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6AANA050 Topics in Philosophy of Science – Philosophy of Biology
This course will look at three main topics: life and organisms, evolution by natural selection, and
molecular biology.
In the first part, we start by discussing what life is and why it’s so hard to define. We will also look at
what is an organism and the problem of biological individuality. Finally, we will look at biological
classification and the question of whether biological taxa are natural kinds.
In the second part, we will look at the logic of evolution by natural selection and the conditions for it
to occur: namely, if there is a population of things that vary among themselves, and this variation is
heritable, and correlates with differences in how many offspring they produce, then evolution by
natural selection always occurs. Organisms are said to be well adapted to their environments. We will
also look at what adaptation means and at different notions of function, and whether we should
think of functions in terms of purpose. We will address the question of what things other than
organisms can be targets of natural selection, discuss cases of evolutionary conflict and cooperation,
major transitions in evolution such as the evolution of multicellularity, and look at ways in which
evolution can sometimes lead to less-than-optimal outcomes.
In the third and final part of the course we will focus on philosophical issues concerning molecular
biology, genes, and information. We will address the question of what is a gene, whether there are
genes for certain features, and in what sense genes contain information. We will also talk about how
biology relates to, and differs from, other sciences such as physics and chemistry, and we will discuss
the issue of reductive explanations in genetics and molecular biology.
Detailed readings for each week are found below and will be available online. There is no textbook
for the course, but the following books are highly recommended.
PHILOSOPHY OF BIOLOGY
Godfrey-Smith, P. (2016) Philosophy of Biology. Princeton University Press.
Okasha, S. (2019). Philosophy of Biology: A Very Short Introduction. Oxford University Press.
Sober, E. (2024). The Philosophy of Evolutionary Theory: Concepts, Inferences, and Probabilities.
Cambridge University Press
Sterelny, K., & Griffiths, P. (1999) Sex and Death: An Introduction to Philosophy of Biology. University
of Chicago Press.
BIOLOGY
Nurse, P. (2020) What is Life David Fickling Books.
Pross, A. (2012) What is Life How Chemistry Becomes Biology. Oxford University Press.
Assessment: The course will be assessed by a two-hour examination in May, in which you will be
asked to answer two questions. There will also be one formative essay (maximum 2,500 words), the
deadline for which is 14:00 on Monday 25 February 2025. Please submit your formative essay by
emailing it to margarida.hermida@kcl.ac.uk before the deadline. Any extensions to formative
deadlines should be requested in advance by email.
PART I: Life and Organisms
1. What is life
What is life Why is life so hard to define Astrobiology and the N=1 problem. Universal biology. Life
as collective phenomenon vs. individual living things.
REQUIRED READINGS:
Cleland, C. E., & Chyba, C. F. (2002). Defining ‘life’. Origins of Life and Evolution of the Biosphere, 32,
387-393.
Schulze-Makuch, D., & Irwin, L. N. (2018). Life in the Universe: Expectations and constraints, third
edition. Springer. Chapter 2: Definition of life (pp. 7-26).
ADDITIONAL SUGGESTED READINGS:
Boden, M. A. (1999). Is metabolism necessary The British Journal for the Philosophy of Science,
50(2): 231-248.
Cockell, C. S. (2016). The similarity of life across the universe. Molecular Biology of the Cell, 27(10),
1553-1555.
Machery, E. (2012). Why I stopped worrying about the definition of life… and why you should as
well. Synthese, 185, 145-164.
Schrodinger, E. (1944). What is life The physical aspect of the living cell. Cambridge University Press.
Chapter 6: Order, disorder and entropy.
Ruiz-Mirazo, K., Peretó, J., & Moreno, A. (2004). A universal definition of life: autonomy and open ended evolution. Origins of Life and Evolution of the Biosphere, 34, 323-346.
Sterelny, K., & Griffiths, P. E. (1999). Sex and Death: An Introduction to Philosophy of Biology.
University of Chicago Press. Chapter 15: What is life (pp. 357-377).
SUGGESTED ESSAY QUESTIONS:
– What is life
– What does the N=1 problem mean for our understanding of life in the universe
– How does the life of a particular organism differ from the historical-collective phenomenon of life
2. Organisms and other biological individuals
The problem of biological individuality. Physiological and evolutionary notions of ‘organism’. Colonial
organisms. Organismality as a continuum.
REQUIRED READINGS:
Clarke, E. (2010). The problem of biological individuality. Biological Theory, 5(4), 312-325.
Okasha, S. (2022). On the very idea of biological individuality. The British Journal for the Philosophy
of Science, forthcoming.
ADDITIONAL SUGGESTED READINGS:
Godfrey-Smith, P. (2013). Darwinian Individuals. In F. Bouchard & P. Huneman (Eds.), From Groups to
Individuals: Evolution and Emerging Individuality (pp. 17-36). MIT Press.
Godfrey-Smith, P. (2014). Philosophy of Biology. Princeton University Press. Chapter 5: Individuals
(pp. 66-80).
Queller, D. C., & Strassmann, J. E. (2009). Beyond society: the evolution of organismality.
Philosophical Transactions of the Royal Society B: Biological Sciences, 364(1533), 3143-3155.
Olson, E. T. (2021) What is the problem of biological individuality In A. S. Meincke & J. Dupré (Eds.),
Biological Identity: Perspectives from Metaphysics and Philosophy of Biology (pp. 63-85). Routledge.
Pepper, J. W., & Herron, M. D. (2008). Does biology need an organism concept Biological Reviews,
83(4), 621-627.
Pradeu, T. (2016). Organisms or biological individuals Combining physiological and evolutionary
individuality. Biology & Philosophy, 31, 797-817.
Wilson, R. A., & Barker, M. J. (2024). Biological Individuals. In E. N. Zalta & U. Nodelman (Eds.), The
Stanford Encyclopedia of Philosophy (Spring 2024 Edition), URL =
.
SUGGESTED ESSAY QUESTIONS:
– What is the problem of biological individuality
– Not all organisms are Darwinian individuals, and not all Darwinian individuals are organisms.
Discuss.
– An organism is simply a unit with high cooperation and very low conflict among its parts. Discuss.
– What does it mean to say that some biological entities are more organismal than others
3. Natural kinds and biological classification
Biological classification. Essentialist and non-essentialist accounts of species and higher taxa. Are
biological taxa natural kinds Species as individuals.
REQUIRED READINGS:
Hull, D. L. (1976). Are species really individuals Systematic Zoology, 25(2), 174-191.
Okasha, S. (2002). Darwinian metaphysics: Species and the question of essentialism. Synthese, 131,
191-213.
ADDITIONAL SUGGESTED READINGS:
Devitt, M. (2008). Resurrecting biological essentialism. Philosophy of Science, 75(3), 344-382.
Godfrey-Smith, P. (2014). Philosophy of Biology. Princeton University Press. Chapter 7: Species and
the Tree of Life (pp. 100-119).
Okasha, S. (2019). Philosophy of Biology: A Very Short Introduction. Oxford University Press. Chapter
5: Species and Classification (pp. 63-82).
Reydon, T. A. C. (2019). Taxa hold little information about organisms: Some inferential problems in
biological systematics. History and Philosophy of the Life Sciences, 41(4):40.
Richards, R. A. (2016) Biological Classification: A Philosophical Introduction. Cambridge University
Press. Chapter 4: Evolutionary Taxonomy and the Cladistic Challenge (pp. 101-145).
Richards, R. A. (2016) Biological Classification: A Philosophical Introduction. Cambridge University
Press. Chapter 7: The Metaphysics of Biological Classification (pp. 209-242).
SUGGESTED ESSAY QUESTIONS:
– Are biological species natural kinds
– What determines whether two organisms belong to the same species
– Does phylogenetic systematics classify organisms into kinds
PART II: Evolution by Natural Selection
4. The logic of evolution by natural selection
Darwin’s theory of evolution by natural selection. Darwinian populations. Lewontin’s criteria of
evolution by natural selection.
REQUIRED READINGS:
Darwin, C. (1859). On the Origin of Species by means of Natural Selection. Chapter XIV:
Recapitulation and Conclusion (pp. 399-425).
Okasha, S. (2019). Philosophy of Biology: A Very Short Introduction. Oxford University Press. Chapter
2: Evolution and Natural Selection (pp. 9-26).
ADDITIONAL SUGGESTED READINGS:
Godfrey-Smith, P. (2009) Darwinian Populations and Natural Selection. Oxford University Press.
Chapter 2: Natural Selection and Its Representation (pp. 17-40); Chapter 3: Variation, Selection, and
Origins (pp. 41-67).
Godfrey-Smith, P. (2014). Philosophy of Biology. Princeton University Press. Chapter 3: Evolution and
Natural Selection (pp. 28-49).
Lewontin, R. C. (1970). The units of selection. Annual Review of Ecology and Systematics, 1: 1-18.
Sober, E. (2024). The Philosophy of Evolutionary Theory: Concepts, Inferences, and Probabilities.
Cambridge University Press. Chapter 1: A Darwinian Introduction (pp. 1-15); Chapter 2: Fitness and
Natural Selection (pp. 16-38).
SUGGESTED ESSAY QUESTIONS:
– How does evolution by natural selection work
– What are the conditions for evolution by natural selection to occur
– What does it mean to say that Darwin’s theory of evolution is variational but Lamarck’s is
transformational
5. Function, purpose, and adaptation
How to understand biological functions. Purpose and teleology. Adaptation and optimality.
REQUIRED READINGS:
Neander, K. (1991). The teleological notion of ‘function’. Australasian Journal of Philosophy, 69(4),
454-468.
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.
ADDITIONAL SUGGESTED READINGS:
Cummins, R. (1975) Functional Analysis. Journal of Philosophy, 72, 741-765.
Garson, J. (2019). What Biological Functions are and Why They Matter. Cambridge University Press.
Chapter 2: Function and Selection (pp. 25-42); Chapter 9: Function Pluralism (pp. 141-151).
Godfrey-Smith, P. (1993). Functions: Consensus Without Unity. Pacific Philosophical Quarterly, 74,
196-208.
Godfrey-Smith, P. (2001). Three Kinds of Adaptationism. In Hecht, O., & Sober, E. (Eds.),
Adaptationism and Optimality (pp. 335-357). Cambridge University Press.
Godfrey-Smith, P. (2014). Philosophy of Biology. Princeton University Press. Chapter 4: Adaptation,
Construction, Function (pp. 50-65).
Lewens, T. (2004). Organisms and Artifacts: Design in nature and elsewhere. MIT Press. Chapter 6:
Deflating Function (pp. 119-138).
Okasha, S. (2019). Philosophy of Biology: A Very Short Introduction. Oxford University Press. Chapter
3: Function and Adaptation (pp. 27-44).
Sober, E. (2024). The Philosophy of Evolutionary Theory: Concepts, Inferences, and Probabilities.
Cambridge University Press. Chapter 8: Adaptationism (pp. 206-229).
SUGGESTED ESSAY QUESTIONS:
– In what sense can organisms’ traits be said to have functions
– Should we understand biological functions in terms of their selected effects, their causal role, or
both
– Does it make sense to say that natural selection designs traits for certain purposes
– Are organisms always optimally adapted to their environments
6. Levels of selection
Levels of selection in evolution. The gene’s eye view. Group selection. Multi-level selection.
REQUIRED READINGS:
Williams, G. C. (1966) Adaptation and Natural Selection. Princeton University Press. Chapter 4:
Group Selection (pp. 92-124).
Sterelny, K., & Griffiths, P. E. (1999). Sex and Death: An Introduction to Philosophy of Biology.
University of Chicago Press. Chapter 3: The Gene’s Eye View of Evolution (pp. 55-76).
ADDITIONAL SUGGESTED READINGS:
gren, J. A. (2021). The Gene’s Eye View of Evolution. Oxford University Press. Chapter 3: Difficulties
of the Theory (pp. 80-115).
Dawkins, R. (1976/2016). The Selfish Gene (40th anniversary edition). Oxford University Press.
Chapter 2: The Replicators (pp 15-25); Chapter 3: Immortal Coils (pp. 26-58).
Lewontin, R. C. (1970). The units of selection. Annual Review of Ecology and Systematics, 1: 1-18.
Okasha, S. (2008). The units and levels of selection. A Companion to the Philosophy of Biology, 138-
156.
Okasha, S. (2019). Philosophy of Biology: A Very Short Introduction. Oxford University Press. Chapter
4: Levels of Selection (pp. 45-62).
Okasha, S. (2006). Evolution and the Levels of Selection. Oxford University Press. Chapter 2: Selection
at Multiple Levels: Concepts and Methods (pp. 40-59).
Sober, E. (2024). The Philosophy of Evolutionary Theory: Concepts, Inferences, and Probabilities.
Cambridge University Press. Chapter 3: Units of Selection (pp. 39-76).
SUGGESTED ESSAY QUESTIONS:
– What is the gene’s eye view of evolution, and what are the main objections to it
– What are the two types of group selection
– Natural selection can operate on multiple levels simultaneously. Discuss.
7. Evolutionary conflict and cooperation
Inter- and intra-organismic conflict. Evolutionary game theory. Cooperation and altruism. Inclusive
fitness. Major evolutionary transitions.
REQUIRED READINGS:
Smith, J. M., & Szathmary, E. (1997). The Major Transitions in Evolution. Oxford University Press.
Chapter 10: Intragenomic Conflict (pp. 171-185).
Bourke, A. F. G. (2011). Principles of Social Evolution. Oxford University Press. Chapter 3: The major
transitions in light of inclusive fitness theory (pp. 74-94).
ADDITIONAL SUGGESTED READINGS:
Bourke, A. F. G. (2011). Principles of Social Evolution. Oxford University Press. Chapter 2: A primer in
inclusive fitness theory (pp. 28-73).
Haig, D. (2019). Cooperation and conflict in human pregnancy. Current Biology, 29(11), R455-R458.
Furness, A. I., Morrison, K. R., Orr, T. J., Arendt, J. D., & Reznick, D. N. (2015). Reproductive mode and
the shifting arenas of evolutionary conflict. Annals of the New York Academy of Sciences, 1360(1),
75-100.
Okasha, S. (2024). Cancer and the levels of selection. The British Journal for the Philosophy of
Science, 75(3), 537-560.
Strassmann, J. E., Queller, D. C., Avise, J. C., & Ayala, F. J. (2011). In the light of evolution V:
Cooperation and conflict. Proceedings of the National Academy of Sciences, 108 (S2), 10787-10791.
SUGGESTED ESSAY QUESTIONS:
– What explains the evolution of altruistic behaviour in many organisms
– Genetic conflict explains some puzzling features of pregnancy. Discuss.
– How does the fact that eukaryotic cells contain both nuclear and mitochondrial DNA help to explain
traits such as sexual reproduction, mate selection, and junk DNA
8. Humans in evolutionary context
What, if anything, can evolution tell us about ourselves Is there such a thing as human nature The
nature vs. nurture debate. Evolutionary psychology and evolutionary medicine.
REQUIRED READINGS:
Nesse, R. M. (2012). Evolution: a basic science for medicine. In Poiani, A. (Ed.). Pragmatic Evolution
(pp. 107-114). Cambridge University Press.
Smith, S. E. (2020). Is evolutionary psychology possible Biological Theory, 15(1), 39-49.
ADDITIONAL SUGGESTED READINGS:
Lewens, T. (2012). Human nature: the very idea. Philosophy & Technology, 25, 459-474.
Machery, E. (2018). Doubling down on the nomological notion of human nature. In Hannon, E., &
Lewens, T. (Eds.), Why We Disagree About Human Nature (pp. 18-39). Oxford University Press.
Chisholm, J. S. (2012). Flourishing, feelings, and fitness: an evolutionary perspective on health
capability. In Poiani, A. (Ed.). Pragmatic Evolution (pp. 188-210). Cambridge University Press.
Sterelny, K., & Griffiths, P. E. (1999). Sex and Death: An Introduction to Philosophy of Biology.
University of Chicago Press. Chapter 13: From Sociobiology to Evolutionary Psychology (pp. 313-
336).
Okasha, S. (2019). Philosophy of Biology: A Very Short Introduction. Oxford University Press. Chapter
7: Human behaviour, mind, and culture (pp. 101-117).
SUGGESTED ESSAY QUESTIONS:
– How is evolution relevant for medicine
– Is there such a thing as human nature
– Can evolutionary mismatch explain the current obesity crisis
PART III: Genes and Molecular Biology
9. Genes and DNA
What is a gene Are there genes for certain features Does DNA literally carry information, and if so,
what kind of information The central dogma of molecular biology. Epigenetics and the interaction
between genes and environment.
REQUIRED READINGS:
Crick, F. (1970). Central dogma of molecular biology. Nature, 227(5258), 561-563.
Griffiths, P., & Stotz, K. (2013) Genetics and Philosophy: An Introduction. Cambridge University Press.
Chapter 6: The gene as information (pp. 143-180).
ADDITIONAL SUGGESTED READINGS:
Watson, J. D., & Crick, F. H. (1953). Molecular structure of nucleic acids: a structure for deoxyribose
nucleic acid. Nature, 171(4356), 737-738.
Godfrey-Smith, P. (2014). Philosophy of Biology. Princeton University Press. Chapter 6: Genes (pp.
81-99); Chapter 9: Information (pp. 144-157).
Griffiths, P., & Stotz, K. (2013) Genetics and Philosophy: An Introduction. Cambridge University Press.
Chapter 2: Mendel’s gene (pp. 9-32); Chapter 3: The material gene (pp. 33-65).
Okasha, S. (2019). Philosophy of Biology: A Very Short Introduction. Oxford University Press. Chapter
6: Genes (pp. 83-100).
Shea, N. (2007). Representation in the genome and in other inheritance systems. Biology &
Philosophy, 22, 313-331.
Maynard Smith, J. (2000). The concept of information in biology. Philosophy of Science, 67(2), 177-
194.
Sterelny, K., & Griffiths, P. E. (1999). Sex and Death: An Introduction to Philosophy of Biology.
University of Chicago Press. Chapter 4: The Organism Strikes Back (pp. 77-93); Chapter 6: Mendel
and Molecules (pp. 112-113).
Kampourakis, K. (2022). Understanding Genes. Cambridge University Press.
SUGGESTED ESSAY QUESTIONS:
– What is a gene
– Do genes contain information, and if so, what kind of information
– Does the genotype determine the phenotype
10. Biology, reduction, and complexity
How does biology relate to other sciences Is biology autonomous from physics and chemistry
Reductive and non-reductive explanations in biology. The complexity of living systems.
REQUIRED READINGS:
Mayr, E. (1996). The autonomy of biology: The position of biology among the sciences. The Quarterly
Review of Biology, 71(1), 97-106.
Weber, M. (2005). Philosophy of Experimental Biology. Cambridge University Press. Chapter 2:
Reductionism and the Nature of Explanations (pp. 18-50).
ADDITIONAL SUGGESTED READINGS:
Hull, D. (1974) Philosophy of Biological Science. Prentice-Hall. Chapter 5: Organicism and
Reductionism (pp.; 125-141).
Kaiser, M. I. (2011). The limits of reductionism in the life sciences. History and Philosophy of the Life
Sciences, 33(4), 453–476.
Nagel, E. (1951). Mechanistic explanation and organismic biology. Philosophy and Phenomenological
Research, 11(3), 327-338.
SUGGESTED ESSAY QUESTIONS:
– Is biology autonomous from other sciences such as physics and chemistry
– Can the properties of a complex system such as an organism be explained in terms of the
properties of its parts and their interactions
– The mechanism of the action potential is a physicochemical explanation of a biological
phenomenon. Discuss.