Dairying, diseases and the evolution of lactase persistence in Europe.

In European and many African, Middle Eastern and southern Asian populations, lactase persistence (LP) is the most strongly selected monogenic trait to have evolved over the past 10,000 years1. Although the selection of LP and the consumption of prehistoric milk must be linked, considerable uncertainty remains concerning their spatiotemporal configuration and specific interactions2,3. Here we provide detailed distributions of milk exploitation across Europe over the past 9,000 years using around 7,000 pottery fat residues from more than 550 archaeological sites. European milk use was widespread from the Neolithic period onwards but varied spatially and temporally in intensity. Notably, LP selection varying with levels of prehistoric milk exploitation is no better at explaining LP allele frequency trajectories than uniform selection since the Neolithic period. In the UK Biobank4,5 cohort of 500,000 contemporary Europeans, LP genotype was only weakly associated with milk consumption and did not show consistent associations with improved fitness or health indicators. This suggests that other reasons for the beneficial effects of LP should be considered for its rapid frequency increase. We propose that lactase non-persistent individuals consumed milk when it became available but, under conditions of famine and/or increased pathogen exposure, this was disadvantageous, driving LP selection in prehistoric Europe. Comparison of model likelihoods indicates that population fluctuations, settlement density and wild animal exploitation-proxies for these drivers-provide better explanations of LP selection than the extent of milk exploitation. These findings offer new perspectives on prehistoric milk exploitation and LP evolution.

Ancient Europeans farmed dairy-but couldn't digest milk.

Giant study of ancient pottery and DNA challenges common evolutionary explanation for lactase persistence.

Dairying enabled Early Bronze Age Yamnaya steppe expansions.

During the Early Bronze Age, populations of the western Eurasian steppe expanded across an immense area of northern Eurasia. Combined archaeological and genetic evidence supports widespread Early Bronze Age population movements out of the Pontic-Caspian steppe that resulted in gene flow across vast distances, linking populations of Yamnaya pastoralists in Scandinavia with pastoral populations (known as the Afanasievo) far to the east in the Altai Mountains1,2 and Mongolia3. Although some models hold that this expansion was the outcome of a newly mobile pastoral economy characterized by horse traction, bulk wagon transport4-6 and regular dietary dependence on meat and milk5, hard evidence for these economic features has not been found. Here we draw on proteomic analysis of dental calculus from individuals from the western Eurasian steppe to demonstrate a major transition in dairying at the start of the Bronze Age. The rapid onset of ubiquitous dairying at a point in time when steppe populations are known to have begun dispersing offers critical insight into a key catalyst of steppe mobility. The identification of horse milk proteins also indicates horse domestication by the Early Bronze Age, which provides support for its role in steppe dispersals. Our results point to a potential epicentre for horse domestication in the Pontic-Caspian steppe by the third millennium BC, and offer strong support for the notion that the novel exploitation of secondary animal products was a key driver of the expansions of Eurasian steppe pastoralists by the Early Bronze Age.

Latitudinal gradient in dairy production with the introduction of farming in Atlantic Europe.

The introduction of farming had far-reaching impacts on health, social structure and demography. Although the spread of domesticated plants and animals has been extensively tracked, it is unclear how these nascent economies developed within different environmental and cultural settings. Using molecular and isotopic analysis of lipids from pottery, here we investigate the foods prepared by the earliest farming communities of the European Atlantic seaboard. Surprisingly, we find an absence of aquatic foods, including in ceramics from coastal sites, except in the Western Baltic where this tradition continued from indigenous ceramic using hunter-gatherer-fishers. The frequency of dairy products in pottery increased as farming was progressively introduced along a northerly latitudinal gradient. This finding implies that early farming communities needed time to adapt their economic practices before expanding into more northerly areas. Latitudinal differences in the scale of dairy production might also have influenced the evolution of adult lactase persistence across Europe.

Dairy pastoralism sustained eastern Eurasian steppe populations for 5,000 years.

Dairy pastoralism is integral to contemporary and past lifeways on the eastern Eurasian steppe, facilitating survival in agriculturally challenging environments. While previous research has indicated that ruminant dairy pastoralism was practiced in the region by circa 1300 BC, the origin, extent and diversity of this custom remain poorly understood. Here, we analyse ancient proteins from human dental calculus recovered from geographically diverse locations across Mongolia and spanning 5,000 years. We present the earliest evidence for dairy consumption on the eastern Eurasian steppe by circa 3000 BC and the later emergence of horse milking at circa 1200 BC, concurrent with the first evidence for horse riding. We argue that ruminant dairying contributed to the demographic success of Bronze Age Mongolian populations and that the origins of traditional horse dairy products in eastern Eurasia are closely tied to the regional emergence of mounted herding societies during the late second millennium BC.

Contrasting patterns of prehistoric human diet and subsistence in northernmost Europe.

Current archaeological evidence indicates the transition from hunting-fishing-gathering to agriculture in Northern Europe was a gradual process. This transition was especially complex in the prehistoric North Fennoscandian landscape where the high latitude posed a challenge to both domestic animal breeding and cereal cultivation. The conditions varied, the coastal dwellers had access to rich marine resources and enjoyed a milder climate due to the Gulf Stream, while those living in the inland Boreal forest zone faced longer and colder winters and less diversity in animal and plant resources. Thus, the coastal area provided more favourable conditions for early agriculture compared to those found inland. Interestingly, a cultural differentiation between these areas is archaeologically visible from the late 2nd millennium BC onwards. This is most clearly seen in regionally distinct pottery styles, offering unique opportunities to probe diet and subsistence through the organic residues preserved in ceramic vessels. Herein, we integrate the lipid biomarker, compound-specific stable carbon isotopes (δ13C), and zooarchaeological evidence to reveal culturally distinct human diets and subsistence patterns. In northern Norway, some of the coastal people adopted dairying as part of their subsistence strategy, while the inhabitants of the interior, in common with northern Finland, continued their hunter-gatherer-fisher lifestyles.

A 100-Year Review: From ascorbic acid to zinc-Mineral and vitamin nutrition of dairy cows.

Mineral and vitamin nutrition of dairy cows was studied before the first volume of the Journal of Dairy Science was published and is still actively researched today. The initial studies on mineral nutrition of dairy cows were simple balance experiments (although the methods available at the time for measuring minerals were anything but simple). Output of Ca and P in feces, urine, and milk was subtracted from intake of Ca and P, and if values were negative it was often assumed that cows were lacking in the particular mineral. As analytical methods improved, more minerals were found to be required by dairy cows, and blood and tissue concentrations became primary response variables. Those measures often were poorly related to cow health, leading to the use of disease prevalence and immune function as a measure of mineral adequacy. As data were generated, mineral requirements became more accurate and included more sources of variation. In addition to milk yield and body weight inputs, bioavailability coefficients of minerals from different sources are used to formulate diets that can meet the needs of the cow without excessive excretion of minerals in manure, which negatively affects the environment. Milk, or more accurately the lack of milk in human diets, was central to the discovery of vitamins, but research into vitamin nutrition of cows developed slowly. For many decades bioassays were the only available method for measuring vitamin concentrations, which greatly limited research. The history of vitamin nutrition mirrors that of mineral nutrition. Among the first experiments conducted on vitamin nutrition of cows were those examining the factors affecting vitamin concentrations of milk. This was followed by determining the amount of vitamins needed to prevent deficiency diseases, which evolved into research to determine the amount of vitamins required to promote overall good health. The majority of research was conducted on vitamins A, D, and E because these vitamins have a dietary requirement, and clinical and marginal deficiencies became common as diets for cows changed from pasture and full exposure to the sun to stored forage and limited sun exposure. As researchers learned new functions of fat-soluble vitamins, requirements generally increased over time. Diets generally contain substantial amounts of B vitamins, and rumen bacteria can synthesize large quantities of many B vitamins; hence, research on water-soluble vitamins lagged behind. We now know that supplementation of specific water-soluble vitamins can enhance cow health and increase milk production in certain situations. Additional research is needed to define specific requirements for many water-soluble vitamins. Both mineral and vitamin research is hampered by the lack of sensitive biomarkers of status, but advanced molecular techniques may provide measures that respond to altered supply of minerals and vitamins and that are related to health or productive responses of the cow. The overall importance of proper mineral and vitamin nutrition is known, but as we discover new and more diverse functions, better supplementation strategies should lead to even better cow health and higher production.

A 100-Year Review: Fat feeding of dairy cows.

Over 100 years, the Journal of Dairy Science has recorded incredible changes in the utilization of fat for dairy cattle. Fat has progressed from nothing more than a contaminant in some protein supplements to a valuable high-energy substitute for cereal grains, a valuable energy source in its own right, and a modifier of cellular metabolism that is under active investigation in the 21st century. Milestones in the use of fats for dairy cattle from 1917 to 2017 result from the combined efforts of noted scientists and industry personnel worldwide, with much of the research published in Journal of Dairy Science. We are humbled to have been asked to contribute to this historical collection of significant developments in fat research over the past 100 years. Our goal is not to detail all the work published as each development moved forward; rather, it is to point out when publication marked a significant change in thinking regarding use of fat supplements. This approach forced omission of critically important names and publications in many journals as ideas moved forward. However, we hope that a description of the major changes in fat feeding during the past 100 years will stimulate reflection on progress in fat research and encourage further perusal of details of significant events.

A 100-Year Review: Protein and amino acid nutrition in dairy cows.

Considerable progress has been made in understanding the protein and amino acid (AA) nutrition of dairy cows. The chemistry of feed crude protein (CP) appears to be well understood, as is the mechanism of ruminal protein degradation by rumen bacteria and protozoa. It has been shown that ammonia released from AA degradation in the rumen is used for bacterial protein formation and that urea can be a useful N supplement when lower protein diets are fed. It is now well documented that adequate rumen ammonia levels must be maintained for maximal synthesis of microbial protein and that a deficiency of rumen-degradable protein can decrease microbial protein synthesis, fiber digestibility, and feed intake. Rumen-synthesized microbial protein accounts for most of the CP flowing to the small intestine and is considered a high-quality protein for dairy cows because of apparent high digestibility and good AA composition. Much attention has been given to evaluating different methods to quantify ruminal protein degradation and escape and for measuring ruminal outflows of microbial protein and rumen-undegraded feed protein. The methods and accompanying results are used to determine the nutritional value of protein supplements and to develop nutritional models and evaluate their predictive ability. Lysine, methionine, and histidine have been identified most often as the most-limiting amino acids, with rumen-protected forms of lysine and methionine available for ration supplementation. Guidelines for protein feeding have evolved from simple feeding standards for dietary CP to more complex nutrition models that are designed to predict supplies and requirements for rumen ammonia and peptides and intestinally absorbable AA. The industry awaits more robust and mechanistic models for predicting supplies and requirements of rumen-available N and absorbed AA. Such models will be useful in allowing for feeding lower protein diets and increased efficiency of microbial protein synthesis.

High-resolution isotopic evidence of specialised cattle herding in the European Neolithic.

Reconstructing stock herding strategies and land use is key to comprehending past human social organization and economy. We present laser-ablation strontium and carbon isotope data from 25 cattle (Bos taurus) to reconstruct mobility and infer herding management at the Swiss lakeside settlement of Arbon Bleiche 3, occupied for only 15 years (3384-3370 BC). Our results reveal three distinct isotopic patterns that likely reflect different herding strategies: 1) localized cattle herding, 2) seasonal movement, and 3) herding away from the site year-round. Different strategies of herding are not uniformly represented in various areas of the settlement, which indicates specialist modes of cattle management. The pressure on local fodder capacities and the need for alternative herding regimes must have involved diverse access to grazing resources. Consequently, the increasing importance of cattle in the local landscape was likely to have contributed to the progress of socio-economic differentiation in early agricultural societies in Europe.

Regional asynchronicity in dairy production and processing in early farming communities of the northern Mediterranean.

In the absence of any direct evidence, the relative importance of meat and dairy productions to Neolithic prehistoric Mediterranean communities has been extensively debated. Here, we combine lipid residue analysis of ceramic vessels with osteo-archaeological age-at-death analysis from 82 northern Mediterranean and Near Eastern sites dating from the seventh to fifth millennia BC to address this question. The findings show variable intensities in dairy and nondairy activities in the Mediterranean region with the slaughter profiles of domesticated ruminants mirroring the results of the organic residue analyses. The finding of milk residues in very early Neolithic pottery (seventh millennium BC) from both the east and west of the region contrasts with much lower intensities in sites of northern Greece, where pig bones are present in higher frequencies compared with other locations. In this region, the slaughter profiles of all domesticated ruminants suggest meat production predominated. Overall, it appears that milk or the by-products of milk was an important foodstuff, which may have contributed significantly to the spread of these cultural groups by providing a nourishing and sustainable product for early farming communities.

Cattle Management for Dairying in Scandinavia's Earliest Neolithic.

New evidence for cattle husbandry practices during the earliest period of the southern Scandinavian Neolithic indicates multiple birth seasons and dairying from its start. Sequential sampling of tooth enamel carbonate carbon and oxygen isotope ratio analyses and strontium isotopic provenancing indicate more than one season of birth in locally reared cattle at the earliest Neolithic Funnel Beaker (EN I TRB, 3950-3500 cal. B.C.) site of Almhov in Scania, Sweden. The main purpose for which cattle are manipulated to give birth in more than one season is to prolong lactation for the production of milk and dairy-based products. As this is a difficult, intensive, and time-consuming strategy, these data demonstrate complex farming practices by early Neolithic farmers. This result offers strong support for immigration-based explanations of agricultural origins in southern Scandinavia on the grounds that such a specialised skill set cannot represent the piecemeal incorporation of agricultural techniques into an existing hunter-gatherer-fisher economy.

Earliest evidence for cheese making in the sixth millennium BC in northern Europe.

The introduction of dairying was a critical step in early agriculture, with milk products being rapidly adopted as a major component of the diets of prehistoric farmers and pottery-using late hunter-gatherers. The processing of milk, particularly the production of cheese, would have been a critical development because it not only allowed the preservation of milk products in a non-perishable and transportable form, but also it made milk a more digestible commodity for early prehistoric farmers. The finding of abundant milk residues in pottery vessels from seventh millennium sites from north-western Anatolia provided the earliest evidence of milk processing, although the exact practice could not be explicitly defined. Notably, the discovery of potsherds pierced with small holes appear at early Neolithic sites in temperate Europe in the sixth millennium BC and have been interpreted typologically as 'cheese-strainers', although a direct association with milk processing has not yet been demonstrated. Organic residues preserved in pottery vessels have provided direct evidence for early milk use in the Neolithic period in the Near East and south-eastern Europe, north Africa, Denmark and the British Isles, based on the δ(13)C and Δ(13)C values of the major fatty acids in milk. Here we apply the same approach to investigate the function of sieves/strainer vessels, providing direct chemical evidence for their use in milk processing. The presence of abundant milk fat in these specialized vessels, comparable in form to modern cheese strainers, provides compelling evidence for the vessels having being used to separate fat-rich milk curds from the lactose-containing whey. This new evidence emphasizes the importance of pottery vessels in processing dairy products, particularly in the manufacture of reduced-lactose milk products among lactose-intolerant prehistoric farming communities.

Earliest date for milk use in the Near East and southeastern Europe linked to cattle herding.

The domestication of cattle, sheep and goats had already taken place in the Near East by the eighth millennium bc. Although there would have been considerable economic and nutritional gains from using these animals for their milk and other products from living animals-that is, traction and wool-the first clear evidence for these appears much later, from the late fifth and fourth millennia bc. Hence, the timing and region in which milking was first practised remain unknown. Organic residues preserved in archaeological pottery have provided direct evidence for the use of milk in the fourth millennium in Britain, and in the sixth millennium in eastern Europe, based on the delta(13)C values of the major fatty acids of milk fat. Here we apply this approach to more than 2,200 pottery vessels from sites in the Near East and southeastern Europe dating from the fifth to the seventh millennia bc. We show that milk was in use by the seventh millennium; this is the earliest direct evidence to date. Milking was particularly important in northwestern Anatolia, pointing to regional differences linked with conditions more favourable to cattle compared to other regions, where sheep and goats were relatively common and milk use less important. The latter is supported by correlations between the fat type and animal bone evidence.

Direct evidence for the existence of dairying farms in prehistoric Central Europe (4th millennium BC).

The molecular and isotopic chemistry of organic residues from archaeological potsherds was used to obtain further insight into the dietary trends and economies at the Constance lake-shore Neolithic settlements. The archaeological organic residues from the Early Late Neolithic (3922-3902 BC) site Hornstaad-Hornle IA/Germany are, at present, the oldest archaeological samples analysed at the Institute of Mineralogy and Geochemistry of the University of Lausanne. The approach includes 13C/12C and 15N/14N ratios of the bulk organic residues, fatty acids distribution and 13C/12C ratios of individual fatty acids. The results are compared with those obtained from the over 500 years younger Neolithic (3384-3370 BC) settlement of Arbon Bleiche 3/Switzerland and with samples of modern vegetable oils and fat of animals that have been fed exclusively on C3 forage grasses. The overall fatty acid composition (C9 to C24 range, maximizing at C14 and C16), the bulk 13C/12C and 15N/14N ratios (delta13C, delta15N) and the 13C/12C ratios of palmitic (C16:0), stearic (C18:0) and oleic acids (C18:1) of the organic residues indicate that most of the studied samples (25 from 47 samples and 5 from 41 in the delta13C18:0 vs. delta13C16:0 and delta13C18:0 vs. delta13C18:1 diagrams, respectively) from Hornstaad-Hornle IA and Arbon Bleiche 3 sherds contain fat residues of pre-industrial ruminant milk, and young suckling calf/lamb adipose. These data provide direct proof of milk and meat (mainly from young suckling calves) consumption and farming practices for a sustainable dairying in Neolithic villages in central Europe around 4000 BC.dagger.

[History and economic importance of cattle (Bos taurus L.) in Switzerland from Neolithic to Early Middle Ages]. / Geschichte und wirtschaftliche Bedeutung des Hausrindes (Bos taurus L.) in der Schweiz von der Jungsteinzeit bis ins frühe Mittelalter.

In Switzerland domestic cattle (Bos primigenius f. taurus resp. Bos taurus L.) first appear with the earliest Neolithic settlements (approximately 5000 BC). With the gradual deforestation of the landscape caused by human exploitation of the environment, cattle were used more intensive and in many ways. There is evidence that cattle were used as draught animal since ca. 3400 BC, probably even earlier milk was regularly used. The size of domestic cattle gradually decreased from Early Neolithic until Iron Age. Only with Roman influence larger animals are found. However, after the withdrawal of Romans the average size of cattle decreased again. Archaeogenetic studies will have to show, whether this is due to novel breeding strategies or the import of breeding stock. First genetic results showed that a female genetic type, which is rare in European breeds, is present in Swiss Evolène cattle and in one animal of Roman time cattle from Augusta Raurica. Is this a sign for influence of Roman cattle on today's Swiss breeds?