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Isotopic analysis on bone collagen and tooth enamel

Bones and teeth consist primarily of two main components, the inorganic bioapatite and the organic collagen. The preservation of collagen in bones depends very much on the taphonomic conditions. However, under good conditions, such as those prevailing in cave sites or permafrost, collagen can be preserved for a very long time and thus provide important information about the diet of the sampled individual, the ecological niches, and also the age. Bioapatite is even longer lasting and is usually sampled from the tooth, where it makes up 97% of the enamel. In this context, stable isotopes can provide important information about the climate and ecological conditions, but also give insight into the diet of the individual.

Bone collagen: Stable carbon and nitrogen isotopes

Primarily, stable carbon and nitrogen isotopes (δ13C and δ15N) will be analyzed from well-preserved bone collagen. However, to make this possible, the collagen must first be chemically extracted from the bone. For this purpose, the method of Bocherens, et al. (1997) is suitable, which includes grinding of the bone samples to powder, and soaking the bone powder in 0.125 M NaOH between the demineralization and solubilization steps to eliminate lipids and humic acids. After extraction, the collagen can be analyzed for the stable δ13C and δ15N isotopes using an elemental analyzer coupled to an isotope ratio mass spectrometer.

The δ13C and δ15N are directly linked to the protein fraction of the diet (Ambrose and Norr, 1993, Tieszen and Fagre, 1993) and increase with the trophic level of the consumer (Krajcarz, et al., 2018). Based on this, it is possible to reconstruct the diet and determine the trophic niche of an individual, as shown by many studies in paleoecology and archaeology (Wißing, et al., 2019, Baumann, et al., 2020a, Krajcarz, et al., 2020, Baumann, et al., 2020b, Baumann, et al., 2021).

Tooth enamel: Stable carbon and oxygen isotopes

In order to analyze the inorganic components, the bioapatite in tooth enamel, the samples are chemically purified over several steps and all organic components, which could contain carbon, will be removed (Bocherens, et al., 1994). A multiflow coupled with an isotope ratio mass spectrometer will be used to measure the ratio between the isotopes of carbon and oxygen atoms. The δ13C and δ18O values obtained are not, as in the case of collagen, only from the protein diet, but from the whole diet and the water drunk by the individual. Thus, it is also possible to determine the average temperatures during the lifetime of the individual by the measured δ18O values (Gehler, et al., 2012).

Schematic overview of the analysis of stable isotopes. A similar approach is used for radiocarbon of C14 dating. Click to read more about C14 dating.

See also palaeopathology to find out more about the effects of diet on health.References Back

Bone collagen: 14C dating of bones

The extracted collagen can be used for the direct dating of archaeological bones as well. Here, however, not stable isotopes such as 13C are measured, but the radioactive 14C, which decays with time to 14N. Similar to 13C, living organisms absorb 14C from food and water over their entire lifetime. With the death of the individual, this uptake ends and the 14C in the body (or bone) decays. Using the known half-life of 14C (5730 ± 40 years), it is possible to calculate the age of the sample. With this method, fossils can be dated up to around 60,000 years before today. To date the collagen, the sample will be prepared according to the protocol of Hajdas (2008) and the 14C/12C ratio will be measured by using an accelerator mass spectrometry. In the last step, the obtained values have to be calibrated against dentrochronology, by using a software such as OxCal (Bronk Ramsey, 2017).

See also the analysis of stable isotopes to learn more about sample preparation and collagen extraction which is similar for both analysisReferencesBack

Ancient protein analysis can be applied to mummified human remains for multiple analytical purposes. Protein preserves longer than DNA, and offers an additional method to study archaeological individuals and biological materials that may no longer contain genetic material. As proteins are comprised of sequences of amino acids, differences in these sequences provide a way to differentiate species-specific animal, plant, and bacterial proteins contained within a sample. The extraction and study of ancient proteins has been used in the past on a diverse range of archaeological materials, such as human and animal bones, leather hides, dental calculus (calcified dental plaque), residues found in ceramic vessels, and historic artistic materials. As mummies are composed of human tissues, their tissues cab be analyzed to investigate questions about the individual’s heath and diet, the composition of their microbiome or necrobiome, and in theory, can be utlized to identify specific tissues. Protein analysis has been used to identify the stomach contacts of Ötzi, better known as the Iceman found in the Tyrolian Alps, and found that his last meal consisted of goat and deer meat, as well as wheat (Maixner citation). Additionally, samples taken from the mouths Peruvian child mummies found high in the Andes demonstrated both the presence of mycobacterial disease (through aDNA) and proteins indicative of the human immune response to these pathogens (Peru citation). Interestingly, an analysis of materials worn around the necks of mummies from the Tarim Basin of Xinjiang were shown to have consisted of dried kefir cheeses made from the processed milk of their domesticated ruminant animals (cow, sheep, goat) herded by the burial population.Protein analysis of mummies can also be combined with aDNA, lipid (fats), stable isotope analysis to gain a better and more thorough understanding of the overall samples and individuals under study. The use of proteomics on mummified individuals is still in its infancy and continued studies in its development are underway, many of which are ongoing here at the Institute for Evolutionary Medicine.

Proteomics

The proteins are extracted from the source, as for example calcified dental plaque, then digested into smaller pieces called peptides. These are then cleaned and analysed via mass spectronomy. This results in a specific profile for each protein and allows them to be distinguished. See also aDNA which also often uses similar sourcesSee also stable isotope analysis which can be also used to investigate diet BackReferences

Maixner, F., Turaev, D., Cazenave-Gassiot, A., Janko, M., Krause-Kyora, B., Hoopmann, M. R., Kusebauch, U., Sartain, M., Guerriero, G., O’Sullivan, N. & Others. The iceman’s last meal consisted of fat, wild meat, and cereals. Curr. Biol. 28, 2348–2355 (2018).

Maixner, F., Overath, T., Linke, D., Janko, M., Guerriero, G., van den Berg, B. H. J., Stade, B., Leidinger, P., Backes, C., Jaremek, M., Kneissl, B., Meder, B., Franke, A., Egarter-Vigl, E., Meese, E., Schwarz, A., Tholey, A., Zink, A. & Keller, A. Paleoproteomic study of the Iceman’s brain tissue. Cell. Mol. Life Sci. 70, 3709–3722 (2013).

Maixner, F., Krause-Kyora, B., Turaev, D., Herbig, A., Hoopmann, M. R., Hallows, J. L., Kusebauch, U., Vigl, E. E., Malfertheiner, P., Megraud, F., O’Sullivan, N., Cipollini, G., Coia, V., Samadelli, M., Engstrand, L., Linz, B., Moritz, R. L., Grimm, R., Krause, J., Nebel, A., Moodley, Y., Rattei, T. & Zink, A. The 5300-year-old Helicobacter pylori genome of the Iceman. Science 351, 162–165 (2016).

Corthals, A., Koller, A., Martin, D. W., Rieger, R., Chen, E. I., Bernaski, M., Recagno, G. & Dávalos, L. M. Detecting the Immune System Response of a 500 Year-Old Inca Mummy. PLoS ONE 7, e41244 (2012).

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Bocherens, H., Billiou, D., Patou-Mathis, M., Bonjean, D., Otte, M., Mariotti, A., 1997. Paleobiological Implications of the Isotopic Signatures (13C, 15N) of Fossil Mammal Collagen in Scladina Cave (Sclayn, Belgium), Quaternary Research 48, 370-380.

Ambrose, S.H., Norr, L., 1993. Experimental evidence for the relationship of the carbon isotope ratios of whole diet and dietary protein to those of bone collagen and carbonate, in: Lambert, J., Grupe, G. (Eds.), Prehistoric Human Bone: Archaeology at the Molecular Level, Springer, Berlin, pp. 1-37.

Tieszen, L.L., Fagre, T., 1993. Effect of diet quality and composition on the isotopic composition of respiratory CO 2, bone collagen, bioapatite, and soft tissues, in: Lambert, J., Grupe, G. (Eds.), Prehistoric Human Bone: Archaeology at the Molecular Level, Springer, Berlin, pp. 121-155.

Krajcarz, M.T., Krajcarz, M., Bocherens, H., 2018. Collagen-to-collagen prey-predator isotopic enrichment ( Δ 13 C, Δ 15 N) in terrestrial mammals - a case study of a subfossil red fox den, Palaeogeography, Palaeoclimatology, Palaeoecology 490, 563-570.

Wißing, C., Rougier, H., Baumann, C., Comeyne, A., Crevecoeur, I., Drucker, D.G., Gaudzinski-Windheuser, S., Germonpré, M., Gómez-Olivencia, A., Krause, J., Matthies, T., Naito, Y.I., Posth, C., Semal, P., Street, M., Bocherens, H., 2019. Stable isotopes reveal patterns of diet and mobility in the last Neandertals and first modern humans in Europe, Scientific Reports 9.

Baumann, C., Starkovich, B.M., Drucker, D.G., Münzel, S.C., Conard, N.J., Bocherens, H., 2020a. Dietary niche partitioning among Magdalenian canids in southwestern Germany and Switzerland Quaternary Science Reviews 227, 106032.

Krajcarz, M., Krajcarz, M.T., Baca, M., Baumann, C., Van Neer, W., Popović, D., Sudoł-Procyk, M., Wach, B., Wilczyński, J., Wojenka, M., Bocherens, H., 2020. Ancestors of domestic cats in Neolithic Central Europe: Isotopic evidence of a synanthropic diet, Proceedings of the National Academy of Sciences, 201918884.

Baumann, C., Bocherens, H., Drucker, D.G., Conard, N.J., 2020b. Fox dietary ecology as a tracer of human impact on Pleistocene ecosystems, PLOS ONE 15, e0235692.

Baumann, C., Pfrengle, S., Münzel, S.C., Molak, M., Feuerborn, T., Breidenstein, A., Reiter, E., Albrecht, G., Kind, C.-J., Verjux, C., Leduc, C., Conard, N.J., Drucker, D.G., Giemsch, L., Thalmann, O., Bocherens, H., Schuenemann, V.J., 2021. A refined proposal for the origin of dogs: The case study of Gnirshöhle, a Magdalenian cave site, Scientific Reports 11:5137.

Hajdas, I., 2008. Radiocarbon dating and its applications in Quaternary studies, Eiszeitalter und Gegenwart Quaternary Science Journal 57, 24.

Bronk Ramsey, C., 2017. OxCal Program, Version 4.3, Oxford Radiocarbon Accelerator Unit: University of Oxford. Available at ….

Bocherens, H., Fizet, M., Mariotti, A., 1994. Diet, physiology and ecology of fossil mammals asinferred from stable carbon and nitrogen isotope biogeochemistry: implications for Pleistocene bears, Paleogeography, Paleoclimatology, Paleoecology 107, 213-225.

Gehler, A., Tütken, T., Pack, A., 2012. Oxygen and carbon isotope variations in a modern rodent community–implications for palaeoenvironmental reconstructions, PLoS One 7, e49531.

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Paleogenetics conducts research on ancient DNA (aDNA), which is defined as DNA extracted from biological remains that are not preserved for later DNA analyses (Hofreiter et al. 2001). This research harbors several challenges as due to nature of aDNA, which is highly fragment, contains chemical modifications such as deaminated cytosines and is only present very low amounts because of its exposure to various environmental conditions (Lindahl, 1993, Hofreiter at al. 2001, Briggs et al. 2007). In recent years, advances in sequencing technologies (Next Generation Sequencing) and simultaneous improvements in the ancient DNA field have allow the reconstruction of full genomes of extinct species, e.g., the woolly mammoth (Miller et al., 2008) or the Neanderthal (Green et al., 2010). Furthermore, NGS based technologies also lead to a better understanding of chemical modifications of aDNA resulting in the establishment of the characteristic ancient DNA profiles and tools for contamination tests (Briggs et al., 2007). Using all these advances, the field of paleogenetics has opened a new means of investigating the genetic diversity and evolution of the human species and other organisms (e.g., Briggs et al. 2007, Mathieson et al. 2018, Bos et al. 2011).Although the first samples trialed were from Egyptian mummies (Pääbo et al. 1985, Pääbo & Wilson 1988), genetic studies on such material are rare due to methodological and contamination issues. Conditions specific to the material such as the hot Egyptian climate, the high humidity levels in many tombs and some of the chemicals used in mummification techniques were thought to damage DNA beyond retrieval and therefore to make the long-term survival of DNA in Egyptian mummies unlikely (Hofreiter et al. 2014, Gilbert et al. 2005). Thus, with the use of state-of-the-art NGS techniques and rigorous contamination tests the first successful paleogenetic investigation of the Nile Valley yielded authentic ancient Egyptian DNA resulting in the reconstruction of the first Egyptian mitochondrial genomes as well as nuclear genomic data (Schuenemann et al. 2017). Following up on this study the first large scale metagenomics analysis of ancient Egyptian material was conducted on the same individuals from the archeological site of Abusir el-Meleq including among others in retrieval of oral microbiome signatures and the reconstruction of two ancient pathogen genomes (Neukamm et al. 2020). Overall, these studies highlight the potential of aDNA research to a better understanding of Egypt’s population history and past health status.

Palaeogenetics

To protect ancient DNA fragments from modern contamination aDNA work has to be carried out in dedicated cleanroom facilities. aDNA can be used to shed light on family relationships, genetic ancestry and to detect DNA of pathogens. Click and read more about the investigation of diseases in mummies. Teeth are often used to extract aDNA. As a closed capsule they protect the DNA fragments against the environment. From the tiny blood vessels in the pulpa it is also possible to detect pathogens that entered the bloodstream. Click to find out what else can be learned from analysing stable isotopes from tooth enamel.

References

Hofreiter, M., Serre, D., Poinar, H. N., Kuch, M., & Pääbo, S. (2001). ancient DNA. Nature Reviews Genetics, 2(5), 353-359.

Lindahl, T. (1993). Instability and decay of the primary structure of DNA. nature, 362(6422), 709-715.

Briggs, A. W., Stenzel, U., Johnson, P. L., Green, R. E., Kelso, J., Prüfer, K., ... & Pääbo, S. (2007). Patterns of damage in genomic DNA sequences from a Neandertal. Proceedings of the National Academy of Sciences, 104(37), 14616-14621.

Miller, W., Drautz, D. I., Ratan, A., Pusey, B., Qi, J., Lesk, A. M., ... & Schuster, S. C. (2008). Sequencing the nuclear genome of the extinct woolly mammoth. Nature, 456(7220), 387-390.

Green, R. E., Krause, J., Briggs, A. W., Maricic, T., Stenzel, U., Kircher, M., ... & Pääbo, S. (2010). A draft sequence of the Neandertal genome. science, 328(5979), 710-722.

Mathieson, I., Alpaslan-Roodenberg, S., Posth, C., Szécsényi-Nagy, A., Rohland, N., Mallick, S., ... & Reich, D. (2018). The genomic history of southeastern Europe. Nature, 555(7695), 197-203.

Bos, K. I., Schuenemann, V. J., Golding, G. B., Burbano, H. A., Waglechner, N., Coombes, B. K., ... & Krause, J. (2011). A draft genome of Yersinia pestis from victims of the Black Death. Nature, 478(7370), 506-510.

Pääbo, S. (1985). Molecular cloning of ancient Egyptian mummy DNA. nature, 314(6012), 644-645.

Pääbo, S., Gifford, J. A., & Wilson, A. C. (1988). Mitochondrial DNA sequences from a 7000-year old brain. Nucleic acids research, 16(20), 9775-9787.

Shapiro, B., & Hofreiter, M. (2014). A paleogenomic perspective on evolution and gene function: new insights from ancient DNA. Science, 343(6169).

Gilbert, M. T. P., Bandelt, H. J., Hofreiter, M., & Barnes, I. (2005). Assessing ancient DNA studies. Trends in ecology & evolution, 20(10), 541-544.

Schuenemann, V. J., Peltzer, A., Welte, B., Van Pelt, W. P., Molak, M., Wang, C. C., ... & Krause, J. (2017). Ancient Egyptian mummy genomes suggest an increase of Sub-Saharan African ancestry in post-Roman periods. Nature communications, 8(1), 1-11.

Neukamm, J., Pfrengle, S., Molak, M., Seitz, A., Francken, M., Eppenberger, P., ... & Schuenemann, V. J. (2020). 2000-year-old pathogen genomes reconstructed from metagenomic analysis of Egyptian mummified individuals. BMC biology, 18(1), 1-18.

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Cultural and ecological context

The people whose remains we might find did not exist in isolation. What kind of society did they live in? Did they have kings or chiefs, arts and crafts, war and peace? And what was the environment they lived in like? How did people get their food? Sometimes these questions can be answered by consulting relevant historical sources or by examining artefacts associated with the remains and interpreting them in light of other archaeological evidence. But even when direct historical or archaeological evidence is absent or inconclusive, we can draw from knowledge of current human diversity to make inferences about cultural and ecological context. How does this work?In the last few hundred years, anthropologists have described the lifeways of numerous cultures around the globe. They documented their kinship and marriage systems, religious beliefs, subsistence practices, and technology. Most of these “ethnographies” are now easily accessible, for example through eHRAF (https://ehrafworldcultures.yale.edu/ehrafe/ ). Here, users can search for records on a specific culture, from A’ani to Zuñi, or on a specific topic, such as religion or warfare. Moreover, the information from many ethnographies has already been coded and compiled for comparative analyses, for example in the globally representative Standard Cross-Cultural Sample or Lewis Binford’s sample of 339 hunter-gatherer societies – these samples, along with relevant ecological variables and information on the phylogenetic relatedness among societies can be accessed at D-PLACE (https://d-place.org/). Many researchers have used these samples to find general patterns that can help us understand how human societies adapt to their environment, and to make informed predictions about an unknown society. For example, hunter-gatherers living farther away from the tropics rely more on meat and those closer to the tropics more on plant food; similarly, those relying on mobile game and other unpredictable foods typically have high mobility, small group size, and relatively egalitarian hierarchies whereas those exploiting rich, predictable foods tend to live in larger, more permanent settlements and have more formal hierarchies – with chiefs, warrior classes, and even slavery. The general trend of these “complex hunter-gatherers” becomes more pronounced with agriculture and animal domestication. Agricultural land and livestock can be monopolized by elites, or contested among groups through warfare. Hence, many agricultural and pastoralist societies are larger, more hierarchical and more complex than hunter-gatherers.Such general patterns in the ethnographic record can to some extent be used to make inferences about past societies. For instance, if human remains clearly stem from a hunter-gatherer context we can expect some similarities in social organization and complexity to ethnographically-documented hunter-gatherers. Similarly, by characterizing the ecological environment we can get a sense for what kinds of resources people would have relied on and how that might have shaped their society. But some caution is warranted: ethnographically-documented societies were not living fossils that have remained unchanged since prehistory. Indeed, many had advanced technologies that would not have been available in earlier times. Similarly, ethnographically-documented societies may well not capture the full range of variation that has existed in the past. For example, it is often argued that people living as hunter-gatherers in recent history occupied poor, marginal environments that were not suitable for farming or herding, and that “complex hunter-gatherers” could have been much more common in the past. Archeological findings associated with human remains can therefore be helpful to ground extrapolation – for instance, many burials as far back as the upper Paleolithic had sophisticated grave goods indicative of advanced craftsmanship and high social status, arguably reflecting relatively complex societies with divisions of labor and formal hierarchies – unlike most mobile foragers of recent history.

The ethnographic record

General cross-cultural patterns

Extrapolating from the present to the past

See also provenance studies to learn more about the context of the Egyptian mummies

https://ehrafworldcultures.yale.edu/ehrafe/https://d-place.org/

Many impressive artefacts and buildings from ancient Egypt have been preserved, which provide information about the life and culture of the people.

Paleopathology, a sub-discipline of evolutionary medicine attempts to describe human diseases in the past and identify changes in disease patterns as a consequence of humankind's history. Thus, paleopathology is the study of ancient diseases in humans but all so in animals and therefore can contribute to a more holistic, One health-like understanding. Its main society, the Paleopathology Association was formed in 1973 by North American scientists. Today, the association is a global network composed of researchers and students with backgrounds in various fields such as physical anthropology, medicine, biology, archaeology, pathology and zoology.

The study of ancient human disease – palaeopathology- is etymologically derived from the ancient Greek words παλαιός (ancient), πάθειν (to suffer) and λόγος (discipline). The term “palaeopathology” was coined in 1893 by Robert W. Shufeldt (1850-1934), yet the very first studies can be dated much earlier; for instance the one by Felix Platter (1536 –1614) on possible gigantism. In the late 19th century, among others, famous Rudolf Virchow (1823 – 1902) or Paul Broca (1824 – 1880) had a notable influence on the field of palaeopathology. Sir Marc A. Ruffer (1845 – 1917) is generally recognised as the founder of palaeopathology as an academic (sub-) discipline particularly due to his posthumously published treatise “Studies in the Palaeopathology of Egypt” (1921).
Some selected research aims within the field are as follows:

- To study the evidence of ill health of the past to understand the nature and extent of disease today in general.

- To analyze skeletal and mummified human remains (with various degree of preservation of soft tissues) for evidence of disease.

- To what extent are sicker or healthier than we were prior to the introduction of modern standards of living, antibiotics, and chemotherapeutics?

Major online links: See also aDNA analysis to identify pathogens or genetic markers for diseasesSee also proteomics or stable isotope analysis to learn more about how to analyse dietary habits Wikipedia: https://en.wikipedia.org/wiki/PaleopathologyGeneral overview with relevant selected publications, authors, journals: https://www.sciencedirect.com/topics/medicine-and-dentistry/paleopathology
Paleopathology Association: https://paleopathology-association.wildapricot.org/

Palaeopathology

Mummies represent a great opportunity for palaeopathological studies since in most cases not only the skeleton but also some soft tissue is preserved. In this way, analyses are not restricted to diseases affecting the bones. On the right picture, X-ray images are used to visualise blood vessels and arteriosclerotic alterations of the mummy of Nes-Schuh. On the left side, the X-ray images show pathological alterations on the knee of a different mummy. Click and read more about X-rays.

Palaeoradiology

Paleoradiology: When investigating ancient skeletonized or mummified human remains, we must follow ethical considerations comparable to those in a clinical setting, where before any analysis, the degree of invasiveness is weighted against the expected progress in knowledge (1). Consequently, paleopathological research places a major focus on non-invasive medical imaging modalities and imaging-guided minimally-invasive sampling techniques, which, in this context, are subsumed as "Palaeoradiology" (2). Radiographic imaging (planar X-rays and CT) complements macroscopic analyses of skeletal remains and is indispensable when examining mummified human remains during field research and excavations (3-5) and is also generally an excellent tool for long-term documentation (6, 7). Currently, the most established diagnostic imaging modalities to study ancient human remains are conventional X-rays and computed tomography (CT). From case reports towards paleo-epidemiological studies: So far, paleoradiological examinations have identified numerous diseases and conditions in ancient human remains, often dating back several thousand years (18). A broader overview of the development and range of the current technologies and methods for investigating mummies has been published by Böni (19) and Lynnerup (20), and for diagnostic imaging methodology specifically by Rühli and Chhem (21). More recently, paleopathology tries increasingly to move towards a more paleo-epidemiological perspective by applying evidence-based examination criteria to larger series of ancient human specimens (22-25).Computed Tomography (CT): While complex anatomical structures, such as the skull base or remnants of internal organs, can often not be adequately represented by planar radiographs due to superimposition artifacts, tomographic imaging techniques such as CT or micro-CT overcome these limitations (12). Additionally, tomographic modalities offer a high spatial resolution and various post-processing options. The first landmark in this field of research was the Manchester Mummy project in the late 1970s (13). Today, CT is often considered the "gold standard" for imaging studies of ancient human remains in general and whole mummies in particular (14-17). Conventional X-rays: Radiographic investigations of ancient human remains began in 1896 with Walter Koenig's (1859-1936) publication of a bandaged ancient Egyptian child mummy (8). Even today, outside of the clinical setting, portable digital conventional radiography is often the only viable imaging modality during archaeological excavations. However, conventional radiography projects 3-dimensional structures onto a 2-dimensional image, resulting in multiple structures being superimposed on one radiograph. Besides using radiography to diagnose specific diseases, also alterations in bone morphology, indicative of a more general health (or growth) status (Harris lines, body mass, osteoporosis), can be identified (9-11).

In the field, a portable X-ray device is used to take images. Several X-ray images are taken and put together later on to have an image of the entire body. In this way also pathological alterations in the interior can be made visible. Click and read more about pathologies.

References

1. Kreissl Lonfat BM, Kaufmann IM, Rühli FJ. A Code of Ethics for Evidence-Based Research With Ancient Human Remains. Anatomical Record. 2015;298(6):1175-81.

2. Notman DN, Anderson L, Beattie OB, Amy R. Arctic paleoradiology: portable radiographic examination of two frozen sailors from the Franklin expedition (1845-1848). AJR Am J Roentgenol. 1987;149(2):347-50.

3. Chhem RK, Brothwell DR. Paleoradiology imaging mummies and fossils. Berlin: Springer; 2008. Online-Ressource p.

4. Seiler R, Eppenberger P, Rühli F. Application of portable digital radiography for dental investigations of ancient Egyptian mummies during archaeological excavations: Evaluation and discussion of the advantages and limitations of different approaches and projections. Imaging Science in Dentistry. 2018;48(3):167-76.

5. Conlogue G, Nelson AJ, Guillen S. The application of radiography to field studies in physical anthropology. Can Assoc Radiol J. 2004;55(4):254-7.

6. Ortner DJ, Hotz G. Skeletal manifestations of hypothyroidism from Switzerland. American journal of physical anthropology. 2005;127(1):1-6.

7. Manchester K, Roberts C. The palaeopathology of leprosy in Britain: a review. World Archaeol. 1989;21(2):265-72.

8. König W. 14 Photographien mit Röntgen-Strahlen. Aufgenommen im physikalischen Verein zu Frankfurt A. M. Leipzig: Verlag von Johann Ambrosius Barth; 1896.

9. Curate F. Osteoporosis and paleopathology: a review. J Anthropol Sci. 2014;92:119-46.

10. Ruff CB. Body mass prediction from skeletal frame size in elite athletes. American Journal of Physical Anthropology. 2000;113(4):507-17.

11. Harris HA. Bone growth in health and disease; the biological principles underlying the clinical, radiological, and histological diagnosis of perversions of growth and disease in the skeleton. London,: Oxford University Press, H. Milford; 1933. xv, 248 p. p.

12. Eppenberger P, Rühli FJ. Imaging in Human Remains. In: López Varela SL, editor. The Encyclopedia of Archaeological Sciences. Hoboken, NJ: John Wiley & Sons; 2018. p. Epub ahead of print.

13. Harwood-Nash DC. Computed tomography of ancient Egyptian mummies. Journal of computer assisted tomography. 1979;3(6):768-73.

14. Conlogue G. Considered Limitations and Possible Applications of Computed Tomography in Mummy Research. Anat Rec. 2015;298(6):1088-98.

15. Lynnerup N, Rühli FJ. Short review: the use of conventional X-rays in mummy studies. Anatomical Record. 2015;298(6):1085-7.

16. Panzer S, Ketterl S, Bicker R, Schoske S, Nerlich AG. How to CT scan human mummies: Theoretical considerations and examples of use. International journal of paleopathology. 2019;26:122-34.

17. Cox SL. A Critical Look at Mummy CT Scanning. Anat Rec (Hoboken). 2015;298(6):1099-110.

18. Sjovold T. The Tyrolean Iceman and excavated human remains as sources of information about the past, the present, and the future. Int J Circumpolar Health. 1998;57 Suppl 1:49-54.

19. Böni T, Rühli FJ, Chhem RK. History of paleoradiology: early published literature, 1896-1921. Canadian Association of Radiologists Journal. 2004;55(4):203-10.

20. Lynnerup N. Mummies. American journal of physical anthropology. 2007;Suppl 45:162-90.

21. Chhem RK, Rühli FJ. Paleoradiology: current status and future challenges. Canadian Association of Radiologists Journal. 2004;55(4):198-9.

22. Camacho M, Araujo A, Morrow J, Buikstra J, Reinhard K. Recovering parasites from mummies and coprolites: an epidemiological approach. Parasit Vectors. 2018;11(1):248.

23. Lalremruata A, Ball M, Bianucci R, Welte B, Nerlich AG, Kun JFJ, et al. Molecular identification of falciparum malaria and human tuberculosis co-infections in mummies from the Fayum depression (Lower Egypt). PloS one. 2013;8(4):e60307.

24. Nerlich AG, Losch S. Paleopathology of human tuberculosis and the potential role of climate. Interdiscip Perspect Infect Dis. 2009;2009:437187.

25. Rühli FJ, Ikram S, Bickel S. New ancient Egyptian human mummies from the valley of the kings, Luxor: anthropological, radiological, and egyptological Iivestigations. BioMed Research International. 2015;2015:530362.

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Over the course of the ancient Egyptian history, additional burial objects accompanied the deceased: a cartonnage mask was put over his/her face, numerous small amulets were placed within the linen bandages and jewellery embellished the body. Then the mummy was placed inside a coffin or sarcophagus made of wood or stone or inside a nest of several coffins within a sarcophagus.

The coffins and sarcophagi were either rectangular or were mummy shaped. The internal organs – the lung, liver, stomach and the intestines - were also embalmed and the viscera placed in separate jars, the so called Canopic jars, or they were put back in the body cavity. Religious texts and figurative scenes on body containers, tomb walls or papyri also accompanied the dead for his/her protection and to safely reach the hereafter.

Mummies are an invaluable historical and scientific source of knowledge on the life, religious practices, cultural traditions as well as health states and pathologies of the ancient Egyptian civilization. The often unique preservation of ancient Egyptian mummies enables humanities, natural and medical sciences to collect and analyze various types of data for interdisciplinary research.

To prevent the body from decaying, the brain and the internal organs – the lung, liver, stomach and intestines - were removed. Then the body was dried out by using salt which removed the body fluids and treated with resin. The heart as the seat of feeling and thinking remained inside the body. During the early years of artificial mummification, the deceased was wrapped in linen bandages and each body part was separately wrapped once the organs had been removed. Over time, the deceased was wrapped in more and more layers of linen resembling a cocoon shape. In ancient Egypt the deceased was mummified to preserve his/her body for the afterlife and to provide an intact container for his/her soul to live for eternity. Mummification procedures helped to protect the dead body from decomposition. In Prehistoric times (ca. 4000-3000 B.C.), the deceased was buried in the desert where dryness, heat and salt lead to his/her natural preservation. When the ancient Egyptians began to bury their dead in coffins and tombs, they discovered that the bodies started to decay. So the embalmers began to experiment with artificial mummification techniques which were gradually elaborated over time.Provenance (lat. word to come from) studies are the studies of the biography and the origin of an object. In the art historical context, provenance is defined as the history of the change of ownership of an object. Provenance studies deal with as complete a reconstruction of the ownership and succession of ownership of an object as possible. In other words, provenance studies are concerned with the history of the collector and the collection and attempts to clarify the ownership of an object through meticulous research of the object. Every object in a museum or archive (a picture, a statue, a mummy, etc.) has a story to tell and leads to questions such as who was the owner, who acquired it when and why? Where was it exhibited? Who saw it, who wrote about it? Who received it as a present and for what reason? A provenance researcher explores and tells these stories. More and more museums engage provenance researchers who check for individual objects of the collection when and how they entered the museum holdings. If the provenance researcher discovers or the museum receives a claim that an object in the collection was illegally acquired, the museum shall look for an appropriate restitution or resolve the matter in an agreeable mutual manner.

Origin of Egyptian Mummies

https://www.iem.uzh.ch/en/research/paleopathology_mummy_studies_group_rühli/project_swiss_mummy_project.html

https://www.mylearning.org/stories/a-step-by-step-guide-to-egyptian-mummification/220?

https://www.ancient.eu/article/44/mummification-in-ancient-egypt/

http://www.ancientegypt.co.uk/mummies/story/main.html

Salima Ikram, Aidan Dodson, The Mummy in Ancient Egypt. Equipping the Dead for Eternity. The American University in Cairo Press 1998.

Provenance

The vast majority of the Egyptian mummies derive from large cemeteries as for example Abusir el Meleq and probably represent the “normal” population. Only a few famous cases were buried in large monuments as seen on the right. Mummy of a child on the left. See also ecological and cultural context to learn more about how to investigate the life circumstances of the Egyptian mummies.

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Picture credits:

Picture 1 Palaeopathology: Clara Saner, Kantonsarchäologie Aargau

Picture 2 Palaeopathology: Prof. Dr. Frank Rühli

Picture 1 & 2 Cultural and ecological context: freepik.com

Title picture and picture 1 Provenance: Dr. Kerttu Majander

Picture 1 Provenance: freepik.com

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Acknowledgments

We thank all authors providing content:

Dr. Chris Baumann (University of Tübingen), Dr. Shevan Wilkin (University of Zurich), Prof. Dr. Verena Schünemann (University of Zurich), Prof. Dr. Frank Rühli (University of Zurich), Dr. Nadja Tomoum, Prof. Dr. Adrian V. Jäggi (University of Tübingen), Dr. med. Patrick Eppenberger (University of Tübingen)

And for setting up the online tool:

Prof. Dr. Verena Schünemann, PD Dr. Nicole Bender, Dr. Anja Furtwängler

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