Raise Your Hand if You’re a Dumbass A/K/A Tinbergen’s Four Questions

About our guest

David Sloan Wilson is an evolutionary biologist, a founder of modern multilevel selection theory, and president of Prosocial World. Books mentioned this episode: This View of Life (2019), Evolution for Everyone (2007) and Darwin’s Cathedral (2002).

Awards

Award	Winner	Stat	For
Best Question	Olin	191	Longest average answer to their questions (words)
Lexicon Award	David	0.671	Most varied vocabulary
Signal-to-Noise Award	David	11.7	Most references per 1,000 words
Laugh Track Award	David	6	Most laughs triggered
Longest Monologue	Olin	302	Longest uninterrupted stretch (words)
Four Questions Evangelist	Andy	16	Said “four questions” the most
Filler King	Andy	5.5	Most filler words per 100 words
The Filibuster Award	Andy	166.6	Most words per point scored
Translator of the Episode	Andy	Judges’ pick	For turning the science into plain English

MVP scoring

Category	David	Andy	Olin
References (+1 each)	62	21	11
Long-answer questions (+2 each)	6	4	6
Callbacks received (+1 each)	13	4	8
Laughs (+1 each)	6	5	1
Vocabulary rank (+3/+2/+1)	3	1	2
Filler penalty (−1 per filler per 100 words)	-2	-5.5	-4.2
Corrections (−5 each)	-10	-5	-5
References per 1,000 words (+1 each)	11.7	4.2	6.0
Blab score (benchmark: 100 words per point)	3.4	-6.7	0.8
MVP score	93.1	22.0	25.6
Place	Gold	Bronze	Silver

How it’s scored: MVP = +1 per reference · +2 per question that drew a 150+ word answer · +1 per callback received · +1 per laugh · +3/+2/+1 for vocabulary rank · −1 per filler per 100 words · −5 per correction · +1 per reference per 1,000 words · +1 for every 10 words per point under 100 (−1 for every 10 over)

Scoring inputs

Input	David	Andy	Olin	How it was measured
Questions that drew a 150+ word answer	3	2	3	Questions followed by 150+ words from the others before the asker spoke again
Callbacks received	13	4	8	”Back to your point,” “Olin’s question,” “great point,” etc., said by someone else
Laughs triggered	6	5	1	Laughter from someone else within two turns
Vocabulary variety (MATTR)	0.671	0.643	0.668	Unique words per rolling 100-word window; fair across amounts of talking
Filler words per 100 words	2	5.5	4.2	”like,” “yeah,” “you know,” “I mean,” “kind of/sort of,” “literally,” “basically”
Words spoken	5,310	4,999	1,834	From the auto-generated transcript (approximate)
Average answer to your questions (words)	47	38	191
Longest uninterrupted stretch (words)	258	261	302
”Four questions” said	10	16	11
References raised	62	21	11	From the scoreboard above
Corrections	2	1	1	From the scoreboard above
Words per point (blab index)	65.6	166.6	91.7	Words spoken ÷ (references + callbacks + laughs). Benchmark: 100. Lower is better.
References per 1,000 words	11.7	4.2	6.0	References raised ÷ words spoken × 1,000

Raise your hand (corrections)

Who	Time	What was said	What the record says	Counts?
David	41:02	”Something like 500,000 generations”	The E. coli experiment is at about 82,000 generations.	Yes
David	1:19:39	”All mammals” have seven neck vertebrae	Nearly all do; sloths and manatees are the exceptions.	Yes
Andy	32:06	Mayr’s proximate/ultimate in “the early fifties”	It was 1961, and David corrected it on air at 32:19.	Yes
Olin	54:55	The W.C. Fields quote	Versions of the joke go back to at least 1936, with many claimants.	Yes
Olin	23:03	Proteins and fats as the proximate reason food tastes good	Judges’ call: that’s the ultimate explanation, as Andy pointed out.	No

More context

• 1:14:51: “Religion” from the Latin “to bind” (religare) is one classical etymology, from Lactantius. Cicero derived it from relegere, “to go over again.”
• 25:42: Eaton et al. (1988) call it ‘discordance’; Neel’s 1962 thrifty genotype is the classic early mismatch argument.
• 57:55–58:01: The song is M.I.A.’s ‘Paper Planes’ (plural). The cocktail was created in 2008.

References raised on air and corrections, by host

Host	References raised	Corrections
David	62	2
Andy	21	1
Olin	11	1

By the numbers

Reference counts

Stat	Value
References	94	Works cited in this episode
Scientific / scholarly	73	Peer-reviewed papers and scholarly books
Years of scholarship spanned	2,069	From Cicero (45 BCE) to 2024
Authors credited across all references	451	108 of them on a single paper (Evershed et al., 2022)
Works by David Sloan Wilson	11	Our guest, cited by us, not by himself
PNAS papers	6
Science papers	4
Nature / Nature Genetics papers	3	Not counting the author correction
Nobel laureates discussed	3	von Frisch, Lorenz, and Tinbergen (1973)
Bible passages	4

Transcript stats

Final tallies for Season 1, Episode 1. Word counts come from the auto-generated final transcript, so treat them as approximate.

Stat	Total	David	Andy	Olin	Note
Words spoken	12,143	5,310	4,999	1,834	David 44% · Andy 41% · Olin 15%
Times speaking	437	136	191	110
Longest uninterrupted stretch (words)		258	261	302	Olin’s four-questions explainer (12:55) wins
”Four questions”	37	10	16	11
”Tinbergen” (any spelling)	23	12	6	5
”Evolution” / “evolutionary”	47	24	17	6
”Multilevel”	7	2	3	2	In a podcast called Everything Is MLS
”Religion” / “religious”	26	23	1	2
”Darwin”	14	12	2	0
”Like”	206	36	133	37
”Amazing”	12	2	10	0
”Dumbass”	3	1	2	0	Including the closing line
F-bombs	7	1	5	1
Cocktails on air (one drink break)	6	2	2	2	Paper Planes (Andy), The Other Words (Olin), gin and tonics (David)
Ingredients per drink		2	4	4	Two equal-parts, four-ingredient cocktails for a four-questions episode, plus a G&T

Where the references come from

Category	Count	Share
Scientific / scholarly	73	78%
Popular / web	13	14%
Historical / literary text	8	9%
Total	94	100%

Popular / web = popular-audience books, websites, and media. Historical / literary = scripture, classical texts, Calvin’s catechism, and Gulliver’s Travels.

Span of references

	Year	Reference
Oldest reference	45 BCE	Cicero (45 BCE)
Newest reference	2024	Birkhead & Schulze-Hagen (2024)
Oldest scientific reference	1871	Darwin (1871)
Newest scientific reference	2024	Birkhead & Schulze-Hagen (2024)
Years spanned	2,069
Median year (all dated references)	1991

References by period

Period	All	Scientific / scholarly	Popular / web	Historical / literary	Share
Pre-1900	6	2	0	4	6%
1900–1950	6	6	0	0	6%
1951–1999	37	33	4	0	39%
2000s (2000–2009)	12	11	1	0	13%
2010s (2010–2019)	18	12	6	0	19%
2020s (2020–)	9	9	0	0	10%
Undated (scripture, web pages)	6	0	2	4	6%
Total	94	73	13	8	100%

References by role

Role	Count
Background	17
Critique / counterpoint	4
Mentioned on-air	4
Optional	1
Original source	43
Original source (reprint)	1
Supporting evidence	10
Update / critique	1
Update / recent work	9
Visual / web	4

References

All 94 works cited, grouped by topic. Link goes to the publisher or DOI; Free goes to an open or legal free copy where one exists.

Cold open: why everything is MLS

01:31–06:41 · Olin, Andy — The hosts introduce the show, David Sloan Wilson, and the idea that understanding evolution properly means understanding multilevel selection.

• Wilson, D. S. (1975). A theory of group selection. Proceedings of the National Academy of Sciences, 72(1), 143–146. Link · Free
  • Introduced the trait-group model, placing selection within and among groups on a single continuum: the founding paper of modern multilevel selection theory. (03:16, raised by Andy)
• Darwin, C. (1871). The Descent of Man, and Selection in Relation to Sex. John Murray. Link · (free)
  • Darwin’s early argument that groups with more cooperative members prevail over other groups, an intuition multilevel selection theory later made rigorous. (03:16, raised by Andy)
• Reflections on Papers Past. (2017). Revisiting Wilson 1975. Reflections on Papers Past (blog). Link · (free)
  • David Sloan Wilson recounts how his 1975 model began, prompted by a paper on the vertical migration of zooplankton. (04:22, raised by Andy)

Tinbergen’s four questions

05:15–06:41; 11:19–22:14 · Olin, David, Andy — Function, mechanism, development, and evolutionary history: the four complementary questions that frame the whole episode.

• Tinbergen, N. (1963). On aims and methods of ethology. Zeitschrift für Tierpsychologie, 20(4), 410–433. Link
  • Proposed four complementary questions (survival value, causation, development, and evolutionary history) for fully explaining any behavior. (05:15–06:41; 11:19–22:14, raised by Olin)
• Tinbergen, N. (2005). On aims and methods of ethology. Animal Biology, 55(4), 297–321.
  • Facsimile reprint of the 1963 paper that established the four questions. (11:19–22:14, raised by Olin)
• Bateson, P., & Laland, K. N. (2013). Tinbergen’s four questions: An appreciation and an update. Trends in Ecology & Evolution, 28(12), 712–718. Link
  • A fifty-year reappraisal arguing that the four questions remain essential, with updates for modern biology. (11:19–22:14, raised by David)
• Bergman, T. J., & Beehner, J. C. (2022). Leveling with Tinbergen: Four levels simplified to causes and consequences. Evolutionary Anthropology, 31(1), 12–19. Link · Free
  • Arranges the four questions on a timescale of causes and consequences, clarifying how they complement one another. (11:19–22:14, raised by David)

Niko Tinbergen (biography)

16:49 · David — Who Tinbergen was: a Dutch pioneer of animal behavior, born in The Hague.

• Kruuk, H. (2003). Niko’s Nature: The Life of Niko Tinbergen and His Science of Animal Behaviour. Oxford University Press.
  • The definitive biography of Niko Tinbergen, from Dutch naturalist to Nobel laureate at Oxford. (16:49, raised by David)
• Hinde, R. A. (1990). Nikolaas Tinbergen, 15 April 1907 – 21 December 1988. Biographical Memoirs of Fellows of the Royal Society, 36, 547–565.
  • The Royal Society’s memoir of Tinbergen’s life and science. (16:49, raised by David)
• Nobel Prize Outreach. (1973). Nikolaas Tinbergen – Biographical. NobelPrize.org. Link · (free)
  • Tinbergen’s own account of his life, written on receiving the Nobel Prize. (16:49, raised by David)

Ethology: definition and origin

17:05–18:16 · David, Andy — Ethology is the study of animal behavior; the key insight was that behaviors evolve just like physical traits.

• Jaynes, J. (1969). The historical origins of “ethology” and “comparative psychology”. Animal Behaviour, 17(4), 601–606.
  • Traces the term ‘ethology’ from its 19th-century French coinage to the modern study of animal behavior. (17:05–18:16, raised by David)
• Wheeler, W. M. (1902). “Natural history,” “oecology” or “ethology”? Science, 15(390), 971–976. (citation details being verified)
  • Popularized ‘ethology’ in English as the name for the scientific study of animal behavior. (17:05–18:16, raised by David)
• Burkhardt, R. W. (2005). Patterns of Behavior: Konrad Lorenz, Niko Tinbergen, and the Founding of Ethology. University of Chicago Press.
  • How Lorenz and Tinbergen built ethology into a science of behavior as an evolved trait. (17:05–18:55, raised by David)

The 1973 Nobel Prize

18:55 · David — Von Frisch (the bee dance), Lorenz (imprinting), and Tinbergen shared the 1973 Nobel Prize for founding ethology.

• Nobel Prize Outreach. (1973). The Nobel Prize in Physiology or Medicine 1973. NobelPrize.org. Link · (free)
  • Awarded jointly to von Frisch, Lorenz, and Tinbergen for discoveries on how individual and social behavior patterns are organized and elicited. (18:55, raised by David)
• von Frisch, K. (1967). The dance language and orientation of bees (L. E. Chadwick, Trans.). Belknap Press of Harvard University Press. (Original work published 1965)
  • Von Frisch’s synthesis showing that honeybees communicate the location of food through dance. (18:55, raised by David)
• Lorenz, K. (1935). Der Kumpan in der Umwelt des Vogels. Der Artgenosse als auslösendes Moment sozialer Verhaltungsweisen. Journal für Ornithologie, 83, 137–213, 289–413. Link
  • Lorenz’s foundational study of imprinting and social behavior in birds. (18:55, raised by David)
• Tinbergen, N. (1951). The Study of Instinct. Clarendon Press.
  • Tinbergen’s classic synthesis of early ethology and the causes of instinctive behavior. (18:55, raised by David)
• Birkhead, T., & Schulze-Hagen, K. (2024). A new foundation for the study of bird behaviour: Konrad Lorenz’s “Kumpan” paper of 1935. Journal of Ornithology, 165(1), 5–14. Link · Free
  • An English-language guide to Lorenz’s 1935 ‘Kumpan’ paper and its lasting influence. (18:55, raised by David)

Mayr: proximate vs. ultimate causation; founder effects

18:55–27:16; 32:19; 1:02:14 · David, Andy, Olin — Mayr’s 1961 split between ‘how’ (proximate) and ‘why’ (ultimate) causes, plus his founder principle.

• Mayr, E. (1961). Cause and effect in biology: Kinds of causes, predictability, and teleology are viewed by a practicing biologist. Science, 134(3489), 1501–1506. Link
  • Distinguished proximate (‘how’) from ultimate (‘why’) causes in biology, a distinction that parallels Tinbergen’s four questions. (18:55; 22:14–27:16; 32:19, raised by Andy)
• Laland, K. N., et al. (2011). Cause and effect in biology revisited: Is Mayr’s proximate-ultimate dichotomy still useful? Science, 334(6062), 1512–1516. Link · Free
  • Argues that Mayr’s proximate–ultimate split obscures how development and behavior can themselves shape evolution. (18:55–27:16, raised by David)
• Mayr, E. (1942). Systematics and the Origin of Species. Columbia University Press.
  • Introduced the founder principle: a population started by a few individuals carries only a chance sample of its parent population’s variation. (1:02:14, raised by Andy)
• Mayr, E. (1954). Change of genetic environment and evolution. In J. Huxley, A. C. Hardy, & E. B. Ford (Eds.), Evolution as a Process (pp. 157–180). Allen and Unwin.
  • Developed founder effects into ‘genetic revolutions’ that can drive rapid divergence of isolated populations. (1:02:14, raised by Andy)

Evolutionary mismatch

22:14–26:38 · Andy, David — Why ice cream tastes so good: traits built for past environments can misfire in new ones.

• Neel, J. V. (1962). Diabetes mellitus: A “thrifty” genotype rendered detrimental by “progress”? American Journal of Human Genetics, 14(4), 353–362. Link · (free)
  • The ‘thrifty genotype’: genes that favor fat storage under scarcity become harmful amid modern abundance. (22:14–25:42, raised by Andy)
• Eaton, S. B., Konner, M., & Shostak, M. (1988). Stone agers in the fast lane: Chronic degenerative diseases in evolutionary perspective. American Journal of Medicine, 84(4), 739–749. Link · (free)
  • Argued that chronic ‘diseases of civilization’ reflect a discordance between Stone Age biology and modern life. (25:42–26:38, raised by David)
• Bowlby, J. (1969). Attachment and Loss: Vol. 1. Attachment. Basic Books. (citation details being verified)
  • Introduced the ‘environment of evolutionary adaptedness,’ the conditions to which a trait is adapted. (26:05, raised by David)
• Griffiths, P. E., & Bourrat, P. (2023). Integrating evolutionary, developmental and physiological mismatch. Evolution, Medicine, and Public Health, 11(1), 277–286. Link · Free
  • Reviews the history of the mismatch concept and integrates its evolutionary, developmental, and physiological forms. (25:42–26:38, raised by David)
• Li, N. P., van Vugt, M., & Colarelli, S. M. (2018). The evolutionary mismatch hypothesis: Implications for psychological science. Current Directions in Psychological Science, 27(1), 38–44. Link
  • Applies evolutionary mismatch to psychology: adaptive minds misfiring in novel environments. (25:42–26:38, raised by David)
• Sih, A., Ferrari, M. C. O., & Harris, D. J. (2011). Evolution and behavioural responses to human-induced rapid environmental change. Evolutionary Applications, 4(2), 367–387. Link
  • Reviews how rapid human-induced environmental change leaves many species’ behavior mismatched to new conditions. (26:24, raised by David)

Dobzhansky and the fruit fly

28:07–29:13; 35:09–37:17 · David — ‘Nothing in biology makes sense except in the light of evolution’, and the irony of a model organism studied without its ecology.

• Dobzhansky, T. (1973). Nothing in biology makes sense except in the light of evolution. The American Biology Teacher, 35(3), 125–129. Link · Free
  • Argued that evolution is the unifying principle without which biology makes no sense. (28:07–29:13, raised by David)
• Dobzhansky, T. (1964). Biology, molecular and organismic. American Zoologist, 4(4), 443–452.
  • The first appearance of Dobzhansky’s famous line, in his presidential address to the American Society of Zoologists. (28:07–29:13, raised by David)
• Dobzhansky, T., & Pavlovsky, O. (1957). An experimental study of interaction between genetic drift and natural selection. Evolution, 11(3), 311–319. Link
  • An experiment on how genetic drift and natural selection interact in small founding populations of fruit flies. (35:09–37:17, raised by David)
• Markow, T. A. (2015). The secret lives of Drosophila flies. eLife, 4, e06793. Link · Free
  • Argues that after a century of lab work, the natural history and ecology of Drosophila remain largely unknown. (35:09–37:17, raised by David)

E. coli Long-Term Evolution Experiment

40:20–44:03 · David, Andy, Olin — Twelve identical E. coli lines: the same function evolved in all of them, but by different mechanisms and along diverging histories.

• Lenski, R. E., et al. (1991). Long-term experimental evolution in Escherichia coli. I. Adaptation and divergence during 2,000 generations. The American Naturalist, 138(6), 1315–1341.
  • Launched the Long-Term Evolution Experiment: twelve identical E. coli populations evolving in parallel. (40:20–44:03, raised by David)
• Lenski, R. E., & Travisano, M. (1994). Dynamics of adaptation and diversification: A 10,000-generation experiment with bacterial populations. Proceedings of the National Academy of Sciences, 91(15), 6808–6814. Link · Free
  • After 10,000 generations, all twelve populations improved similarly in fitness yet diverged in other traits. (42:58–43:30, raised by David)
• Blount, Z. D., Borland, C. Z., & Lenski, R. E. (2008). Historical contingency and the evolution of a key innovation in an experimental population of Escherichia coli. Proceedings of the National Academy of Sciences, 105(23), 7899–7906. Link
  • One population evolved to use citrate only after earlier ‘potentiating’ mutations, direct evidence of historical contingency. (43:30–44:03, raised by David)

Path dependence / historical contingency

45:30–45:37 · Olin, David — Early chance events set the path for everything that follows.

• David, P. A. (1985). Clio and the economics of QWERTY. The American Economic Review, 75(2), 332–337.
  • The QWERTY keyboard as a case study in path dependence: early accidents can lock in lasting outcomes. (45:30–45:37, raised by Olin)
• Arthur, W. B. (1989). Competing technologies, increasing returns, and lock-in by historical events. The Economic Journal, 99(394), 116–131.
  • Modeled how small chance events, amplified by increasing returns, lock in one technology over its rivals. (45:30–45:37, raised by Olin)
• Liebowitz, S. J., & Margolis, S. E. (1990). The fable of the keys. The Journal of Law and Economics, 33(1), 1–25.
  • Challenged the QWERTY story, questioning whether inferior standards are really locked in. (45:30–45:37, raised by Olin)
• Gould, S. J. (1989). Wonderful Life: The Burgess Shale and the Nature of History. W. W. Norton.
  • Argued that ‘replaying the tape of life’ would yield a different history: evolution as deeply contingent. (45:30–45:37, raised by Olin)

Lactose tolerance (gene-culture coevolution)

45:42–51:08 · David, Andy, Olin — Dairying came first, then genes for adult milk digestion, with different mutations in different places.

• Simoons, F. J. (1969). Primary adult lactose intolerance and the milking habit: A problem in biological and cultural interrelations. I. Review of the medical research. American Journal of Digestive Diseases, 14(12), 819–836. Link
  • Reviewed medical evidence that most adults worldwide cannot digest lactose. (45:54–46:55, raised by David)
• Simoons, F. J. (1970). Primary adult lactose intolerance and the milking habit: A problem in biologic and cultural interrelations. II. A culture historical hypothesis. American Journal of Digestive Diseases, 15(8), 695–710. Link
  • Proposed that adult lactose tolerance spread after, and because of, dairying: culture first, genes second. (45:54–46:55, raised by David)
• McCracken, R. D. (1971). Lactase deficiency: An example of dietary evolution. Current Anthropology, 12(4/5), 479–517. Link
  • Framed adult lactose tolerance as an example of dietary evolution in human populations. (45:54–46:55, raised by David)
• Itan, Y., et al. (2010). A worldwide correlation of lactase persistence phenotype and genotypes. BMC Evolutionary Biology, 10, 36. Link
  • Compiled global data on lactase persistence and mapped its worldwide distribution. (48:56–50:12, raised by David)
• Tishkoff, S. A., et al. (2007). Convergent adaptation of human lactase persistence in Africa and Europe. Nature Genetics, 39(1), 31–40. Link
  • Showed that lactase persistence evolved independently, through different mutations, in African and European herders. (46:55, raised by David)
• Enattah, N. S., et al. (2002). Identification of a variant associated with adult-type hypolactasia. Nature Genetics, 30(2), 233–237. Link
  • Identified the main genetic variant associated with adult lactase persistence in Europeans. (46:55, raised by David)
• Evershed, R. P., et al. (2022). Dairying, diseases and the evolution of lactase persistence in Europe. Nature, 608(7922), 336–345. Link
  • Found that prehistoric milk use does not explain the spread of lactase persistence; famine and disease likely drove selection. (45:42–51:08, raised by David)
• Evershed, R. P., et al. (2022). Author Correction: Dairying, diseases and the evolution of lactase persistence in Europe. Nature, 609(7927), E9. Link
  • Author correction to Evershed et al. (2022). (45:42–51:08, raised by David)
• Burger, J., et al. (2007). Absence of the lactase-persistence-associated allele in early Neolithic Europeans. Proceedings of the National Academy of Sciences, 104(10), 3736–3741. Link · Free
  • Ancient DNA showed that early Neolithic Europeans lacked the persistence variant: dairying came before the gene. (45:54–46:55, raised by David)
• Burger, J., et al. (2020). Low prevalence of lactase persistence in Bronze Age Europe indicates ongoing strong selection over the last 3,000 years. Current Biology, 30(21), 4307–4315.e13. Link
  • Lactase persistence was still rare in Bronze Age Europe, implying strong selection within the last 3,000 years. (45:42–51:08, raised by David)
• Gerbault, P., et al. (2011). Evolution of lactase persistence: An example of human niche construction. Philosophical Transactions of the Royal Society B, 366(1566), 863–877. Link
  • Presents lactase persistence as human niche construction, with culture reshaping selection on genes. (45:42–51:08, raised by David)
• Durham, W. H. (1991). Coevolution: Genes, Culture, and Human Diversity. Stanford University Press.
  • A classic account of gene–culture coevolution, including the case of lactose absorption. (45:42–51:08, raised by David)
• Obinu, D. A., et al. (2010). Prevalence of lactase persistence and the performance of a non-invasive genetic test in adult Sardinian patients. e-SPEN, the European e-Journal of Clinical Nutrition and Metabolism, 5(1), e1–e5. Link
  • Found lactase-persistence variants to be rare among Sardinian adults. (50:12, raised by Olin)
• Schirru, E., et al. (2007). Decline of lactase activity and C/T-13910 variant in Sardinian childhood. Journal of Pediatric Gastroenterology and Nutrition, 45(4), 503–506.
  • Found that most Sardinian children lose lactase activity by around age nine. (50:12, raised by Olin)
• Mead, J. S. (2017). Talking Evolution: Are We Still Evolving? National Geographic Education Blog. Link · (free)
  • Source of the lactose intolerance map shown in this episode (map by Jerome M. Cookson, National Geographic). (48:56–50:12, raised by Andy)
• Howard Hughes Medical Institute. (n.d.). The making of the fittest: Got lactase? The co-evolution of genes and culture [Short film]. HHMI BioInteractive. Link
  • A short film on the coevolution of lactase persistence and dairying. (45:42–51:08, raised by Andy)

Darwin’s Cathedral

1:02:14–1:22:31 · Andy, David — David’s book treating religion, and Calvin’s Geneva in particular, as a group-level adaptation.

• Wilson, D. S. (2002). Darwin’s Cathedral: Evolution, Religion, and the Nature of Society. University of Chicago Press.
  • Treats religion as a group-level adaptation, with Calvin’s Geneva as the central case study. (1:02:14–1:22:31, raised by Andy)

Andy’s dissertation: cultural founder effects

1:02:14 · Andy — Applying Mayr’s founder effect to culture: implicit Puritanism in American moral cognition.

• Poehlman, T. A. (2007). Ideological Inheritance: Implicit Puritanism in American Moral Cognition [Doctoral dissertation, Yale University].
  • Proposed a cultural founder effect: Puritan values implicitly shape American moral cognition today. (1:02:14, raised by Andy)
• Uhlmann, E. L., et al. (2011). Implicit Puritanism in American moral cognition. Journal of Experimental Social Psychology, 47(2), 312–320. Link
  • Reported that Americans implicitly link work and sexual restraint to morality, consistent with a Puritan cultural inheritance. (1:02:14, raised by Andy)
• Tierney, W., et al. (2021). A creative destruction approach to replication: Implicit work and sex morality across cultures. Journal of Experimental Social Psychology, 93, 104060. Link
  • A multi-nation replication testing the implicit Puritanism effects across cultures. (1:02:14, raised by Andy)

Symbotype

1:18:28 · Andy — David’s term for a group’s network of symbols, which works like a genotype for culture.

• Wilson, D. S., et al. (2014). Evolving the future: Toward a science of intentional change. Behavioral and Brain Sciences, 37(4), 395–416. Link · Free
  • Coined ‘symbotype’ for the networks of symbols that guide human behavior, much as a genotype guides development. (1:18:28, raised by Andy)
• Wilson, D. S., et al. (2023). Multilevel cultural evolution: From new theory to practical applications. Proceedings of the National Academy of Sciences, 120(16), e2218222120. Link · Free
  • Applies multilevel cultural evolution to real-world change, including how symbolic systems are inherited and transformed. (1:18:28, raised by Andy)

David’s books mentioned on-air

10:20; 40:56 · David, Olin — Evolution for Everyone and This View of Life.

• Wilson, D. S. (2007). Evolution for Everyone: How Darwin’s Theory Can Change the Way We Think About Our Lives. Delacorte Press.
  • David Sloan Wilson’s case that evolutionary thinking can illuminate everyday life, for everyone. (10:20, raised by David)
• Wilson, D. S. (2019). This View of Life: Completing the Darwinian Revolution. Pantheon Books. (citation details being verified)
  • David Sloan Wilson’s call to complete the Darwinian revolution by applying evolution to human affairs. (40:56, raised by Olin)

Huxley: ‘How extremely stupid not to have thought of that!’

10:25; 34:44 · David — T. H. Huxley’s reaction to the Origin of Species.

• Huxley, T. H. (1887). On the reception of the “Origin of Species”. In F. Darwin (Ed.), The Life and Letters of Charles Darwin, Including an Autobiographical Chapter (Vol. 2, pp. 179–204). John Murray. Link · (free)
  • Huxley recalls his reaction to the Origin of Species: ‘How extremely stupid not to have thought of that!’ (10:25; 34:44, raised by David)

Mendel, the ‘eclipse of Darwinism’, and the modern synthesis

27:22–28:07 · David — Why it took until the 1940s for genetics and Darwinism to come together.

• Bowler, P. J. (1983). The Eclipse of Darwinism: Anti-Darwinian Evolution Theories in the Decades Around 1900. Johns Hopkins University Press.
  • The decades around 1900 when rival theories eclipsed Darwinian natural selection. (27:22–28:07, raised by David)
• Huxley, J. (1942). Evolution: The modern synthesis. George Allen & Unwin. (Definitive edition reissued 2010, MIT Press)
  • Named and synthesized ‘the modern synthesis’ of Mendelian genetics and natural selection. (28:07, raised by David)

Two bird brains: caching vs. star navigation

37:50 · David — Why you can’t understand a brain without knowing the animal’s ecology.

• Krebs, J. R., et al. (1989). Hippocampal specialization of food-storing birds. Proceedings of the National Academy of Sciences, 86(4), 1388–1392. Link · Free
  • Food-storing birds have a larger hippocampus than non-storers: brain structure shaped by ecology. (37:50, raised by David)
• Emlen, S. T. (1970). Celestial rotation: Its importance in the development of migratory orientation. Science, 170(3963), 1198–1201.
  • Young indigo buntings learn a star compass from the rotation of the night sky. (37:50, raised by David)

‘Not everything is adaptive’

21:37; 37:50; 1:11:19 · David — Byproducts, drift, and mismatch: the reasons a trait might exist without being an adaptation.

• Gould, S. J., & Lewontin, R. C. (1979). The spandrels of San Marco and the Panglossian paradigm: A critique of the adaptationist programme. Proceedings of the Royal Society of London. Series B, Biological Sciences, 205(1161), 581–598. Link
  • Warned against assuming every trait is an adaptation, highlighting byproducts and constraints. (21:37; 37:50; 1:11:19, raised by David)

Design principles for cooperative groups

51:38–53:27 · David — Ostrom’s core design principles, generalized through evolutionary theory.

• Ostrom, E. (1990). Governing the Commons: The Evolution of Institutions for Collective Action. Cambridge University Press.
  • Identified design principles that let communities govern shared resources without collapse. (51:38–53:27, raised by David)
• Wilson, D. S., Ostrom, E., & Cox, M. E. (2013). Generalizing the core design principles for the efficacy of groups. Journal of Economic Behavior & Organization, 90, S21–S32. Link · Free
  • Generalized Ostrom’s core design principles to groups of all kinds using multilevel selection theory. (51:38–53:27, raised by David)
• Atkins, P. W. B., Wilson, D. S., & Hayes, S. C. (2019). Prosocial: Using Evolutionary Science to Build Productive, Equitable, and Collaborative Groups. New Harbinger Publications.
  • Turns the core design principles into a practical method for building cooperative groups. (51:38–53:27, raised by David)

Multilevel selection and superorganisms

1:04:41–1:06:05 · David — Can a group be functionally organized like an organism? The podcast’s namesake idea.

• Wilson, D. S., & Sober, E. (1989). Reviving the superorganism. Journal of Theoretical Biology, 136(3), 337–356.
  • Revived the superorganism concept: groups become adaptive units when selection operates among them. (1:04:41–1:06:05, raised by David)
• Sober, E., & Wilson, D. S. (1998). Unto Others: The Evolution and Psychology of Unselfish Behavior. Harvard University Press.
  • The book-length case that selection operates at multiple levels, explaining genuine altruism in nature and in human psychology. (1:04:41–1:06:05, raised by David)
• Wilson, D. S., & Wilson, E. O. (2007). Rethinking the theoretical foundation of sociobiology. The Quarterly Review of Biology, 82(4), 327–348.
  • Rebuilt sociobiology on multilevel selection, showing how a more careful formulation overturned the 1960s rejection of group-level selection. (1:04:41–1:06:05, raised by David)

Holism and mid-century individualism

02:52; 1:07:03–1:08:01 · David — From Durkheim’s social organism to selfish genes and rational actors.

• Durkheim, É. (1995). The elementary forms of religious life (K. E. Fields, Trans.). Free Press. (Original work published 1912)
  • Argued that religion is fundamentally social, binding people into moral communities. (1:07:03, raised by David)
• Williams, G. C. (1966). Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought. Princeton University Press.
  • The influential critique of group selection that made gene-centered explanations the default for decades. (1:07:03–1:08:01, raised by David)
• Dawkins, R. (1976). The Selfish Gene. Oxford University Press.
  • Popularized the gene’s-eye view of evolution that dominated late-20th-century biology. (02:52; 1:07:03–1:08:01, raised by Andy)

‘Body of Christ’ and the etymology of ‘religion’

1:06:05–1:07:03; 1:14:51 · David, Andy — Religious communities describe themselves as one body, and ‘religion’ may derive from the Latin ‘to bind’.

• 1 Corinthians 12:12–27. New Revised Standard Version Updated Edition. Link
  • Paul’s image of the church as one body with many members. (1:06:52, raised by Andy)
• Romans 12:4–5. New Revised Standard Version Updated Edition. Link
  • Paul’s parallel image of many members forming one body. (1:06:52, raised by Andy)
• Colossians 1:18. New Revised Standard Version Updated Edition. Link
  • Christ as the head of the body, the church. (1:06:05, raised by David)
• Ephesians 4:15–16. New Revised Standard Version Updated Edition. Link
  • The body joined and knit together under Christ as its head. (1:06:05, raised by David)
• Cicero, M. T. (45 BCE). De natura deorum [On the nature of the gods], Book 2, §72. (citation details being verified)
  • Derived religio from relegere, ‘to go over again.’ (1:14:51, raised by David)
• Lactantius. (c. 311). Divinae institutiones [The divine institutes], Book 4, ch. 28. (citation details being verified)
  • Derived religio from religare, ‘to bind,’ the etymology cited in this episode. (1:14:51, raised by David)

Calvin’s Geneva Catechism

1:21:13 · David — The catechism as a cultural transmission device that ‘arranges the furniture of the mind’.

• Calvin, J. (1545). Catechism of the Church of Geneva. (French original 1542) (citation details being verified)
  • Calvin’s question-and-answer catechism, a tool for transmitting Geneva’s beliefs and norms. (1:21:13, raised by David)

The Axial Age

1:15:46 · David — The great religious and philosophical traditions of roughly 800–200 BCE.

• Jaspers, K. (1953). The origin and goal of history (M. Bullock, Trans.). Yale University Press. (Original work published 1949)
  • Coined the ‘Axial Age’ for the era (c. 800–200 BCE) when major world religions and philosophies emerged. (1:15:46, raised by David)
• Bellah, R. N. (2011). Religion in Human Evolution: From the Paleolithic to the Axial Age. Belknap Press of Harvard University Press.
  • Traces the evolution of religion from the Paleolithic through the Axial Age. (1:15:46, raised by David)

Seven neck vertebrae

1:19:39 · David — Some traits are locked in (even giraffes have seven neck vertebrae), just as core doctrines are in religions.

• Galis, F. (1999). Why do almost all mammals have seven cervical vertebrae? Developmental constraints, Hox genes, and cancer. Journal of Experimental Zoology (Molecular and Developmental Evolution), 285(1), 19–26.
  • Explains why nearly all mammals, giraffes included, have seven neck vertebrae: changing the number carries developmental costs. (1:19:39, raised by David)

Gulliver’s Travels

1:21:05 · David — Big-Endians vs. Little-Endians: disputes that look trivial but aren’t.

• Swift, J. (1726). Gulliver’s Travels. Benjamin Motte.
  • Satirizes a war between Big-Endians and Little-Endians over which end of an egg to break. (1:21:05, raised by David)

Moral foundations (Andy’s cancel culture research)

1:09:46 · Andy — Coding moral language in text with moral foundations dictionaries.

• Graham, J., Haidt, J., & Nosek, B. A. (2009). Liberals and conservatives rely on different sets of moral foundations. Journal of Personality and Social Psychology, 96(5), 1029–1046. Link
  • Showed that liberals and conservatives draw on different moral foundations, measured partly through moral language. (1:09:46, raised by Andy)
• Hopp, F. R., et al. (2021). The extended Moral Foundations Dictionary (eMFD): Development and applications of a crowd-sourced approach to extracting moral intuitions from text. Behavior Research Methods, 53(1), 232–246. Link
  • A crowd-sourced dictionary for detecting moral intuitions in text. (1:09:46, raised by Andy)

The cocktail segment

05:42 (preview); 53:27–1:00:31 · Andy, Olin, David — The Paper Plane, the Last Word, the Cocktail Codex, and a W.C. Fields quote that wasn’t his.

• Day, A., Fauchald, N., & Kaplan, D. (2018). Cocktail Codex: Fundamentals, Formulas, Evolutions. Ten Speed Press.
  • Organizes cocktails into a handful of root recipes and their variations: a phylogeny of drinks. (59:16, raised by Olin)
• Difford’s Guide. (n.d.). Paper Plane. Difford’s Guide. Link · (free)
  • The Paper Plane: Sam Ross’s equal-parts modern classic, a riff on the Last Word. (55:17–58:51, raised by Andy)
• Quote Investigator. (2010). Part Went for Liquor, Part for Women; The Rest I Spent Foolishly. Quote Investigator. Link · (free)
  • Traces the ‘half my money’ joke, often credited to W.C. Fields, back to at least 1936. (54:55, raised by Olin)

Music

• Intro: “Your Way to Tell Me Go” by Plastic Penny
• Outro: “Wants for Everyone” by Slippers

Credits

Everything Is MLS is Olin and Andy Poehlman, with David Sloan Wilson. Reference tracking and fact-checking by Andy Poehlman. All timestamps are from the full (YouTube) cut.