References for The Predatory Stresses of Hierarchy and Socioeconomic Inequality

References & Notes for:

The Predatory Stresses of Hierarchy and Socioeconomic Inequality

Please note the following: In the context of the interplay of genes and environments–unless the article states otherwise–cultural, historical, political, economic, natural and biophysical factors are viewed as dynamic and interwoven environmental conditions and not as isolated variables.

In addition, the use of the word ‘interplay’ in the context of ‘gene-environment interplay’ and other similar sentences is defined according the following article: Allegrini, A. G., Karhunen, V., Coleman, J. R., Selzam, S., Rimfeld, K., von Stumm, S., … & Plomin, R. (2020). Multivariable GE interplay in the prediction of educational achievement. PLoS Genetics, 16(11), e1009153.

“Quantitative genetic theory distinguishes two types of interplay between genetic and environmental effects, genotype-environment correlation (rGE) and genotype-environment interaction (GxE).”

1. Tilly, C. (2011). Cities, states, and trust networks: Chapter 1 of Cities and states in world history. In Contention and trust in cities and states (pp. 1-16). Springer Netherlands.

“No states existed anywhere in the world before 4000 BCE.”

“Cities, then, first appeared in the same periods and regions as states. Like cities, states can only exist in symbiosis with agriculture that produces enough to support significant non-agricultural populations. Cities differ from strictly agricultural settlements, furthermore, by virtue of substantial populations, differentiated and specialized activities, and location as nodes in far-reaching networks of trade and political coordination. Cities and states maintain ambivalent relations: urban merchants and intellectuals seek the protection that states can provide, but resist the extraction and control that states’ rulers impose on them. Rulers of states, on their side, commonly try to combat urbanites’ independence, but also seek to benefit from concentrations of resources in cities as well as from the relative defensibility of compact cities as compared with scattered rural populations.”

“What of the state? A state is a structure of power involving four distinctive elements: 1) major concentrated means of coercion, especially an army, 2) organization that is at least partly independent of kinship and religious relations, 3) a defined area of jurisdiction, and 4) priority in some regards over all other organizations operating within that area. Although the four elements had existed separately for some time, no one put all four of them together before the Middle East’s creation of both cities and states. No states existed anywhere in the world before 4000 BCE. By the era of Gilgamesh’s Uruk, however, full-fledged cities and states were flourishing across significant parts of the Middle East, and possibly forming in other parts of Eurasia as well.”

2. Pinker, S. (2012). The better angels of our nature: Why violence has declined. Penguin Books. Kindle version.

This citation is used to reinforce the point the first ‘true states’ and ‘civilizations’ emerged some 5000 years ago.

According to Pinker: “Archaeologists tell us that humans live in a state of anarchy until the emergence of civilization some five thousand years ago…” …location 1062

“It took around five thousand years after the origin of agriculture for true states to appear on the scene”26 …location 1206

The following are references from Pinker’s book concerning cities and states–Pinker’s Reference 26: First states: Diamond, 1977; Gat 2006; Kurthz, 2001; Otterbein, 2004.

3. Whenever social hierarchy is referred to, the reader should note there is a difference between ‘top-down hierarchy’ and ‘reverse dominance hierarchy’ and their evolutionary context.

In addition, whether top-down or bottom-up, social hierarchies are dominance structures.

Cultural anthropologist Christopher Boehm introduced the theory of ‘reverse dominance hierarchy.’^3a

According to Boehm, “Egalitarianism involves a very special type of hierarchy”^3b that “can stay in place only with the vigilant and active suppression of bullies, who as free riders could otherwise openly take what they wanted from others who were less selfish or less powerful.”^3c

Boehm writes, “Humans naturally form hierarchies when they live in groups.”^3d “In despotic social dominance hierarchies the pyramid of power is pointed upward with one or a few individuals…at the top. In egalitarian hierarchies, the pyramid of power is turned upside down, with a politically united rank and file dominating…”

3a. Boehm, Christopher. 1993. “Egalitarian Behavior and Reverse Dominance Hierarchy.” Current Anthropology, Vol. 34, No. 3, pp 227-254. Also see: Christopher Boehm. Hierarchy in the Forest: The Evolution of Egalitarian Behavior (Kindle Location 908).

3b. Boehm, Christopher. Hierarchy in the Forest: The Evolution of Egalitarian Behavior. Kindle Location 178-179.

3c. Boehm, Christopher (2012-05-01). Moral Origins: The Evolution of Virtue, Altruism, and Shame (p. 204). Basic Books. Kindle Edition.

3d. Boehm, Christopher. Hierarchy in the Forest: The Evolution of Egalitarian Behavior. Kindle Location 573.

4. Martin, C. L., Ghastine, L., & Lodge, E. K. (2022). Understanding health inequalities through the lens of social epigenetics. Annual Review of Public Health, 43, 235-254.

Willems, Y.E., Rezaki, A.D., Aikins, M. et al. (2026). Social determinants of health and epigenetic clocks: a systematic review and meta-analysis of 140 studies. Nat Hum Behav https://doi.org/10.1038/s41562-026-02477-6

T.W. McDade,C. Ryan,M.J. Jones,J.L. MacIsaac,A.M. Morin,J.M. Meyer,J.B. Borja,G.E. Miller,M.S. Kobor, & C.W. Kuzawa, (2017). Social and physical environments early in development predict DNA methylation of inflammatory genes in young adulthood, Proc. Natl. Acad. Sci. U.S.A. 114 (29) 7611-7616, https://doi.org/10.1073/pnas.1620661114

Aristizabal, M. J., Anreiter, I., Halldorsdottir, T., et al. (2020). Biological embedding of experience: A primer on epigenetics. Proceedings of the National Academy of Sciences, 117(38), 23261-23269.

Schmitz, L. L., et al. (2021). The Socioeconomic Gradient in Epigenetic Aging Clocks: Evidence from the Multi-Ethnic Study of Atherosclerosis and the Health and Retirement Study. Epigenetics, 16(7), 658-668.

Notterman, D. A., & Mitchell, C. (2015). Epigenetics and Understanding the Impact of Social Determinants of Health. Pediatric Clinics of North America, 62(5), 1227-1240.

Nusslock, R., & Miller, G. E. (2016). Early-Life Adversity and Physical and Emotional Health Across the Lifespan: A Neuroimmune Network Hypothesis. Biological Psychiatry, 80(1), 23-32.

Lappé, M., & Jeffries Hein, R. (2021). You are what your mother endured: Intergenerational epigenetics, early caregiving, and the temporal embedding of adversity. Medical Anthropology Quarterly, 35(4), 458–475.

Belsky, D. W., Caspi, A., Corcoran, D. L., Sugden, K., Poulton, R., Arseneault, L., … & Moffitt, T. E. (2022). DunedinPACE, a DNA methylation biomarker of the pace of aging. elife, 11, e73420.441.

Motsinger-Reif, A. A., Reif, D. M., Akhtari, F. S., House, J. S., Campbell, C. R., Messier, K. P., … & Woychik, R. (2024). Gene-environment interactions within a precision environmental health framework. Cell Genomics, 4(7).

5. Please note: …the inheritance of acquired epigenetic marks in humans remains challenged.

Allegrini, A. G., Karhunen, V., Coleman, J. R., Selzam, S., Rimfeld, K., von Stumm, S., Pingault, JB, Plomin, R. (2020). Multivariable GE interplay in the prediction of educational achievement. PLoS Genetics, 16(11), e1009153.

“…quantitative genetics findings indicate that measures of the environment are themselves heritable.” They continue: “Our findings have relevance for genomic and environmental prediction models alike, as they show the way in which individuals’ genetic predispositions and environmental effects are intertwined.”

Herrera-Luis, E., Benke, K., Volk, H., Ladd-Acosta, C., & Wojcik, G. L. (2024). Gene-environment interactions in human health. Nature Reviews Genetics, 25(11), 768–784.

Abdellaoui, A., Martin, H. C., Kolk, M., Rutherford, A., Muthukrishna, M., Tropf, F. C., … & Visscher, P. M. (2025). Socio-economic status is a social construct with heritable components and genetic consequences. Nature human behaviour, 9(5), 864-876.

Erola, J., Lehti, H., Baier, T., & Karhula, A. (2022). Socioeconomic background and gene–environment interplay in social stratification across the early life course. European Sociological Review, 38(1), 1-17.

Thayer, Z. M., & Kuzawa, C. W. (2011). Biological memories of past environments: Epigenetic pathways to health disparities. Epigenetics, 6(7), 798–803.

Meloni, M. (2015). Epigenetics for the social sciences: Justice, embodiment, and inheritance in the postgenomic age. New Genetics and Society, 34(4), 439–461.

Scorza, P., et al. (2019). Intergenerational transmission of disadvantage: Epigenetics and parents’ childhoods as the first exposure. Journal of Child Psychology and Psychiatry, 60(2), 119-132.

Rothstein, M. A., et al. (2017). Transgenerational epigenetics and environmental justice. Environmental Epigenetics, 3(3), dvx011.

Tucker-Drob, E. M., & Bates, T. C. (2016). Large cross-national differences in gene × socioeconomic status interaction on intelligence. Psychological Science, 27(2), 138–149.

McEwen, B. S., & Gianaros, P. J. (2011). Stress-and allostasis-induced brain plasticity. Annual review of medicine, 62(1), 431-445.

Hertzman, C. (2012). Putting the concept of biological embedding in a historical perspective. American Journal of Public Health, 102(10), 1843-1844.

Sapolsky, R. M. (2017). Behave: The biology of humans at our best and worst. Penguin Press.

Raffington, L., Tanksley, P. T., Sabhlok, A., et al. (2023). Socially stratified epigenetic profiles are associated with cognitive functioning in children and adolescents. Psychological Science, 34(2), 170–185.

7. Motsinger-Reif, A. A., Reif, D. M., Akhtari, F. S., House, J. S., Campbell, C. R., Messier, K. P., … & Woychik, R. (2024). Gene-environment interactions within a precision environmental health framework. Cell Genomics, 4(7).

8. von Stumm, S., & d’Apice, K. (2022). From genome-wide to environment-wide: Capturing the environome. Perspectives on Psychological Science, 17(1), 30–40.

“There is broad consensus that people’s differences in affect, behavior and cognition result from the interplay between genetic propensities and environmental conditions. However, the mechanisms and processes that drive this interplay are not yet well understood.”

“People select themselves into, adapt to, and shape the environments that correspond to their genotypes.”

“…most of people’s differences in affect, behavior and cognition are influenced by both genetic and environmental factors…”

Psychologist Sophie Von Stumm and Katrina d’Apice write: “…we conceive here the environome, akin to the genome, that encompasses all environmental influences that give rise to people’s differences in affect, behavior and cognition.”

9. Laland, K. N., Odling-Smee, J., & Feldman, M. W. (2000). Niche construction, biological evolution, and cultural change. Behavioral and Brain Sciences, 23(1), 131–146.

Saltz, J. B., & Nuzhdin, S. V. (2014). Genetic variation in niche construction: Implications for development and evolutionary genetics. Trends in Ecology & Evolution, 29(1), 8–14.

Laland, K. N., Uller, T., Feldman, M. W., Sterelny, K., Müller, G. B., Moczek, A., Jablonka, E., & Odling-Smee, J. (2015). The extended evolutionary synthesis: Its structure, assumptions and predictions. Proceedings of the Royal Society B: Biological Sciences, 282(1813), 20151019.

Odling-Smee, J., Erwin, D. H., Palkovacs, E. P., Feldman, M. W., & Laland, K. N. (2013). Niche construction theory: A practical guide for ecologists. The Quarterly Review of Biology, 88(1), 3–28.

10. Briley, D. A., Livengood, J., & Derringer, J. (2018). Behaviour genetic frameworks of causal reasoning for personality psychology. European Journal of Personality, 32(3), 202-220.

“…individuals actively create or select environmental experiences aligned with their genetically influenced preferences and desires.”

von Stumm, S., & d’Apice, K. (2022). From genome-wide to environment-wide: Capturing the environome. Perspectives on Psychological Science, 17(1), 30-40.

“People select themselves into, adapt to, and shape the environments that correspond to their genotypes.”

“…most of people’s differences in affect, behavior and cognition are influenced by both genetic and environmental factors…”

Beam, C. R., Turkheimer, E., Dickens, W. T., & Davis, D. W. (2015). Twin differentiation of cognitive ability through phenotype to environment transmission: The Louisville Twin Study. Behavior Genetics, 45, 622-634.

“People do not randomly select environments, but maneuver and position themselves into environments and milieus where they can thrive, as well as react to environments provided to them (e.g., by caregivers). That is, people select into certain environments (e.g., niches) to reinforce innate or learned abilities…”

Scarr, S., & McCartney, K. (1983). How people make their own environments: A theory of genotype → environment effects. Child Development, 54(2), 424-435.

Mann, F. D., et al. (2019). Social-relational exposures and well-being. Journal of Research in Personality, 83, 103880.

“…individuals are not randomly assigned to social-relational environments. Rather, individuals select into and evoke responses from environments based on their heritable characteristics.”

Avinun, R. (2020). The E is in the G. Perspectives on Psychological Science, 15(1), 81-89.

Hufer, A., Kornadt, A. E., Kandler, C., & Riemann, R. (2020). Genetic and environmental variation in political orientation in adolescence and early adulthood: A nuclear twin family analysis. Journal of Personality and Social Psychology, 118(4), 762–776. https://doi.org/10.1037/pspp0000258

11. Alexander, R. D. (1974). The evolution of social behavior. Annual Review of Ecology and Systematics, 5(1), 325–383.

“When man developed his weapons, culture, and population sizes to levels that essentially erased the significance of predators of other species, he simultaneously created a new predator: groups and coalitions within his own species.”

Cosmides, L. & Tooby, J. (2010). Groups in mind: The coalitional roots of war and morality. Human morality and sociality: Evolutionary and comparative perspectives (pp. 191-234).

Kurzban, R., Tooby, J., & Cosmides, L. (2001). Can race be erased? Coalitional computation and social categorization. Proceedings of the National Academy of Sciences, 98(26), 15387-15392.

Pietraszewski, D. (2016). How the mind sees coalitional and group conflict: The evolutionary invariances of n-person conflict dynamics. Evolution and Human Behavior, 37(6), 470-480.

12. Sidanius, J., & Pratto, F. (2001). Social dominance: An intergroup theory of social hierarchy and oppression. Cambridge University Press.

Tooby, J., & Cosmides, L. (2010). Groups in mind: The coalitional roots of war and morality. Human morality and sociality: Evolutionary and comparative perspectives, 191-234.

13. Sidanius, J., & Pratto, F. (2001). Social dominance: An intergroup theory of social hierarchy and oppression. Cambridge University Press.

Alexander, R. D. (1974). The evolution of social behavior. Annual review of ecology and systematics, 5(1), 325-383.

Tooby, J., & Cosmides, L. (2010). Groups in mind: The coalitional roots of war and morality. Human morality and sociality: Evolutionary and comparative perspectives, 191-234.

15. van de Werfhorst, H. G. (2026). The Sociogenomics of Social Stratification and General Theories of Inequality. The British journal of sociology. DOI: 10.1111/1468-4446.70138

16. Erola, J., Lehti, H., & Baier, T. (2022). Socioeconomic background and gene–environment interplay in social stratification across the early life course. European Sociological Review, 38(1), 1–17

Raffington, L., Tanksley, P. T., Sabhlok, A., Vinnik, L., Mallard, T., King, L. S., Goosby, B., Harden, K. P., & Tucker-Drob, E. M. (2023). Socially stratified epigenetic profiles are associated with cognitive functioning in children and adolescents. Psychological Science, 34(2), 170–185. https://doi.org/10.1177/09567976221122760

Castagné, R., Ménard, S., & Delpierre, C. (2023).The epigenome as a biological candidate to incorporate the social environment over the life course and generations. Epigenomics, 15(1), 7–10.

Schmitz, C., H. T., et al. (2021). The Socioeconomic Gradient in Epigenetic Aging Clocks: Evidence from the Multi-Ethnic Study of Atherosclerosis and the Health and Retirement Study. Epigenetics, 16(7), 658-668.

Cerutti, R. et al. (2021). Associations between indicators of socioeconomic position and DNA methylation: a scoping review. Clinical Epigenetics, 13, 221. https://doi.org/10.1186/s13148-021-01189-0

Petrovic, D., Carmeli, C., Sandoval, J. L., Bodinier, B., Chadeau-Hyam, M., Schrempft, S., … & Stringhini, S. (2023). Life-course socioeconomic factors are associated with markers of epigenetic aging in a population-based study. Psychoneuroendocrinology, 147, 105976.

17. Abdellaoui, A., Martin, H. C., Kolk, M., Rutherford, A., Muthukrishna, M., Tropf, F. C., … & Visscher, P. M. (2025). Socio-economic status is a social construct with heritable components and genetic consequences. Nature human behaviour, 9(5), 864-876.

Belsky, Jay. (2024). The nature of nurture: Darwinian and mendelian perspectives. Development and Psychopathology: 1-10.

Kong, A., et al. (2018). The nature of nurture: Effects of parental genotypes. Science, 359(6374), 424-428.

Sauce, Bruno, and Louis D. Matzel. The paradox of intelligence: Heritability and malleability coexist in hidden gene-environment interplay. Psychological bulletin 144.1 (2018): 26.

“…because intelligence is demonstrably heritable, independent environmental effects cannot possibly run the show. This leads us to the conclusion that gene– environment interplay is the ring master.” “Even though the heritability of intelligence is high (at least in some populations), evidence from multiple lines of research suggests that variation in intelligence is greatly affected by normal environmental variation. In other words, one can say that IQ has a high heritability and a high malleability.”

Erola, Jani, et al. (2021). Socioeconomic Background and Gene–Environment Interplay in Social Stratification across the Early Life Course. European Sociological Review, 1 17.

“It is possible to test for the presence of gene-environment correlation, and one method to do this is using the twin design to estimate the heritability of environmental experiences. A systematic review of gene-environment correlation twin studies estimated that the average heritability of measures of the environment was as high as 27%.”

Kendler, Kenneth S., and Jessica H. Baker (2007). Genetic influences on measures of the environment: a systematic review. Psychological medicine 37.5: 615-626.

Bartels, M., de Geus, E. J. C., Vinkers, C. H., Bartels, M. (2022). Exploring the biological basis for happiness. In J. Blom, A. Tarnoki, D. Tarnoki, H. Harris, & N. Segal (Eds.), Twin Research for Everyone (pp. 105–126). Academic Press. doi:10.1016/B978-0-12-821514-2.00016-7

18. Scarr, S., & McCartney, K. (1983). How people make their own environments: A theory of genotype → environment effects. Child Development, 54(2), 424–435.

Plomin, R., DeFries, J. C., Knopik, V. S., & Neiderhiser, J. M. (2016). Top 10 replicated findings from behavioral genetics. Perspectives on Psychological Science, 11(1), 3–23.

Reinforces that sociability and many other behavioral traits are heritable and influence environmental selection.

19. Plomin, R., DeFries, J. C., Knopik, V. S., & Neiderhiser, J. M. (2016). Top 10 replicated findings from behavioral genetics. Perspectives on Psychological Science, 11(1), 3–23.

Genotype–Environment Correlation (rGE) is a central concept in their work. They outline the three types—passive, evocative, and active—where individuals’ genetically influenced traits affect the environments they experience. Environmental factors such as parenting and peer relationships are not purely external influences—they are often shaped by the child’s own genetically influenced traits. This is especially emphasized in their explanation of evocative and active rGE.

20. von Stumm, S., Smith-Woolley, E., Ayorech, Z., et al. (2020). Predicting educational achievement from genomic measures and socioeconomic status. Developmental Science, 23(3), e12925.

“…SES is often assumed to represent solely environmental advantages of wealth and privilege, but it is actually just as heritable as most other complex traits, with estimates from twin studies of about 50%. The main ingredients in most SES scores are parents’ educational attainment and occupational status, both of which are substantially heritable.”

21. Allegrini, A. G., Karhunen, V., Coleman, J. R., Selzam, S., Rimfeld, K., von Stumm, S., et al. (2020). Multivariable GE interplay in the prediction of educational achievement. PLoS Genetics, 16(11), e1009153.

22. Briley, D. A., Livengood, J., & Derringer, J. (2018). Behaviour genetic frameworks of causal reasoning for personality psychology. European Journal of Personality, 32(3), 202-220.

“…individuals actively create or select environmental experiences aligned with their genetically influenced preferences and desires.” “People select themselves into, adapt to, and shape the environments that correspond to their genotypes.”

Avinun, R. (2020). The E is in the G: gene–environment–trait correlations and findings from Genome-Wide Association Studies. Perspectives on Psychological Science, 15(1), 81-89.

“Active rGE refers to instances in which individuals choose their environment (e.g. friends, activities) based on genetically influenced traits.”

Rimfeld, K., Ayorech, Z., Dale, P. S., Kovas, Y., Plomin, R. (2016). Genetics affects choice of academic subjects as well as achievement. Scientific reports, 6(1), 26373.

“The findings that DNA differences substantially affect differences in appetites as well as aptitudes suggest a genetic way of thinking about education in which individuals actively create their own educational experiences in part based on their genetic propensities.

Plomin, R. (2019). Blueprint: How DNA makes us who we are. MIT Press. Kindle version, pp. 100–101.

“We actively perceive, modify and even create environments correlated with our genetic propensities.”

24. Cavalli, G., & Heard, E. (2019). Advances in epigenetics link genetics to the environment and disease. Nature, 571(7766), 489-499.

Aristizabal, M. J., Anreiter, I., Halldorsdottir, T., Odgers, C. L., McDade, T. W., Goldenberg, A., … & O’Donnell, K. J. (2020). Biological embedding of experience: A primer on epigenetics. Proceedings of the National Academy of Sciences, 117(38), 23261-23269.

Heijmans, B. T., Tobi, E. W., Stein, A. D., Putter, H., Blauw, G. J., Susser, E. S., Lumey, L. H., & Slagboom, P. E. (2008). Persistent epigenetic differences associated with prenatal exposure to famine in humans. Proceedings of the National Academy of Sciences, 105(44), 17046–17049.

25. Aristizabal, M. J., Anreiter, I., Halldorsdottir, T., Odgers, C. L., McDade, T. W., Goldenberg, A., … & O’Donnell, K. J. (2020). Biological embedding of experience: A primer on epigenetics. Proceedings of the National Academy of Sciences, 117(38), 23261-23269.

Cecil, C. A., Zhang, Y., & Nolte, T. (2020). Childhood maltreatment and DNA methylation: A systematic review. Neuroscience & Biobehavioral Reviews, 112, 392-409.

26. Bush, N. R., Edgar, R. D., Park, M., et al. (2018). The biological embedding of early-life socioeconomic status and family adversity in children’s genome-wide DNA methylation. Epigenomics, 10(11), 1445–1461.

27. Kuzawa, C. W., & Sweet, E. (2009). Epigenetics and the embodiment of race: Developmental origins of US racial disparities in cardiovascular health. American Journal of Human Biology, 21(1), 2-15.

Gkiouleka, M., Karalexi, M., Sergentanis, T. N., Nouvakis, D., Proikaki, S., Kornarou, E., & Vassilakou, T. (2025). The Epigenetic Role of Nutrition Among Children and Adolescents: A Systematic Literature Review. Children, 12(2), 143.

Scorza, P., et al. (2019). Research Review: Intergenerational transmission of disadvantage: epigenetics and parents’ childhoods as the first exposure. Journal of Child Psychology and Psychiatry, 60(2), 119-132.

Bowers, M. E., & Yehuda, R. (2016). Intergenerational transmission of stress in humans. Neuropsychopharmacology, 41(1), 232-244.

Holuka, C., Grova, N., Charalambous, E. G., Le Cléac’H, J., Turner, J. D., & Mposhi, A. (2024). Transgenerational impacts of early life adversity: from health determinants, implications to epigenetic consequences. Neuroscience & Biobehavioral Reviews, 164, 105785.

Zhou, A., & Ryan, J. (2023). Biological embedding of early-life adversity and a scoping review of the evidence for intergenerational epigenetic transmission of stress and trauma in humans. Genes, 14(8), 1639.

Conching, A. K. S., & Thayer, Z. (2019). Biological pathways for historical trauma to affect health: A conceptual model focusing on epigenetic modifications. Social Science & Medicine, 230, 74-82.

Chervova, O., Panteleeva, K., Chernysheva, E., Widayati, T. A., Baronik, Ž. F., Hrbková, N., … & Voloshin, V. (2024). Breaking new ground on human health and well-being with epigenetic clocks: A systematic review and meta-analysis of epigenetic age acceleration associations. Ageing Research Reviews, 102, 102552.

DiMarzio, K., Rojo-Wissar, D. M., Hernandez Valencia, E., Ver Pault, M., Denherder, S., Lopez, A., … & Parent, J. (2025). Childhood adversity and adolescent epigenetic age acceleration: the role of adolescent sleep health. Sleep Advances, 6(1), zpaf003.

Mozhui, K., Starlard-Davenport, A., Sun, Y., Shadyab, A. H., Casanova, R., Thomas, F., … & Johnson, K. C. (2026). Epigenetic entropy, socioeconomic differences, and health and lifespan in the Women’s Health Initiative. Clinical Epigenetics.

Hamlat, E. J., Mayer, S. E., Laraia, B., Moffitt, T. E., Surachman, A., Dutcher, E. G., … & Epel, E. S. (2025). Maternal childhood adversity accelerates epigenetic aging of children. Health Psychology, 44(5), 479.

Dieckmann, L., & Czamara, D. (2024). Epigenetics of prenatal stress in humans: the current research landscape. Clinical Epigenetics, 16(1), 20.

28. Fiorito, G., Polidoro, S., Dugué, P. A., Kivimaki, M., Ponzi, E., Matullo, G., … & Vineis, P. (2017). Social adversity and epigenetic aging: a multi-cohort study on socioeconomic differences in peripheral blood DNA methylation. Sci Rep 7, 16266 (2017). https://doi.org/10.1038/

Schmitz, L. L., et al. (2021). The socioeconomic gradient in epigenetic aging clocks: Evidence from the Multi-Ethnic Study of Atherosclerosis and the Health and Retirement Study. Epigenetics, 16(7), 658–668.

29. Willems, Y.E., Rezaki, A.D., Aikins, M. et al. (2026). Social determinants of health and epigenetic clocks: a systematic review and meta-analysis of 140 studies. Nat Hum Behav. https://doi.org/10.1038/s41562-026-02477-6

30. Slavich, G. M., Mengelkoch, S., & Cole, S. W. (2023). Human social genomics: Concepts, mechanisms, and implications for health. Lifestyle Medicine, 4(2), e75.

Cole, S. W. (2014). Human social genomics. PLoS Genetics, 10(8), e1004601

31. von Stumm, S., Kandaswamy, R., & Maxwell, J. (2022). Gene–environment interplay in early life cognitive development. Intelligence, 98, 101748

“Children’s differences in early life cognitive development are driven by the interplay of genetic and environmental factors.”

“By the time they start formal education, children’s differences in cognitive ability are powerful predictors of their contemporaneous and future academic achievement.”

32. Choi, S. W., Mak, T. S. H., & O’Reilly, P. F. (2020). Tutorial: a guide to performing polygenic risk score analyses. Nature protocols, 15(9), 2759-2772.

33. Kweon, H., Burik, C., Karlsson Linnér, R., De Vlaming, R., Okbay, A., Martschenko, D., … & Koellinger, P. (2020). Genetic fortune: Winning or losing education, income, and health. SSRN Electronic Journal. https://doi.org/10.2139/ssrn.3682041

34. Cavalli, G., & Heard, E. (2019). Advances in epigenetics link genetics to the environment and disease. Nature, 571(7766), 489-499.

Belsky, D. W., Caspi, A., Arseneault, L., Baccarelli, A. A., Corcoran, D. L., Gao, X., … & Moffitt, T. E. (2020). Quantification of the pace of biological aging in humans through a blood test, the DunedinPoAm DNA methylation algorithm. eLife, 9, e54870.

Dunn, E. C., Soare, T. W., Zhu, Y., Simpkin, A. J., Suderman, M. J., Klengel, T., … & Relton, C. L. (2019). Sensitive periods for the effect of childhood adversity on DNA methylation: results from a prospective, longitudinal study. Biological Psychiatry, 85(10), 838-849.

Swartz, J. R., Hariri, A. R., & Williamson, D. E. (2017). An epigenetic mechanism links socioeconomic status to changes in depression-related brain function in high-risk adolescents. Molecular Psychiatry, 22(2), 209-214.

Keller, M., Vogel, M., Garten, A., Svensson, S. I. A., Rossi, E., Kovacs, P., … & Kiess, W. (2025). Epigenetics of childhood obesity. Hormone Research in Paediatrics

Harden, K. P., & Koellinger, P. D. (2020). Using genetics for social science. Nature Human Behaviour, 4(6), 567-576.

Schuurmans, I. K., Smajlagic, D., Baltramonaityte, V., Malmberg, A. L., Neumann, A., Creasey, N., … & Cecil, C. A. (2025). Genetic susceptibility to neurodevelopmental conditions associates with neonatal DNA methylation patterns in the general population: an individual participant data meta-analysis. Biological Psychiatry.

Dolcini, J., Chiavarini, M., Firmani, G., Brennan, K. J., Cardenas, A., Baccarelli, A. A., & Barbadoro, P. (2025). Methylation biomarkers of lung Cancer risk: A systematic review and Meta-Analysis. Cancers, 17(4), 690.

Bartels, M., van de Weijer, M. P., Azcona Granada, N., Baselmans, B. M., Suderman, M., Soerensen, M., … & van Dongen, J. (2025). Epigenome-wide association study meta-analysis of wellbeing. bioRxiv, 2025-09.

Richardson, T. G., Zheng, J., Davey Smith, G., Timpson, N. J., Gaunt, T. R., Relton, C. L., & Hemani, G. (2017). Mendelian randomization analysis identifies CpG sites as putative mediators for genetic influences on cardiovascular disease risk. American Journal of Human Genetics, 101(4), 590-602.

Hannon, E., Dempster, E., Viana, J., Burrage, J., Smith, A. R., Macdonald, R., … & Mill, J. (2016). An integrated genetic-epigenetic analysis of schizophrenia: Evidence for co-localization of genetic associations and differential DNA methylation. Genome Biology, 17(1), 1-16.

Wawrzczak‐Bargieła, A., Bilecki, W., & Maćkowiak, M. (2023). Epigenetic targets in schizophrenia development and therapy. Brain Sciences, 13(3), 426.

35. Mavrommatis, C., Belsky, D. W., Ying, K., Moqri, M., Campbell, A., Richmond, A., … & Marioni, R. E. (2025). An unbiased comparison of 14 epigenetic clocks in relation to 174 incident disease outcomes. Nature communications, 16(1), 11164.

Kuo, P. L., Moore, A. Z., Tanaka, T., Belsky, D. W., Lu, A. T. H., Horvath, S., … & Ferrucci, L. (2026). Longitudinal changes in epigenetic clocks predict survival in the InCHIANTI cohort. Nature Aging, 1-7.

36. Heijmans, B., Tobi, E., Stein, A., Putter, H., Blauw, G., Susser, E., … & Lumey, L. (2008). Persistent epigenetic differences associated with prenatal exposure to famine in humans. Proceedings of the National Academy of Sciences, 105(44), 17046-17049.

Eichenauer, H. and Ehlert, U. (2023). The association between prenatal famine, dna methylation and mental disorders: a systematic review and meta-analysis. Clinical Epigenetics, 15(1).

Bind, M. A., Lepeule, J., Zanobetti, A., Gasparrini, A., Baccarelli, A. A., Coull, B. A., … & Schwartz, J. (2014). Air pollution and gene-specific methylation in the Normative Aging Study: Association, effect modification, and mediation analysis. Epigenetics, 9(3), 448-458.

Cosin-Tomas, M., Hoang, T., Qi, C., Monasso, G. S., Langdon, R., Merid, S. K., … & Bustamante, M. (2025). Association of exposure to second-hand smoke during childhood with blood DNA methylation. Environment international, 195, 109204.

Plusquin, M., Guida, F., Polidoro, S., Vermeulen, R., Raaschou-Nielsen, O., Campanella, G., … & Chadeau-Hyam, M. (2017). DNA methylation and exposure to ambient air pollution in two prospective cohorts. Environment International, 108, 127-136.

Rider, C. F., & Carlsten, C. (2019). Air pollution and DNA methylation: Effects of exposure in humans. Clinical Epigenetics, 11(1), 131.

Vineis, P., Robinson, O., Chadeau-Hyam, M., Dehghan, A., Mudway, I., & Dagnino, S. (2020). What is new in the exposome? Environment International, 143, 10588737.

Tobi, E. W., Slieker, R. C., Luijk, R., Dekkers, K. F., Stein, A. D., Xu, K. M., … & Heijmans, B. T. (2018). DNA methylation as a mediator of the association between prenatal adversity and risk factors for metabolic disease in adulthood. Science Advances, 4(1), eaao4364.

37. Conching, A. K. S., & Thayer, Z. (2019). Biological pathways for historical trauma to affect health: A conceptual model focusing on epigenetic modifications. Social Science & Medicine, 230, 74-82.

Newell, M. E., Aravindan, A., Babbrah, A., & Halden, R. U. (2025). Epigenetic Biomarkers Driven by Environmental Toxins Associated with Alzheimer’s Disease, Parkinson’s Disease, and Amyotrophic Lateral Sclerosis in the United States: A Systematic Review. Toxics, 13(2), 114.

Juster, R., McEwen, B., & Lupien, S. (2010). Allostatic load biomarkers of chronic stress and impact on health and cognition. Neuroscience & Biobehavioral Reviews, 35(1), 2-16.

Gallego-Fabrega, C., Cullell, N., & Fernández-Cadenas, I. (2025). How epigenetics impacts stroke risk and outcomes through DNA methylation: A systematic review. Journal of Cerebral Blood Flow & Metabolism, 0271678X251322032. https://doi.org/10.1177/0271678X251322032

Chambers, J., Roscoe, C. M., Chidley, C., Wisniewska, A., & Duggirala, A. (2025). Molecular Effects of Physical Activity and Body Composition: A Systematic Review and Meta-Analysis. International Journal of Environmental Research and Public Health, 22(4), 637

Prisco, S. Z., Umar, S., Potus, F., & Agrawal, V. (2025). Exploring the role of epigenetic modifications in pulmonary vascular disease pathogenesis. Frontiers in Medicine, 12, 1618278.

Willems, Y.E., Rezaki, A.D., Aikins, M. et al. (2026). Social determinants of health and epigenetic clocks: a systematic review and meta-analysis of 140 studies. Nat Hum Behav. https://doi.org/10.1038/s41562-026-02477-6

Perera, F., & Herbstman, J. (2011). Prenatal environmental exposures, epigenetics, and disease. Reproductive Toxicology, 31(3), 363–373.

Thayer, Z. M., & Kuzawa, C. W. (2011). Biological memories of past environments: Epigenetic pathways to health disparities. Epigenetics, 6(7), 798–803.

Rothstein, M. A., et al. (2017). Transgenerational epigenetics and environmental justice. Environmental Epigenetics, 3(3), dvx011.

38. Deaton, A. (2013). The Great Escape: Health, Wealth, and the Origins of Inequality. Princeton University Press.

39. Stiglitz, J. E. (2016). Inequality and economic growth. Political Quarterly, 86(S1), 134-155.

Bowles, S., & Gintis, H. (2002). The inheritance of inequality. Journal of Economic Perspectives, 16(3), 3-30.

Hornborg, A. (2024). Beyond the veil of market prices: The implications of ecologically unequal exchange for conceptualizations of technological development. Contributions to Political Economy, 43(1).

Beckert, J. (2009) The social order of markets. Theory and Society, 38(3), 245–269.

Piketty, T., & Saez, E. (2014). Inequality in the long run. Science, 344(6186), 838–843.

40. Diesendorf, M., Davies, G., & Wiedmann, T. (2024). Sustainability scientists’ critique of neoclassical economics. Global Sustainability, Cambridge University Press.

Cook, E. (2020). Naturalizing Inequality: The Problem of Economic Fatalism in the Age of Piketty. Capitalism: A Journal of History and Economics, 1(2), 338-378.

Spash, C. L. (2020). A Tale of Three Paradigms: Realising the Revolutionary Potential of Ecological Economics. Ecological Economics, 169.

Bowles, S., & Gintis, H. (2000). Walrasian economics in retrospect. The Quarterly Journal of Economics, 115(4), 1411-1439.

41. Amis, J., Munir, K., & Mair, J. (2017). Institutions and economic inequality. Research in the Sociology of Organizations, 52, 3–34.

42. Intergovernmental Panel on Climate Change. (2021). Climate change 2021: The physical science basis. Contribution of Working Group I to the Sixth Assessment Report. Cambridge University Press.

43. Kenner, D. (2019). Carbon inequality: The role of the richest in climate change. Routledge.

Wiedmann, T., Lenzen, M., Keyßer, L. T., & Steinberger, J. K. (2020). Scientists’ warning on affluence. Nature Communications, 11(1), 3107.

Otto, I. M., Kim, K. M., Dubrovsky, N., & Lucht, W. (2019). Shift the focus from the super-poor to the super-rich. Nature Climate Change, 9(2), 82-84.

Oxfam. (2023). Climate equality: A planet for the 99%. Oxfam International.

44. Piketty, T. (2014). Capital in the twenty-first century. Harvard University Press.

45. Chancel, L. (2022). Global carbon inequality over 1990–2019. Nature Sustainability, 5(11), 931–938.

Diffenbaugh, N. S., & Burke, M. (2019). Global warming has increased global economic inequality. Proceedings of the National Academy of Sciences, 116(20), 9808-9813.

Schöngart, S., Nicholls, Z., Hoffmann, R., Pelz, S., & Schleussner, C.-F. (2025). High-income groups disproportionately contribute to climate extremes worldwide. Nature Climate Change. Volume 15, Issue 6, pages 627-633

46. Clark, B. & York, R. (2005). Carbon metabolism: Global capitalism, climate change, and the biospheric rift. Theory and Society, 34(4), 391-428.

Hickel, J., & Kallis, G. (2020). Is green growth possible? New Political Economy, 25(4), 469-486.

Lamb, W. F., Wiedmann, T., Pongratz, J., Andrew, R., Crippa, M., Olivier, J. G., … & Minx, J. C. (2021). A review of trends and drivers of greenhouse gas emissions by sector from 1990 to 2018. Environmental Research Letters, 16(7), 073005.

47. Tooze, A. (2014). The deluge: The Great War, America and the remaking of the global order, 1916-1931. Viking Press.

Beckert, S. (2014). Empire of cotton: A global history. Alfred A. Knopf.

Malm, A. (2016). Fossil capital: The rise of steam power and the roots of global warming. Verso Books.

48. Nielsen, K. S., Nicholas, K. A., Creutzig, F., Dietz, T., & Stern, P. C. (2021). The role of high-socioeconomic-status people in locking in or rapidly reducing energy-driven greenhouse gas emissions. Nature Energy, 6(11), 1011–1016.

49. Wiedmann, T., Lenzen, M., Keyßer, L. T., & Steinberger, J. K. (2020). Scientists’ warning on affluence. Nature Communications, 11, 3107.

50. Ripple, W. J., Wolf, C., Gregg, J. W., Rockström, J., Newsome, T. M., Law, B. E., … & King, S. D. A. (2023). The 2023 state of the climate report: Entering uncharted territory. BioScience, 73(12), 841–850.

51. World Bank. (2022, December 8). Half of the global population lives on less than US$6.85 per person per day. World Bank Blogs. https://blogs.worldbank.org/en/developmenttalk/half-global-population-lives-less-us685-person-day

52. Ritchie, H., et al. (2023). Population Growth. Our World in Data. https://ourworldindata.org/population-growth

53. Shonkoff, J. P., Garner, A. S., Committee on Psychosocial Aspects of Child and Family Health, & Committee on Early Childhood, Adoption, and Dependent Care. (2012). The lifelong effects of early childhood adversity and toxic stress. Pediatrics, 129(1), e232-e246.

Hertzman, C. (2012). Putting the concept of biological embedding in a historical perspective. Proceedings of the National Academy of Sciences, 109 (Supplement 2), 17160-17167.

Chetty, R., Grusky, D., Hell, M., Hendren, N., Manduca, R., & Narang, J. (2017). The fading American dream: Trends in absolute income mobility since 1940. Science, 356(6336), 398-406.

Chetty, R., Hendren, N., Kline, P., & Saez, E. (2014). Where is the land of opportunity? The geography of intergenerational mobility in the United States. Quarterly Journal of Economics, 129(4), 1553-1623.

Blanden, J., Doepke, M., & Stuhler, J. (2022). Educational inequality. In Handbook of Labor, Human Resources and Population Economics (pp. 1-40). Springer.

Pager, D., & Shepherd, H. (2008). The sociology of discrimination: Racial discrimination in employment, housing, credit, and consumer markets. Annual Review of Sociology, 34, 181-209.

Harris, J., & Nisbett, N. (2020). The basic determinants of malnutrition: resources, structures, ideas and power. International Journal of Health Policy and Management, 10(12), 817.

Delisle, H. (2008). Poverty: The double burden of malnutrition in mothers and the intergenerational impact. Annals of the New York Academy of Sciences, 1136(1), 172-184.

Solt, F. (2008). Economic inequality and democratic political engagement. American Journal of Political Science, 52(1), 48-60.

Krieger, N. (2012). Methods for the scientific study of discrimination and health: An ecosocial approach. American Journal of Public Health, 102(5), 936–944.

Marmot, M. (2005). Social determinants of health inequalities. The Lancet, 365(9464), 1099–1104.

54. World Bank. (2023). World Development Indicators: Population total. Data.worldbank.org

55. Our World in Data. (2023). Daily calorie supply per person. Retrieved from ourworldindata.org/grapher/food-supply-kcal

56. Our World in Data. (2023). Life expectancy. Retrieved from ourworldindata.org/grapher/life-expectancy

57. Ritchie, H. (2017). How much of the world’s land would we need in order to feed the global population with the average diet of a given country? Our World in Data.

Based on 2011 data from 161 countries.

58. Chetty, R., Stepner, M., Abraham, S., Lin, S., Scuderi, B., Turner, N., et al. (2016). The association between income and life expectancy in the United States, 2001–2014. JAMA, 315(16), 1750–1766.

59. Deaton, A. (2016). On Death and Money, Commentary on “The association between income and life expectancy…” JAMA.

Also see:

Deaton, A. (2006). Global patterns of income and health: Facts, interpretations, and policies. National Bureau of Economic Research Working Paper No. 12735.

Deaton, A. (2013). The Great Escape: Health, wealth, and the origins of inequality. Princeton University Press. (Kindle Edition)

60. Papanicolas, I., Woskie, L. R., & Jha, A. K. (2018). Health care spending in the United States and other high-income countries. JAMA, 319(10), 1024–1039.

61. Emanuel, E. J., et al. (2021). Comparing health outcomes of privileged US citizens with those of average residents of other developed countries. JAMA Internal Medicine, 181(3), 339–344.

62. OECD. (2019). Health at a glance 2019: United States – How does it compare? World Health Systems Facts. https://healthsystemsfacts.org/the-us-health-system/

63. ‘System-justified harms–harms that function as social and moral goods’–is introduced in the article The Gene-Environment Roots of Inequality.

64. Knafo-Noam, A. (2015). The developmental and genetic architecture of prosociality. Current Opinion in Psychology, 6, 60-64.

Gelfand, M. J., Raver, J. L., Nishii, L., Leslie, L. M., Lun, J., Lim, B. C., … & Yamaguchi, S. (2011). Differences between tight and loose cultures. A 33-nation study. Science, 332(6033), 1100-1104

65. The ‘Jack-in-the-Box’ Pattern: Development and Influences

The following sources provide a range of historical, structural, and socio-genetic context for the concept…demonstrating the many dimensions of Jack.

I. Core Historical & Cyclical Influences

These works establish the recurring nature of hierarchical reset through violent “leveling” events and elite-driven disintegration cycles.

Scheidel, W. (2017). The Great Leveler: Violence and the history of inequality from the Stone Age to the twenty-first century. Princeton University Press.

Turchin, P., & Nefedov, S. A. (2009). Secular cycles. Princeton University Press.

Turchin, P. (2023). End times: Elites, counter-elites, and the path of political disintegration. Penguin Press.

Wallerstein, I. (1974-1989). The Modern World-System (Vols. 1-3). Academic Press.

II. Structural Mechanisms of Systemic Failure

These sources identify the mechanical “tension” and specific vulnerabilities that lead to a failure or a reset of the hierarchy.

Scheidel, W. (2024). Beyond technology and wages: Power and the history of inequality. Oxford Open Economics, 3 (Supplement_1), 1212–1216.

Turchin, P., Currie, T. E., Whitehouse, H., François, P., Feeney, K., Mullins, D., … & Spencer, C. (2018). Quantitative historical analysis uncovers a single dimension of complexity that structures global variation in human social organization. Proceedings of the National Academy of Sciences, 115(2), E144-E151.

Piketty, T. (2014). Capital in the twenty-first century (A. Goldhammer, Trans.). Belknap Press of Harvard University Press.

Tainter, J. A. (1988). The collapse of complex societies. Cambridge University Press.

Kennedy, P. (1987). The rise and fall of the great powers: Economic change and military conflict from 1500 to 2000. Random House.

III. The “Jack-in-the-Box” Replacement Dynamics

Khaldun, I. (2015). The Muqaddimah: An introduction to history (N. J. Dawood, Ed.; F. Rosenthal, Trans.). Princeton University Press.

Describes Asabiyyah (social cohesion) as the social force that allows a new group to replace a failing elite.

IV. Socio-Genetic & Genomic Evidence

These references provide evidence in the archaeological record of large-scale population change and the hereditary transmission of status.

Theory & Mechanism:

Plomin, R., & von Stumm, S. (2018). The new genetics of intelligence. Nature Reviews Genetics, 19(3), 148–159.

Polderman, Tinca JC, et al. “Meta-analysis of the heritability of human traits based on fifty years of twin studies.” Nature genetics 47.7 (2015): 702-709.

Causadias, J. M., & Korous, K. M. (2017). How are genes related to culture? An introduction to the field of cultural genomics. The handbook of culture and biology, 151-177.

Boyce, W. T., Sokolowski, M. B., & Robinson, G. E. (2020). Genes and environments, development and time. Proceedings of the National Academy of Sciences (PNAS), 117(38), 23235–23241.

Population Turnover & Heritable Status:

Žegarac, A., Winkelbach, L., Blöcher, J., Diekmann, Y., Krečković Gavrilović, M., Porčić, M., … & Burger, J. (2021). Ancient genomes provide insights into family structure and the heredity of social status in the early Bronze Age of southeastern Europe. Scientific Reports, 11(1), 10072.

Allentoft, M.E., Sikora, M., Refoyo-Martínez, A. et al. Population genomics of post-glacial western Eurasia. Nature 625, 301–311 (2024). https://doi.org/10.1038/s41586-023-06865-0

Gretzinger, J., et al. (2022). The Anglo-Saxon migration and the formation of the early English gene pool. Nature, 610, 112–119.

66. Cannadine, D. (1999). The rise and fall of class in Britain. Columbia University Press.

Jump to:

References & Notes for:

The Predatory Stresses of Hierarchy and Socioeconomic Inequality