest and most intensively affected by influenza;
  2. The resulting process of genetic adaptation may among others have promoted lactose tolerance, which substantially increases influenza resistance. Rather than cow-herders, reindeer herders could have been the first group that effectively benefitted from the new genetic trait. Given that reindeer are also affected by small pox, it is probably no coincidence that pox resistance (CCR5-32 allele, see my previous post) has the highest prevalence in Finnish and adjacent Russian populations.
Good thinking. Flue could have accelerated faster spreading of lactose persistent gene.

Influenza prevalence in already reasonably resistant (pre-) Funnelbeaker populations should have promoted substantial influenza epidemics in less-resistant cultures further south, especially during the epi-Rössen and epi-Lengyel (Schöningen) phase (4,100-3,800 BC) when the contact intensified. These should primarily have been equestrian / maritime strains (H7N7, H3N3, H3N8) yet mostly unknown to central European farmers. The latter may of course have "paid back" with swine flu (H1N1, H3N2), but since the H1N1 strain also occurs with deer, Funnelbeaker people might have been already prepared.
Interesting thought. EEF and HG would have been attacked by same strains with different intensity according to genetic immunity of both groups. The most important questions would be how often both groups were decimated by really bad cases of flu? Once in generation, once in 100 years or even more rarely?
 
Thanks for your extensive reply, LeBrok. A few comments:
There is no doubt that catching flu from killed animals is minimal, however it still can occur occasionally from bodily fluids during butchering. Giving thousands of years of hunting wild boar and wild birds and catching occasional flu gave HG some immunity to this pathogen.
Direct human infection from birds is rare, though there have been a few cases reported from workers on large chicken farms and chicken slaughterhouses, but none of the European cases spread that flu further to other humans. Conversion from bird to human flu in general needs a mammal vessel. So far, pigs, seals and bats are proven to have acted as such a vessel, dogs at least seem to have the potential, and I personally think more research on rodents, especially squirrels, is required in this respect.

It's not clear whether pig/boar were original hosts, or only became so with the spread of agriculture. Equestrian flu isn't well researched, but the one and only study that I have seen, from Germany in 1980, had 11 out of 20 deer samples showing antibodies to in total 3 different flu strains, one of which later spread to humans (via horses?). HGs catching the flu when butchering deer is of course a possibility, but when the deer is already dead, and outside temperatures are above freezing, the risk is rather minimal. I think the main issue for HGs in warmer climates is caves (cool, dry, no sunlight) and bats (which also aren't well researched for their flu reservoir and transmission potential). And, in fact, cave-dwelling Mesolithic Europeans appear to have been quite resilient to the spread of EEFs (and their diseases?) - Iberia / SW France is a point in case here.

My take on this would be that EEF could have brought intensity and faster mutation of pathogens due to much higher population density and closeness and interaction of various domesticated animals. Although these new strains affected them and surrounding HGs equally, HGs might have not been exposed to all of them (living far away), later paying higher consequences being attacked by completely (for them) new strains, unlike EEF.

EEF and HG would have been attacked by same strains with different intensity according to genetic immunity of both groups. The most important questions would be how often both groups were decimated by really bad cases of flu? Once in generation, once in 100 years or even more rarely?
I will comment on this in more detail I the thread related to the recent Brotherton study on Neolithic population exchange in the Elbe-Saale region, which actually inspired me to look deeper into the possible effect of diseases on the haplogroup composition.
http://www.eupedia.com/forum/thread...-Mid-Neolithic?p=435557&viewfull=1#post435557

However, here my current line of thinking as concerns NHG:
From 5,500 BC on, the Baltic Sea became flooded from the west, around 4,500 BC the former sweetwater lake had turned brackish. Seals immigrated from the North Sea and quickly became the main food source along the SW Baltic coast (25-40 % of all animal bones, second only to deer). Given that seals are potent flu conversion vessels, and flu viruses can survive much longer in brackish than in saltwater, Ertebolle culture HGs should have gained substantial exposition to new strains [Under a "out of Doggerland" scenario, one might assume that Ertebolle populations had already gained substantial seal flu resistance much earlier,.]
Hazelnut was cultivated around Ertebolle settlements, as food, but also as the branches were used for various devices such as fish traps. This should have increased exposition to (flu-bearing) rodents. The same applies to attested rasp- and blackberry cultivation, and flu-bearing bats. Several late Ertebolle settlements on the Danish isles show substantial shares (~30%) of bones of fur-bearing animals (another known source of flu infections), apparently hunted for their pelts. Dogs typically make up for 1-4% of all mammal bones. Dog burials together with their "master" are known from some Ertebolle sites (Skateholm), quite common in Funnelbeaker graves, and the tradition appears to have continued uninterruptedly to early Medieval Saxon graves. As such it is fair to assume that the Ertebolle culture had already domesticized dogs. Genetic analysis of excavated pig bones shows that from 4,700 BC on, the Ertebolle culture did not any more rely exclusively on boar, but had imported domesticated pigs (Anatolian DNA) from the south, and was most likely raising them systematically. In short, latest by the mid-5th milennium BC the Ertebolle culture had assembled an impressive package of substantial exposition to almost all known influenza hosts.
http://www.nature.com/ncomms/2013/130827/ncomms3348/full/ncomms3348.html
http://etheses.whiterose.ac.uk/1712/5/chapter_4.pdf

Flu infections are difficult to trace from the Ertebolle archaeological record, as most sites were only used for a limited period before they became submerged by the rising Baltic Sea. Nevertheless, Scandinavia witnessed at least two periods of volcanic winters around 4,500-4350 BC, and it would be a miracle if those hadn't caused flu epidemics. A rapid cooling sets in in Scandinavia around 4,100 BC, contemporarily with the transition to the Funnelbeaker culture, which is described as quick in the south (Mecklenburg/ Holstein), abrupt in the north (Jutland), but as subtle blend-over in Southern Sweden. Demic expansion is genetically proven, but there is no evidence of violent conflict associated to it. Did a (swine) flu epidemic pave the way for EEF on the Cimbrian peninsula, but not make it across the Öresund?

Aside from swine import, there are other indications of regular contact between the Ertebolle culture and LBK/ Rössen further south, The oldest "Ertebolle"-pottery found (Schlamersdorf near Bad Segeberg) dates to around 5,000 BC, has apparently been produced in the Rhineland, and corresponds date-wise to the earliest imported amber found there. Flint adzes and, less frequently, axes were imported from the Harz and Silesia during the 5th millennium BC, a few copper items (Slovakia) have also been found at Ertebolle sites. There has also been cultural interchange between SW Sweden and the Narva culture in Lithuania and Latvia, e.g. Swedish flint sickles apparently produced for export across the Baltic Sea during the 5th millennium BC. Locally produced Ertebolle ceramic has several parallels to Narva culture ceramic, both in ornamental design and in function (blubber lamps). Moreover, cereal imprints have occasionally (5 times) been found on Ertebolle sherds, interpreted as signs of grain import across the Baltic Sea.
http://what-when-how.com/ancient-eu...urope-transition-to-agriculture-70004000-b-c/

Whether these contacts were intensive enough to disseminate "northern" influenza strains to EEF further south may be debated. Aside from humans, boar may have served as transmission channel. Domesticized pigs were commonly left grazing in open oak forests, and there is genetic evidence of substantial incorporation of European boar DNA into the originally Anatolian swine gene pool. In any case, interaction intensified during the Funnelbeaker period. Around 4,000-3,700 BC, Central Europe experienced a cold and dry phase, during which EEF should at latest have gotten the flu. The Brotherton study documents a massive genetic break around this period in the Elbe-Saale region: mtDNA H subclades of near eastern origin, which dominate the LBK and Rössen samples, get extinct, to be later replaced by subclades that are today mainly West Atlantic (H3, H4) or (south-)east European (H2, H5, H6, H10).
Another study comes to the same result with respect to mtDNA U5 EEF: Outside the basal clade, only one out of nine early Neolithic sub-clades made it to the late Neolithic. Obviously, NHG (Funnelbeaker) U5b wasn't concerned, otherwise they wouldn't have become the dominating mtDNA haplogroup in the Elbe-Saale region during the late third millennium BC Bernburg culture (though U5b may have had other health issues, which subsequently affected its frequency).

Coming back to the OT: The Funnelbeaker itself, and associated ceramic innovations such as the first jugs, point to a drink-based culture. Aside from fresh milk, beer might of course also be considered, but cattle herding appears to have predated systematic cereal production by several centuries. As one paper puts it:
http://www.uni-kiel.de/ufg/bereiche/dateienJMueller/ber_rgk_89_mueller.pdf
A combination of flat-based Gatersleben elements [Elbe-Saale] and funnel-necked Michelsberg types [Rhine-Weser] should have resulted in funnel beakers
Add to this the world's most renowned dairy cow breed, the Holstein-Frisian, and the area where lactose tolerance matured (though not necessarily originated) becomes clear. Holstein Frisians have "only" been bred systematically for some two millennia, but they represent a combination of the original Anatolian/ LBK cattle )mtDNA T3), with Italian / Alpine Aurochs (mtDNA T*) crossed in via Bavarian breeds. Their yDNA is exclusively Anatolian Y1.
http://www.pnas.org/content/103/21/8113.full#T1
F2.large.jpg
 
Thanks for your extensive reply, LeBrok. A few comments:
Lol, if mine is extensive your's has to be called colossal and exhaustive. I wish I had more time to indulge in my hobby these days.
Your posts are always full of knowledge, well written and easy to follow, and I read them with great interest.

Forgive me if I don't elaborate on all points from lack of time.


Dog burials together with their "master" are known from some Ertebolle sites (Skateholm), quite common in Funnelbeaker graves, and the tradition appears to have continued uninterruptedly to early Medieval Saxon graves. As such it is fair to assume that the Ertebolle culture had already domesticized dogs.
According to newest finds on dogs, their domestication can go back as far as 30 thousand years in Europe.
http://www.eupedia.com/forum/threads/29250-Dogs-domesticated-over-20-thousands-years-ago-in-Europe



Whether these contacts were intensive enough to disseminate "northern" influenza strains to EEF further south may be debated. Aside from humans, boar may have served as transmission channel. Domesticized pigs were commonly left grazing in open oak forests, and there is genetic evidence of substantial incorporation of European boar DNA into the originally Anatolian swine gene pool. In any case, interaction intensified during the Funnelbeaker period. Around 4,000-3,700 BC, Central Europe experienced a cold and dry phase, during which EEF should at latest have gotten the flu. The Brotherton study documents a massive genetic break around this period in the Elbe-Saale region: mtDNA H subclades of near eastern origin, which dominate the LBK and Rössen samples, get extinct, to be later replaced by subclades that are today mainly West Atlantic (H3, H4) or (south-)east European (H2, H5, H6, H10).
Another study comes to the same result with respect to mtDNA U5 EEF: Outside the basal clade, only one out of nine early Neolithic sub-clades made it to the late Neolithic. Obviously, NHG (Funnelbeaker) U5b wasn't concerned, otherwise they wouldn't have become the dominating mtDNA haplogroup in the Elbe-Saale region during the late third millennium BC Bernburg culture (though U5b may have had other health issues, which subsequently affected its frequency).
Mitochondria is an energy generator of every cell. Therefore I have a feeling that changeover of MtDNA in the cold phase in Europe went towards MtDNA which could produce more heat in the body. Other words, some MtDNA which came with farmers from Near East didn't survive when European climate turned cold. To forefront came many native ones found among local HGs.
It is just my supposition based on own observation of people around. Some like winter but sweat profusely in summer. Some love summer and freeze terribly in winter. (adjusted for body size) And I think it has a lot to do with mitochondria.

Add to this the world's most renowned dairy cow breed, the Holstein-Frisian, and the area where lactose tolerance matured (though not necessarily originated) becomes clear. Holstein Frisians have "only" been bred systematically for some two millennia, but they represent a combination of the original Anatolian/ LBK cattle )mtDNA T3), with Italian / Alpine Aurochs (mtDNA T*) crossed in via Bavarian breeds. Their yDNA is exclusively Anatolian Y1.
http://www.pnas.org/content/103/21/8113.full#T1
Cool map. Did they say something how they compare autosomally?
 
Cool map. Did they say something how they compare autosomally?
No, they just looked at cattle mtDNA (I took the yDNA info from another page). Nevertheless, their study was interesting enough. It showed, a.o.:
  1. Aside from the British Aurochs, which is a different genus, there have been at least 2 Italian kinds of Aurochs, one possibly with a previously unknown additional T subclade, the other one with an ancient mutation of T3. Further "northern" varieties may show up when bones are screened systematically.
  2. For Italian cattle breeds, some intermixture of Anatolian domesticates with local Aurochs has been proven. For many other European cattle breeds, intermixture is likely. This implies that a lot of the "wild vs. domesticized" analysis of Mesolithic/ Neolithic bone remains needs review. Older genetic screening just recognised the British, but not the South European T3 Aurochs as wild. Phylogenic screening may have mistaken in-bred Aurochs features into domesticates as indication of wild Aurochs, or vice versa. Actually, we could ultimately end up with a somewhat different geographical distinction between EEF and HGs, especially in Italy, Iberia and around the Alps.
  3. North African cattle is almost exclusively T1, with high internal diversity. Independent domestication is possible. In any case, North Africa has a pastoralist and breeding history that is quite different from Europe.
  4. Presence of North African T1 on the north Mediterranean coast indicates trans-Mediterranean cattle shipment (most likely accompanied by human migration). Respective signs are particularly strong in Iberia, and can, from their ancient DNA analysis, be at least traced back to the Bronze Age.
  5. Different ratios of the Anatolian T2/T3 mix should reflect different Neolithisation paths. A maritime path from Anatolia to Greece, central Italy, Southern France and Southern Spain is clearly distinguishable. In addition, there seem to have been at least two continental waves - one "pure T3" one via Thrakia into NW Europe and the Alps, and a mixed one on the western Pontic shore that unfortunately cannot be traced further into Eastern Europe. Both waves appear to have crossed path and intermixed on the middle Danube. T* intermixture may reflect in-breeding of Aurochs, or have been part of the original Neolithic package that was enhanced in some local breeds (note that most Italian Aurochs had T3 and is thus not distinguishable at this hg level).
It is actually a fascinating and promising approach that may potentially tell us a lot about the spread of farming and associated population movements.
 
It is actually a fascinating and promising approach that may potentially tell us a lot about the spread of farming and associated population movements.
Yes, this is amazing. Soon we'll know more about their cattle than they did.
 
Mitochondria is an energy generator of every cell. Therefore I have a feeling that changeover of MtDNA in the cold phase in Europe went towards MtDNA which could produce more heat in the body. Other words, some MtDNA which came with farmers from Near East didn't survive when European climate turned cold. To forefront came many native ones found among local HGs.
It is just my supposition based on own observation of people around. Some like winter but sweat profusely in summer. Some love summer and freeze terribly in winter. (adjusted for body size) And I think it has a lot to do with mitochondria.
That may indeed have been an issue. Digestion efficiency is another one: Farmers in moderate climate zones need to cope with increased seasonal fluctuation in food supply, and there is the occasional crop failure resulting in a hunger winter. Some starve, other survive, depending on how effectively the body can build up reserves and use the few calories that are available [Later on, social status, which is closely linked to body height as proven from bronze-age graves, becomes also important, but that might be more of an yDNA issue].

The role of Mitochondria even goes far beyond. Essentially, it controls which substances are allowed to enter which cells, and that covers nutrients as well as pathogens. So, the whole immune system is strongly shaped by mtDNA. For example, recent research has yielded that susceptibility to HIV/AIDS, but also diabetes risk, varies considerably between mtDNA haplogroups. Haplogroup U, and especially U5, e.g., has been found to bear greater risk of diabetes, liver cirrhosis and capturing liver-related diseases like hepatitis. While detailed mechanisms aren't understood yet, this is all linked to digestion of starch, sugar and lipids. Considering that U5 is the classical European HG haplogroup, it seems plausible that digesting starch-based food(cereals) is working less well with them than with typical EEF HGs like, e.g., H / HV (low risk of cardio-vascular and liver diseases). mtDNA X, another EEF hg, has in some studies found to be associated with high body-mass and body-fat indexes - historically a good trait to survive food shortages, though maybe less desirable today.
http://www.aidsreviews.com/files/2013_15_4_213-220.pdf (I don't understand the technical details either, but a look at the headlines is nevertheless informative)

There are clearly trade-offs between the advantages and disadvantages of specific immunological traits:
http://www.pnas.org/content/104/42/16645.full
Infectious diseases exert a constant evolutionary pressure on the genetic makeup of our innate immune system. Polymorphisms in Toll-like receptor 4 (TLR4) have been related to susceptibility to Gram-negative infections and septic shock. Here we show that two polymorphisms of TLR4, Asp299Gly and Thr399Ile, have unique distributions in populations from Africa, Asia, and Europe. Genetic and functional studies are compatible with a model in which the nonsynonymous polymorphism Asp299Gly has evolved as a protective allele against malaria, explaining its high prevalence in subSaharan Africa. However, the same allele could have been disadvantageous after migration of modern humans into Eurasia, putatively because of increased susceptibility to severe bacterial infections. In contrast, the Asp299Gly allele, when present in cosegregation with Thr399Ile to form the Asp299Gly/Thr399Ile haplotype, shows selective neutrality. Polymorphisms in TLR4 exemplify how the interaction between our innate immune system and the infectious pressures in particular environments may have shaped the genetic variations and function of our immune system during the out-of-Africa migration of modern humans.

I have in a previous post mentioned the CCR5-Δ32 allele, which is highly protective against AIDS/HIV and (cattle-related) smallpox. I have come across a paper that analyses its current distribution:
http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.0030339
originalimage

They first tried to explain the distribution by homogenous spreading from a Ferroscandian origin. While such model works technically, it yields a far too young age of the mutation (700-3500 years) which in the meantime has been identified to have been present with Swedish HGs 7,000 years ago.
Alternatively, they established a model of strong selective pressure working in favour as well as against the mutation. The underlying rationale was that that the CCR5-Δ32 allele blocks mechanisms that are not only used by smallpox and HIV viruses, but also by the human immune system. "Studies with mice show that the CCR5 knock-outs have poor immune response to various pathogene infections". Among others, CCR5 has been found to play a strong role in the immune defence against tetanus and similar bacterial wound infections. They thus assumed that CCR5-Δ32 may actually become disadvantageous in hot climates that promote bacterial infections. A second simulation that included a north-westerly selection gradient in favour of the mutation identified Spain and the eastern Caspian Sea as most likely origins of the mutation. Given that it was yet unknown at the time of their analysis that CCR5-Δ32 was present 7,000 years ago in Sweden, and that their sampling and subsequent modelling spared out Britain, this will probably not be the final word on this mutation.
Still, the study illustrates that the value of specific immunologic traits may differ considerably according to climate and related infection risks. Population movement into other climate zones, and/or climate change, may turn a selection advantage into a disadvantage. To the extent those traits are tied to mtDNA haplogroups, we should expect resulting selection processes being reflected in an increase or decrease of the share of the haplogroups in question. Not all, possibly not even the majority of changes in the mtDNA structure are linked to demic processes.
 
Here is another issue that has crossed my mind: A major problem of early farmers (and not only them, it also has been diagnosed among bronze-age populations) was scurvy, i.e. undersupply with vitamin C. Vitamin C content of meat or fish typically ranges around 20mg/kg, which is insufficient to cover the daily demand of 75 mg. Cereals don't contain vitamin C at all. Most vitamin C sources used today (citrus, tomato, red pepper/paprika, potatoes) aren't native to Europe.
In late spring and summer, EEF could cover their vitamin C demand from fruit and berries, especially black currant, elderberry, sea-buckthorn, or rose-hip, provided they were cultivated nearby. But most berries are quickly perishable and don't last until the winter. That leaves mainly apples, cabbage, turnip and beet, possibly also sloe, as vitamin C sources during winter. Modern apples have been bred from Caucasian and Central Asian varieties, and could already have been part of the "Neolithic package". The northern border of their cultivation is approximately the same as for wheat. Cabbage grows wildly in maritime Europe up to at least Helgoland, sea-beet even up to the southern Norwegian coast, and the Swedish turnip occurred wildly in - yes - Sweden. So far, so good - Vitamin C supply shouldn't have been much of a problem for HGs and EEF up to around 60° northern latitude, provided they were aware of their respective demand.
With cattle nomads, be it in northern Scandinavia, or in the Eastern European steppes, the situation is different. Here, lactose tolerance comes in handy. Fresh cow milk contains some 10mg of vitamin C per litre, which isn't enough to satisfy the vitamin C demand during winter, but better than nothing, especially if there is some preserved lingonberry available as additional vitamin C source. Much better is sheep milk (42 mg vitamin C per litre); horse milk even contains 130 mg/ litre! The vitamin C content decreases by some 25% in yoghurt and by much more ih cheese, so being able to consume fresh milk can be quite an advantage for pastoralist nomads and in northern climate zones.
 
Here is another issue that has crossed my mind: A major problem of early farmers (and not only them, it also has been diagnosed among bronze-age populations) was scurvy, i.e. undersupply with vitamin C. Vitamin C content of meat or fish typically ranges around 20mg/kg, which is insufficient to cover the daily demand of 75 mg. Cereals don't contain vitamin C at all. Most vitamin C sources used today (citrus, tomato, red pepper/paprika, potatoes) aren't native to Europe.
In late spring and summer, EEF could cover their vitamin C demand from fruit and berries, especially black currant, elderberry, sea-buckthorn, or rose-hip, provided they were cultivated nearby. But most berries are quickly perishable and don't last until the winter. That leaves mainly apples, cabbage, turnip and beet, possibly also sloe, as vitamin C sources during winter. Modern apples have been bred from Caucasian and Central Asian varieties, and could already have been part of the "Neolithic package". The northern border of their cultivation is approximately the same as for wheat. Cabbage grows wildly in maritime Europe up to at least Helgoland, sea-beet even up to the southern Norwegian coast, and the Swedish turnip occurred wildly in - yes - Sweden. So far, so good - Vitamin C supply shouldn't have been much of a problem for HGs and EEF up to around 60° northern latitude, provided they were aware of their respective demand.
With cattle nomads, be it in northern Scandinavia, or in the Eastern European steppes, the situation is different. Here, lactose tolerance comes in handy. Fresh cow milk contains some 10mg of vitamin C per litre, which isn't enough to satisfy the vitamin C demand during winter, but better than nothing, especially if there is some preserved lingonberry available as additional vitamin C source. Much better is sheep milk (42 mg vitamin C per litre); horse milk even contains 130 mg/ litre! The vitamin C content decreases by some 25% in yoghurt and by much more ih cheese, so being able to consume fresh milk can be quite an advantage for pastoralist nomads and in northern climate zones.
Great point, another plus for explanation of lactose persistence in northern latitudes. Even if crops fail and not many veggies to eat, there is always grass to feed cows to get fresh milk, which includes fats, sugar, proteins and many vitamins. The best miracle comes in winter when is cold and nothing grows to eat. Feed cows dry grass and you will still receive 10 or 20 liters of milk a day.

If it comes to vitamin C there is a huge tradition in central and north Europe to pickle cabbage and cucumbers. I never checked vitamin C value in them, but I have a strong feeling that it was/is the main purpose behind pickling them.
 
If it comes to vitamin C there is a huge tradition in central and north Europe to pickle cabbage and cucumbers. I never checked vitamin C value in them, but I have a strong feeling that it was/is the main purpose behind pickling them.
The problem with vitamin C appears to be that it is water soluble and doesn't stand most traditional conservation techniques: Heat it, and its gone (no jams), let it ferment, and its gone (no fruit wine), expose it to vinegar, and its gone (no pickles). There seem to be two exceptions, however: One is Sauerkraut - cabbage not pickled in vinegar, but salted, which makes it ferment in lactic acid. This destroys some of the vitamin C, but preserves part of it. 100g Sauerkraut contain 25mg vitamin C, compared to 40-45 mg in fresh cabbage. However, to stand the lactic acid included in Sauerkraut, you better be lactose tolerant, otherwise it "speeds up" your digestion. So, I guess a Sauerkraut consumption map is also not too bad as indicator of lactase persistence.

The other way to preserve part of the Vitamin C is conserving it in pure alcohol, i.e. preparing liqueurs. For their high vitamin C content (but also as they are quite tasty), black current, sloe and elderberry would be the first choice here. The good old fashioned North German / Danish Rumtopf should work as well. Unfortunately, it seems that distilling technology only entered Europe during the Medieval. It may be that a reasonably strong alcoholic beverage, e.g. strong mead, also does the trick. Beer apparently isn't sufficient, in Belgian Kriek (cherry beer), vitamin C content is reduced to zero.
390px-Rumtopf.jpg


But, when it is sufficiently cold outside, the best is probably to store a lot of apples, cabbage and beet underground and regularly add it to your winter diet. And I believe that is what most of Northern and Eastern Europeans have traditionally been doing...

P.S: I have come across another Vitamin C source for the winter: Horseradish - whopping 175mg vitamin C /100g. Eastern Europe, Scandinavia & Baltic Sea, again (plus Bavaria, plus Japan)!
 
The problem with vitamin C appears to be that it is water soluble and doesn't stand most traditional conservation techniques: Heat it, and its gone (no jams), let it ferment, and its gone (no fruit wine), expose it to vinegar, and its gone (no pickles). There seem to be two exceptions, however: One is Sauerkraut - cabbage not pickled in vinegar, but salted, which makes it ferment in lactic acid. This destroys some of the vitamin C, but preserves part of it. 100g Sauerkraut contain 25mg vitamin C, compared to 40-45 mg in fresh cabbage. However, to stand the lactic acid included in Sauerkraut, you better be lactose tolerant, otherwise it "speeds up" your digestion. So, I guess a Sauerkraut consumption map is also not too bad as indicator of lactase persistence.

The other way to preserve part of the Vitamin C is conserving it in pure alcohol, i.e. preparing liqueurs. For their high vitamin C content (but also as they are quite tasty), black current, sloe and elderberry would be the first choice here. The good old fashioned North German / Danish Rumtopf should work as well. Unfortunately, it seems that distilling technology only entered Europe during the Medieval. It may be that a reasonably strong alcoholic beverage, e.g. strong mead, also does the trick. Beer apparently isn't sufficient, in Belgian Kriek (cherry beer), vitamin C content is reduced to zero.


But, when it is sufficiently cold outside, the best is probably to store a lot of apples, cabbage and beet underground and regularly add it to your winter diet. And I believe that is what most of Northern and Eastern Europeans have traditionally been doing...

P.S: I have come across another Vitamin C source for the winter: Horseradish - whopping 175mg vitamin C /100g. Eastern Europe, Scandinavia & Baltic Sea, again (plus Bavaria, plus Japan)!
This is a great info. Yes I meant pickling with sult in a barrel, no vinegar, the traditional way of Northern and Eastern Europeans. I have a great appetite for anything made of cabbage, love pickles and horseradish and ordinary radish too. I'm not a fan of vinegar pickles and can't stand wine vinegar. Generally I like fresh organic veggies and salads, but I can vouch for many men from this part of Europe, they won't touch fresh veggies with exception of sauerkraut and pickles.
 
The problem with vitamin C appears to be that it is water soluble and doesn't stand most traditional conservation techniques: Heat it, and its gone (no jams), let it ferment, and its gone (no fruit wine), expose it to vinegar, and its gone (no pickles). There seem to be two exceptions, however: One is Sauerkraut - cabbage not pickled in vinegar, but salted, which makes it ferment in lactic acid. This destroys some of the vitamin C, but preserves part of it. 100g Sauerkraut contain 25mg vitamin C, compared to 40-45 mg in fresh cabbage. However, to stand the lactic acid included in Sauerkraut, you better be lactose tolerant, otherwise it "speeds up" your digestion. So, I guess a Sauerkraut consumption map is also not too bad as indicator of lactase persistence.

The other way to preserve part of the Vitamin C is conserving it in pure alcohol, i.e. preparing liqueurs. For their high vitamin C content (but also as they are quite tasty), black current, sloe and elderberry would be the first choice here. The good old fashioned North German / Danish Rumtopf should work as well. Unfortunately, it seems that distilling technology only entered Europe during the Medieval. It may be that a reasonably strong alcoholic beverage, e.g. strong mead, also does the trick. Beer apparently isn't sufficient, in Belgian Kriek (cherry beer), vitamin C content is reduced to zero.
390px-Rumtopf.jpg


But, when it is sufficiently cold outside, the best is probably to store a lot of apples, cabbage and beet underground and regularly add it to your winter diet. And I believe that is what most of Northern and Eastern Europeans have traditionally been doing...

P.S: I have come across another Vitamin C source for the winter: Horseradish - whopping 175mg vitamin C /100g. Eastern Europe, Scandinavia & Baltic Sea, again (plus Bavaria, plus Japan)!

When I was growing up in the country, we lived in an old stone farmhouse with one wood stove for heat, and the cellar was of course unheated. We could store cabbages, potatoes and apples down there and they would last for most of the winter. Carrots spoiled more easily, so we would bury them outside in layers of straw, and dig them up a bit at a time. And some fruit, vegetables and meat were stored in the basement in glass jars sealed with wax - probably the same could have been done with ceramic jars. And although we didn't smoke meat in order to store it for the winter, some of our neighbours did. If you live in a cold climate with low technology, you need to be clever about food storage for the winter season, but the cold weather helps to make it easier. Of course, all those old ways are no doubt forgotten, since farmers are much more prosperous now and all have refrigerators, freezers and cars that make it easy to drive into town for shopping.
 
When I was growing up in the country, we lived in an old stone farmhouse with one wood stove for heat, and the cellar was of course unheated. We could store cabbages, potatoes and apples down there and they would last for most of the winter. Carrots spoiled more easily, so we would bury them outside in layers of straw, and dig them up a bit at a time. And some fruit, vegetables and meat were stored in the basement in glass jars sealed with wax - probably the same could have been done with ceramic jars. And although we didn't smoke meat in order to store it for the winter, some of our neighbours did. If you live in a cold climate with low technology, you need to be clever about food storage for the winter season, but the cold weather helps to make it easier. Of course, all those old ways are no doubt forgotten, since farmers are much more prosperous now and all have refrigerators, freezers and cars that make it easy to drive into town for shopping.
In east European villages there was storage in the ground by every house. Small mound of earth with a wooden door . Poor village houses didn't have basements/cellars. Anyway, roughly same idea for cold storage.
 
In east European villages there was storage in the ground by every house. Small mound of earth with a wooden door . Poor village houses didn't have basements/cellars. Anyway, roughly same idea for cold storage.
Underground storage pits ere typical for LBK settlements/ houses, and have also been found in many middle and late Neolithic settlements. And the Hallstatt peolpe are known to have stored their meat directly in the salt mine, which had three advantages - they were cool, smoky (from all the fatwood burning), and the air was of course salt-filled. Quite clever!

I have tried to find out whether pickling was already common practice during the Neolithics. Many claim so, but there is little evidence for Northern Europe (though olive pickling around the Mediterranean appears to be quite well documented). The earliest evidence I could find is from Norway from the mid 3rd millennium BC - fishermen had taken pots with a stew/porridge of pickled meat (probably intestines) and unidentified starch sources (nuts?, beets?, cabbage?) with them to their summer fishing, seal- and beaver-hunting camp on the Oslo fjord.
http://www.archaeology.su.se/polopo.../standard/file/LA9.Ostmo.Hulthen.Isaksson.pdf

The problem appears to be that lactic acid fermentation, plus subsequent heating (plus a few millennia of depositing in the soil) destroys the cell structure of vegetables to such an extent that their original nature cannot be established anymore. Anyway, I looked around for Neolithic pottery that resembles 20th century pottery for household sauerkraut preparation:
514223B01.JPG


This is what I have found - Austrian LBK:
A.A_005_mit.jpg

Definitely not a cooking pot, and rather impractical for storing grain or serving drinking water. In order to be useable for pickling, a container should have low porosity to prevent brine evaporation over time. Whether Central European early Neolithic pottery already was sufficiently water-tight is unclear, but at latest Funnelbeaker pottery (i.e. early Chalcolithic) is documented to have included such vessels.
 
In east European villages there was storage in the ground by every house. Small mound of earth with a wooden door . Poor village houses didn't have basements/cellars. Anyway, roughly same idea for cold storage.

Some of the farms in the area where I grew up had such storage sheds, usually in addition to cellars - these storage sheds were called root cellars because were mostly used to store root vegetables, such as potatoes and turnips. I forgot about that because we didn't need one - we had a large cellar and it stayed quite cold all winter.
 
As far as pots are concerned, it's probably difficult to overestimate just how important glazed pottery would have been for survival in a cold or even a temperate climate, anywhere far enough north that crops are seasonal. Without the ability to store food for long periods, survival must have been difficult. Glazed pottery would probably have given late Neolithic and Copper Age people a big advantage as compared to early Neolithic people who only had unglazed pottery.
 
This is what I have found - Austrian LBK:
A.A_005_mit.jpg

Definitely not a cooking pot, and rather impractical for storing grain or serving drinking water. In order to be useable for pickling, a container should have low porosity to prevent brine evaporation over time. Whether Central European early Neolithic pottery already was sufficiently water-tight is unclear, but at latest Funnelbeaker pottery (i.e. early Chalcolithic) is documented to have included such vessels.

That's an interesting vessel. I wonder what are the holes at the bottom for? Some sort of carrying ropes for transportation, for a neat hanging contraption, or perhaps to hang it upside down for drying after a wash?
I don't think sealing porous clay containers was a big problem, in absence of glaze. One can moist inside walls with grease or olive oil, any edible hydrophobic substance.

Going back to this vessel above. It might be actually for carrying water from the well or river. The bottom holes are needed for stabilization of the vessel, so it doesn't swing while being carried. It doesn't mean that it couldn't have had a double, or many uses. It might have been the closest thing related to a jar they've used.
 
Mongols are very interesting example of high dairy diet, and surprisingly only being around 30% lactose persistent. (if all the maps are correct).

This is from the trip of Lieutenant-Colonel Nikolai Mikhailovich Przhevalsky in 1870:


Looks like they used to have diet only made of dairy and meat. Makes you think how they manage doing it without being lactose tolerant. Granted they consume most dairy as cheeses and butter, but in quantities they do one would think they should be lactose tolerant.
One explanation is that they must have adequate bacterial flora in their digestive organs to help them deal with all the lactose, or we are missing one more gene or allele taking part in lactose digestion.

If it is about bacterial flora, it would be a good news for people who are not LP but like eating dairy. Start with small quantities of natural products containing good bacteria. With time build up consumption and your good bacterial flora will grow together adequately.

One thought from the Colonel's quote:

"When it comes to "white foods" (anything made from milk), almost everything is heated due to the brucellosis problem within the country."

http://en.wikipedia.org/wiki/Brucellosis


Needing to always heat milk would limit the benefit of LP so I wonder if that had something to do with it. The lower the risk of Brucellosis (for whatever reason) the more people might drink milk raw and the bigger the benefit from LP?

 
Fascinating posts on this thread re mtdna, LP and connection with disease resistance. Ty all.

"I have a great appetite for anything made of cabbage, love pickles and horseradish and ordinary radish too. I'm not a fan of vinegar pickles and can't stand wine vinegar. Generally I like fresh organic veggies and salads, but I can vouch for many men from this part of Europe, they won't touch fresh veggies with exception of sauerkraut and pickles."

I wonder if a liking for certain foods necessary to survival in a particular region has been selected for in people from that region?
 
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