The genomic history of southeastern Europe.

Farming was first introduced to Europe in the mid-seventh millennium bc, and was associated with migrants from Anatolia who settled in the southeast before spreading throughout Europe. Here, to understand the dynamics of this process, we analysed genome-wide ancient DNA data from 225 individuals who lived in southeastern Europe and surrounding regions between 12000 and 500 bc. We document a west-east cline of ancestry in indigenous hunter-gatherers and, in eastern Europe, the early stages in the formation of Bronze Age steppe ancestry. We show that the first farmers of northern and western Europe dispersed through southeastern Europe with limited hunter-gatherer admixture, but that some early groups in the southeast mixed extensively with hunter-gatherers without the sex-biased admixture that prevailed later in the north and west. We also show that southeastern Europe continued to be a nexus between east and west after the arrival of farmers, with intermittent genetic contact with steppe populations occurring up to 2,000 years earlier than the migrations from the steppe that ultimately replaced much of the population of northern Europe.

Rapid Cortical Adaptation and the Role of Thalamic Synchrony during Wakefulness.

Rapid sensory adaptation is observed across all sensory systems, and strongly shapes sensory percepts in complex sensory environments. Yet despite its ubiquity and likely necessity for survival, the mechanistic basis is poorly understood. A wide range of primarily in vitro and anesthetized studies have demonstrated the emergence of adaptation at the level of primary sensory cortex, with only modest signatures in earlier stages of processing. The nature of rapid adaptation and how it shapes sensory representations during wakefulness, and thus the potential role in perceptual adaptation, is underexplored, as are the mechanisms that underlie this phenomenon. To address these knowledge gaps, we recorded spiking activity in primary somatosensory cortex (S1) and the upstream ventral posteromedial (VPm) thalamic nucleus in the vibrissa pathway of awake male and female mice, and quantified responses to whisker stimuli delivered in isolation and embedded in an adapting sensory background. We found that cortical sensory responses were indeed adapted by persistent sensory stimulation; putative excitatory neurons were profoundly adapted, and inhibitory neurons only modestly so. Further optogenetic manipulation experiments and network modeling suggest this largely reflects adaptive changes in synchronous thalamic firing combined with robust engagement of feedforward inhibition, with little contribution from synaptic depression. Taken together, these results suggest that cortical adaptation in the regime explored here results from changes in the timing of thalamic input, and the way in which this differentially impacts cortical excitation and feedforward inhibition, pointing to a prominent role of thalamic gating in rapid adaptation of primary sensory cortex.SIGNIFICANCE STATEMENT Rapid adaptation of sensory activity strongly shapes representations of sensory inputs across all sensory pathways over the timescale of seconds, and has profound effects on sensory perception. Despite its ubiquity and theoretical role in the efficient encoding of complex sensory environments, the mechanistic basis is poorly understood, particularly during wakefulness. In this study in the vibrissa pathway of awake mice, we show that cortical representations of sensory inputs are strongly shaped by rapid adaptation, and that this is mediated primarily by adaptive gating of the thalamic inputs to primary sensory cortex and the differential way in which these inputs engage cortical subpopulations of neurons.

Stimulus Context and Reward Contingency Induce Behavioral Adaptation in a Rodent Tactile Detection Task.

Behavioral adaptation is a prerequisite for survival in a constantly changing sensory environment, but the underlying strategies and relevant variables driving adaptive behavior are not well understood. Many learning models and neural theories consider probabilistic computations as an efficient way to solve a variety of tasks, especially if uncertainty is involved. Although this suggests a possible role for probabilistic inference and expectation in adaptive behaviors, there is little if any evidence of this relationship experimentally. Here, we investigated adaptive behavior in the rat model by using a well controlled behavioral paradigm within a psychophysical framework to predict and quantify changes in performance of animals trained on a simple whisker-based detection task. The sensory environment of the task was changed by transforming the probabilistic distribution of whisker deflection amplitudes systematically while measuring the animal's detection performance and corresponding rate of accumulated reward. We show that the psychometric function deviates significantly and reversibly depending on the probabilistic distribution of stimuli. This change in performance relates to accumulating a constant reward count across trials, yet it is exempt from changes in reward volume. Our simple model of reward accumulation captures the observed change in psychometric sensitivity and predicts a strategy seeking to maintain reward expectation across trials in the face of the changing stimulus distribution. We conclude that rats are able maintain a constant payoff under changing sensory conditions by flexibly adjusting their behavioral strategy. Our findings suggest the existence of an internal probabilistic model that facilitates behavioral adaptation when sensory demands change.SIGNIFICANCE STATEMENT The strategy animals use to deal with a complex and ever-changing world is a key to understanding natural behavior. This study provides evidence that rodent behavioral performance is highly flexible in the face of a changing stimulus distribution, consistent with a strategy to maintain a desired accumulation of reward.

The ROCEEH Out of Africa Database (ROAD): A large-scale research database serves as an indispensable tool for human evolutionary studies.

Large scale databases are critical for helping scientists decipher long-term patterns in human evolution. This paper describes the conception and development of such a research database and illustrates how big data can be harnessed to formulate new ideas about the past. The Role of Culture in Early Expansions of Humans (ROCEEH) is a transdisciplinary research center whose aim is to study the origins of culture and the multifaceted aspects of human expansions across Africa and Eurasia over the last three million years. To support its research, the ROCEEH team developed an online tool named the ROCEEH Out of Africa Database (ROAD) and implemented its web-based applications. ROAD integrates geographical data as well as archaeological, paleoanthropological, paleontological and paleobotanical content within a robust chronological framework. In fact, a unique feature of ROAD is its ability to dynamically link scientific data both spatially and temporally, thereby allowing its reuse in ways that were not originally conceived. The data stem from published sources spanning the last 150 years, including those generated by the research team. Descriptions of these data rely on the development of a standardized vocabulary and profit from online explanations of each table and attribute. By synthesizing legacy data, ROAD facilitates the reuse of heritage data in novel ways. Database queries yield structured information in a variety of interoperable formats. By visualizing data on maps, users can explore this vast dataset and develop their own theories. By downloading data, users can conduct further quantitative analyses, for example with Geographic Information Systems, modeling programs and artificial intelligence. In this paper, we demonstrate the innovative nature of ROAD and show how it helps scientists studying human evolution to access datasets from different fields, thereby connecting the social and natural sciences. Because it permits the reuse of "old" data in new ways, ROAD is now an indispensable tool for researchers of human evolution and paleogeography.

Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe.

How modern humans dispersed into Eurasia and Australasia, including the number of separate expansions and their timings, is highly debated [1, 2]. Two categories of models are proposed for the dispersal of non-Africans: (1) single dispersal, i.e., a single major diffusion of modern humans across Eurasia and Australasia [3-5]; and (2) multiple dispersal, i.e., additional earlier population expansions that may have contributed to the genetic diversity of some present-day humans outside of Africa [6-9]. Many variants of these models focus largely on Asia and Australasia, neglecting human dispersal into Europe, thus explaining only a subset of the entire colonization process outside of Africa [3-5, 8, 9]. The genetic diversity of the first modern humans who spread into Europe during the Late Pleistocene and the impact of subsequent climatic events on their demography are largely unknown. Here we analyze 55 complete human mitochondrial genomes (mtDNAs) of hunter-gatherers spanning ∼35,000 years of European prehistory. We unexpectedly find mtDNA lineage M in individuals prior to the Last Glacial Maximum (LGM). This lineage is absent in contemporary Europeans, although it is found at high frequency in modern Asians, Australasians, and Native Americans. Dating the most recent common ancestor of each of the modern non-African mtDNA clades reveals their single, late, and rapid dispersal less than 55,000 years ago. Demographic modeling not only indicates an LGM genetic bottleneck, but also provides surprising evidence of a major population turnover in Europe around 14,500 years ago during the Late Glacial, a period of climatic instability at the end of the Pleistocene.

The Nature of Culture: an eight-grade model for the evolution and expansion of cultural capacities in hominins and other animals.

Tracing the evolution of human culture through time is arguably one of the most controversial and complex scholarly endeavors, and a broad evolutionary analysis of how symbolic, linguistic, and cultural capacities emerged and developed in our species is lacking. Here we present a model that, in broad terms, aims to explain the evolution and portray the expansion of human cultural capacities (the EECC model), that can be used as a point of departure for further multidisciplinary discussion and more detailed investigation. The EECC model is designed to be flexible, and can be refined to accommodate future archaeological, paleoanthropological, genetic or evolutionary psychology/behavioral analyses and discoveries. Our proposed concept of cultural behavior differentiates between empirically traceable behavioral performances and behavioral capacities that are theoretical constructs. Based largely on archaeological data (the 'black box' that most directly opens up hominin cultural evolution), and on the extension of observable problem-solution distances, we identify eight grades of cultural capacity. Each of these grades is considered within evolutionary-biological and historical-social trajectories. Importantly, the model does not imply an inevitable progression, but focuses on expansion of cultural capacities based on the integration of earlier achievements. We conclude that there is not a single cultural capacity or a single set of abilities that enabled human culture; rather, several grades of cultural capacity in animals and hominins expanded during our evolution to shape who we are today.

Radiocarbon dating the late Middle Paleolithic and the Aurignacian of the Swabian Jura.

Many lines of evidence point to the period between roughly 40 and 30 ka BP as the period in which modern humans arrived in Europe and displaced the indigenous Neandertal populations. At the same time, many innovations associated with the Upper Paleolithic--including new stone and organic technologies, use of personal ornaments, figurative art, and musical instruments--are first documented in the European archaeological record. Dating the events of this period is challenging for several reasons. In the period about six to seven radiocarbon half-lives ago, variable preservation, pre-treatment, and sample preparation can easily lead to a lack of reproducibility between samples and laboratories. A range of biological, cultural, and geological processes can lead to mixing of archaeological strata and their contents. Additionally, some data sets point to this period as a time of significant spikes in levels of atmospheric radiocarbon. This paper assesses these questions in the context of the well-excavated and intensively studied caves of Geissenklösterle and Hohle Fels in the Swabian Jura of southwestern Germany. We conclude that variable atmospheric radiocarbon production contributes to the problems of dating the late Middle Paleolithic and the early Upper Paleolithic. To help establish a reliable chronology for the Swabian Aurignacian, we are beginning to focus our dating program on short-lived, stratigraphically secure features to see if they yield reproducible results. This approach may help to test competing explanations for the noisy and often non-reproducible results that arise when trying to date the transition from the Middle to the Upper Paleolithic.

Hammer or crescent wrench? Stone-tool form and function in the Aurignacian of southwest Germany.

The early Upper Paleolithic of Europe is associated with the appearance of blade/bladelet technology (e.g., Aurignacian). These industries include a wider range of formal tool types than seen in the Middle Paleolithic. Greater diversity in tool types is often interpreted as specialized tools created for specific tasks. This, in turn, is said to reflect dramatic behavioral shifts between Neandertals and modern humans. In order to test previous interpretations, it is necessary to have a detailed understanding of early Upper Paleolithic stone-tool function. Toward this end, analyses of microscopic residue and use-wear were undertaken on 109 stone tools from three Aurignacian sites in southwest Germany (Hohle Fels, Geissenklösterle, and Vogelherd). These cave sites evidenced remarkable residue preservation, with approximately 82% of the sample showing some form of functional evidence. Residues observed included hair, feathers, bone/antler, wood, plant tissue, phytoliths, starch grains, and resin. The results suggest that tool typology is not strongly linked to the processing of specific materials. For example, endscrapers from the sample show evidence of processing wood, charred wood, plants, starchy plants, birds, bone/antler, and animals (hair). Hairs are found on tools typologically classified as blades, flakes, borers, pointed blades, and combination tools (nosed endscraper-borer, burin-laterally-retouched blade). In the early Upper Paleolithic of southwest Germany, a wide range of tool types appears to have been used to process a diverse array of materials. These results suggest that the interpretation of behavioral patterns from stone tools must consider more than tool typology.

Radiocarbon dating the appearance of modern humans and timing of cultural innovations in Europe: new results and new challenges.

New radiocarbon dates from the sites of Bockstein-Törle, Geissenklösterle, Hohle Fels, Hohlenstein-Stadel, Sirgenstein, and Vogelherd in the Swabian Jura of southwestern Germany indicate that the Aurignacian of the region spans the period from ca. 40-30ka BP. If the situation at Vogelherd, in which skeletal remains from modern humans underlie an entire Aurignacian sequence, is viewed as representative for the region, the dates from the Swabian Jura support the hypothesis that populations of modern humans entered the region by way of the "Danube Corridor." The lithic technology from the lower Aurignacian of Geissenklösterle III is fully developed, and classic Aurignacian forms are well represented. During the course of the Aurignacian, numerous assemblages rich in art works, jewelry, and musical instruments are documented. By no later than 29ka BP the Gravettian was well established in the region. These dates are consistent with the "Kulturpumpe" hypothesis that important cultural innovations of the Aurignacian and Gravettian in Swabia predate similar developments in other regions of Europe. The radiocarbon dates from Geissenklösterle corroborate observations from other non-archaeological data sets indicating large global fluctuations in the atmospheric concentrations of radiocarbon between 30 and 50ka calendar years ago. These fluctuations lead to complications in building reliable chronologies during this period and cause the "Middle Paleolithic Dating Anomaly" and the "Coexistence Effect," which tend to exaggerate the temporal overlap between Neanderthals and modern humans.