COURSE TITLE: Darwinian Medicine
NUMBER OF CREDITS/HOURS: 3 credits; 54-60 hrs.
NAME OF COORDINATOR: Angel A. Román-Franco
COORDINATOR’S OFFICE: A-359, Dept. of Pathology
MEETING PLACE: Dept of Pathology meeting room

Charles Darwin


Darwinian medicine is the application of modern evolutionary theory to human health and illness, lying at the crossroads between theoretical and applied with human biology, medical anthropology, psychology and physiology. It begins examining the overall concept of evolutionary medicine, advancing through a series of topics showing the scope of impact that evolutionary theory has on medicine today. A continuing theme will be the disparity between proximate and the ultimate causes, approached through an evolutionary interpretation. It will lay emphasis upon those situations and conditions of health (or illness) that necessitate both proximate and ultimate causality. It will address Homo sapiens as a microecosystem grounded on the complex interaction of genotype, phenotype, environment, and behavior that influence health and illness. It will also address recent re-evaluations of the biological locus of Homosapiens, its evolvability and the current status of human evolution.


  • To acquaint the student with the ecology and evolution of host –pathogens interactions and the current status of the evolution of humans and non-human primates,
  • To bring to the fore evolutionary causes of high- impact late-onset diseases, including senescence and cancer.
  • To introduce the concepts needed to understand human pathology in terms of ultimate causes vis –à– vis proximate ones.
  • To deepen understanding of the dyad of ongoing human biological and cultural evolution and the ongoing co-evolution of human pathogens.


New trends in the teaching of the life sciences are being increasingly grounded of evolutionary principles. The great evolutionist Theodosius Dobzhansky stated in a 1973 essay: “Nothing in Biology Makes Sense Except in the Light of Evolution.” Today we can safely state the nothing in the life sciences makes sense if not in the light of evolution. The increasing presence of the evolutionary paradigm makes it essential that life-sciences students become thoroughly acquainted with its foundational principles and their application.

COURSE CONTENT (subject to yearly updates)

First session: Evolutionary medicine-general aspects

  1. Losos, J. B., Arnold, S. J., Bejerano, G., Brodie III, E. D., Hibbett, D., Hoekstra, H. E., … & Turner, T. L. (2013). Evolutionary biology for the 21st century. PLoS biology, 11(1), e1001466.
  2. Manry, J., & Quintana-Murci, L. (2013). A Genome-Wide Perspective of Human Diversity and Its Implications in Infectious Disease. Cold Spring Harbor perspectives in medicine, 3(1).
  3. Anastasiou, E., & Mitchell, P. D. (2013). Paleopathology and genes: investigating the genetics of infectious diseases in excavated human skeletal remains and mummies from past populations. Gene. -38713; No. of pages: 8; 4C

Second session: The human lineage

  1. Cela-Conde, C. J., & Nadal, M. (2012). Taxonomical uses of the species concept in the human lineage. Human Origins Research, 2(1), e1.
  2. Schwartz, G. T. (2012). Growth, Development, and Life History throughout the Evolution of Homo. Current Anthropology, 53(S6), S395-S408.
  3. O’Bleness, M., Searles, V. B., Varki, A., Gagneux, P., & Sikela, J. M. (2012). Evolution of genetic and genomic features unique to the human lineage. Nature Reviews Genetics, 13(12), 853-866.

Third session: Human evolutionary plasticity

  1. Alves I, Šrámková Hanulová A, Foll M, Excoffier L (2012) Genomic Data Reveal a Complex Making of Humans. PLoS Genet 8(7): e1002837. doi:10.1371/journal.pgen.1002837
  2. Reich D, et. al., Genetic history of an archaic hominin group from Denisova Cave in Siberia. Nature. 2010 Dec 23;468(7327):1053-60.
  3. Hochberg Z. Evo-devo of child growth II: human life history and transition between its phases. Eur J Endocrinol. 2009 Feb;160(2):135-41. Epub 2008 Nov 20
  4. Hochberg Z Evo-Devo of child growth III: premature juvenility as an evolutionary trade-off. Horm Res Paediatr. 2010;73(6):430-7. Epub 2010 Apr 15

Fourth session: Evolvability and robustness

  1. Whitacre, J. M. (2012). Biological robustness: paradigms, mechanisms, and systems principles. Frontiers in genetics, 3. 67. May 11. doi:       10.3389/fgene.2012.00067
  2. Tian T, Olson S, Whitacre JM, Harding A. The origins of cancer robustness and evolvability. Integr Biol (Camb). 2011 Jan 11;3(1):17-30. Epub 2010 Oct 14
  3. Kitano H, Oda K, Kimura T, Matsuoka Y, Csete M, Doyle J, Muramatsu M. Metabolic syndrome and robustness tradeoffs. Diabetes. 2004 Dec;53 Suppl 3:S6-S15.

Fifth session: Human evolution and emerging diseases.

  1. Martínez, J. L. (2013), Bacterial pathogens: from natural ecosystems to human hosts. Environmental Microbiology, 15: 325–333. doi: 10.1111/j.1462-2920.2012.02837.x
  2. Rook GA. 99th Dahlem conference on infection, inflammation and chronic inflammatory disorders: darwinian medicine and the ‘hygiene’ or ‘old friends’ hypothesis. Clin Exp Immunol. 2010 Apr;160(1):70-9.
  3. Bourke CD, Maizels RM, Mutapi F. Acquired immune heterogeneity and its sources in human helminth infection. Parasitology. 2011 Feb;138(2):139-59. Epub 2010 Oct 15

Sixth session: Human evolution and nutrition.

  1. Milton K The critical role played by animal source foods in human (Homo) evolution. J Nutr. 2003 Nov;133(11 Suppl 2):3886S-3892S.
  2. Cordain L, Miller JB, Eaton SB, Mann N, Holt SH, Speth JD. Plant-animal subsistence ratios and macronutrient energy estimations in worldwide hunter-gatherer diets. Am J Clin Nutr. 2000 Mar;71(3):682-92.
  3. Cordain L, Eaton SB, Miller JB, Mann N, Hill K. The paradoxical nature of hunter-gatherer diets: meat-based, yet non-atherogenic. Eur J Clin Nutr. 2002 Mar;56 Suppl
  4. Turner, B. L. and Thompson, A. L. (2013), Beyond the Paleolithic prescription: incorporating diversity and flexibility in the study of human diet evolution. Nutrition Reviews, 71: 501–510. doi: 10.1111/nure.12039

Seventh session: Adaptations to temperature and altitude.

  1. Huerta-Sánchez, E., DeGiorgio, M., Pagani, L., Tarekegn, A., Ekong, R., Antao, T., … & Nielsen, R. (2013). Genetic signatures reveal high-altitude adaptation in a set of Ethiopian populations. Molecular biology and evolution. Vol. 71(8):501–510 doi:10.1093/molbev/mst089
  2. Bigham, A. W., Wilson, M. J., Julian, C. G., Kiyamu, M., Vargas, E., Leon‐Velarde, F., … & Shriver, M. D. (2013). Andean and Tibetan patterns of adaptation to high altitude. American Journal of Human Biology. 25:190–197,2013.pdf
  3. Hancock AM, Witonsky DB, Gordon AS, Eshel G, Pritchard JK, Coop G, Di Rienzo A. Adaptations to climate in candidate genes for common metabolic disorders. PLoS Genet. 2008 Feb;4(2):e32. rticle/info%3Adoi%2F10.1371%2Fjournal.pgen.0040032

Eighth session: Gene-culture coevolution: the case of domestication.

  1. Laland, K. N., Odling-Smee, J., & Myles, S. (2010). How culture shaped the human genome: bringing genetics and the human sciences together. Nature Reviews Genetics, 11(2), 137-148.
  2. O’Brien, M. J., & Laland, K. N. (2012). Genes, Culture, and Agriculture. Current Anthropology, 53(4), 434-470.
  3. Enattah NS, et. al., Independent introduction of two lactase-persistence alleles into human populations reflects different history of adaptation to milk culture. Am J Hum Genet. 2008 Jan;82(1):57-72

Ninth session: Evolutionary medicine of hypertension and renal disease.

  1. Young JH, Chang YP, Kim JD, Chretien JP, Klag MJ, Levine MA, Ruff CB, Wang NY, Chakravarti A. Differential susceptibility to hypertension is due to selection during the out-of-Africa expansion. PLoS Genet. 2005 Dec;1(6):e82. Epub 2005 Dec 30
  2. Freedman BI Susceptibility genes for hypertension and renal failure. J Am Soc Nephrol. 2003 Jul;14(7 Suppl 2):S192-4.
  3. Genovese G et al., Association of trypanolytic ApoL1 variants with kidney disease in African Americans. Science. 2010 Aug 13;329(5993):841-5. Epub 2010 Jul 15.
  4. Gravlee CC, Non AL, Mulligan CJ (2009) Genetic Ancestry, Social Classification, and Racial Inequalities in Blood Pressure in Southeastern Puerto Rico. PLoS ONE 4(9): e6821. doi:10.1371/journal.pone.0006821

Tenth session: The human as, and in, an ecosystem.

  1. Rook, G. A., Raison, C. L., & Lowry, C. A. (2013). Childhood microbial experience, immunoregulation, inflammation and adult susceptibility to psychosocial stressors and depression in rich and poor countries. Evolution, Medicine, and Public Health, 2013(1), 14-17.
  2. Munch, E. M., Harris, R. A., Mohammad, M., Benham, A. L., Pejerrey, S. M., Showalter, L., … & Aagaard, K. (2013). Transcriptome Profiling of microRNA by Next-Gen Deep Sequencing Reveals Known and Novel miRNA Species in the Lipid Fraction of Human Breast Milk. PloS one, 8(2), e50564.
  3. Zhang, L., Hou, D., Chen, X., Li, D., Zhu, L., Zhang, Y., … & Zhang, C. Y. (2011). Exogenous plant MIR168a specifically targets mammalian LDLRAP1: evidence of cross-kingdom regulation by microRNA. Cell Research, 22(1), 107-126.
  4. Hehemann, J. H., Correc, G., Barbeyron, T., Helbert, W., Czjzek, M., & Michel, G. (2010). Transfer of carbohydrate-active enzymes from marine bacteria to Japanese gut microbiota. Nature, 464(7290), 908-912.

Eleventh session: Evolutionary medicine and human reproduction.

  1. Voland E. Evolutionary ecology of human reproduction. Annu Rev Anthropol. 1998;27:347-74,5
  2. Lukas, D., & Clutton-Brock, T. H. (2013). The Evolution of Social Monogamy in Mammals. Science, 341(6145), 526-530.
  3. Davis, J. A., & Gallup, G. G., Jr. (2006). Preeclampsia and other pregnancy complications as an adaptive response to unfamiliar semen. In S. M. Platek & T. K. Shackelford (Eds.), Female infidelity and paternal uncertainty (pp. 191–04). New York: Cambridge University Press.
  4. Koelman CA, Coumans AB, Nijman HW, Doxiadis II, Dekker GA, Glaas FH. Correlation between oral sex and a low incidence of preeclampsia: a role for soluble HLA in seminal fluid? J Reprod Immunol 2000; 46: 155–66.
  5. Thornhill, R., & Palmer, C. T. (2000). „The Evolutionary Biology of Rape. von Gerfried Stocker und Christine Schöpf, Wien und New York.

Twelfth session: Human life-span and evolution.

  1. Mace R. Evolutionary ecology of human life history. Anim Behav. 2000 Jan;59(1):1-10.,5
  2. Bochdanovits Z, de Jong G. Antagonistic pleiotropy for life-history traits at the gene expression level. Proc Biol Sci. 2004 Feb 7;271 Suppl 3:S75-8.
  3. Partridge L. The new biology of ageing. Philos Trans R Soc Lond B Biol Sci. 2010 Jan 12;365(1537):147-54.


  • Group Discussion
  • Seminars
  • Critical review of the scientific papers assigned.


  • Participation in seminars
  • Discussion of assigned papers (Rubric and URL included).


            Similar to Medical Sciences requirements for admission to its several faculties and programs.


            Pathology conference room.


  • Participation in seminars
  • Discussion of assigned papers (Rubric and URL included).

GRADING SYSTEM: Medical students: Letter grade: (A,B,C, F)

Biomedical Sciences students –MS/PhD- Pass/Fail

Students with a health condition or situation that, according to the law, make then eligible for reasonable accommodation have the right to submit a written application to the professor and the Dean of their Faculty, according to the procedures established in the document, Submittal Process for Reasonable Accommodation of the Medical Science Campus. A free copy of this document may be obtained at the Office of the Dean for Students Affairs. A copy may also be obtained at the Office of the Faculty Deans as well as in the MSC web page. The application does not exempt student from complying with the academic requirements pertaining to the programs of the Medical Science Campus.

SUGGESTED READINGS: Classics (these are highly recommended publications):

  • Darwin, Charles (1859). On the Origin Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. London: John Murray, 1st Edition. (The Complete Works of Charles Darwin on line: )
  • Schrodinger, Erwin (2002). What is Life? Cambridge: Cambridge University Press. (Re-issue of the 1944 Edition).
  • Monod, Jacques (1971). Chance and Necessity: an Essay on the Natural Philosophy of Modern Biology. New York: Alfred A Knopf.
  • Dawkins, R. (2006). The Selfish Gene: 30th Anniversary Edition. London: Oxford University Press.
  • Mayr, Ernst (2001). What evolution is? New York: Basic Books.


  • Stearns, S.C., & Koella, J.K.. (2007). Evolution in Health and Disease. 2nd Edition. Oxford: Oxford University Press.
  • Trevathan, W. R (2008). Evolutionary Medicine. 2nd Edition. New York: Oxford University Press.
  • O’Higgings, P. & Elton, S. (2012) Medicine and Evolution: Current Applications, Future Prospects, Society for the Study of Human Biology. V.48. London: Taylor and Francis
  • Hall B K and Hallgrímsson B (ed.) (2008) Strickberger’s evolution.The integration of genes, organisms, and populations. 4th edition (Sudbury, MA: Jones and Bartlett, Publishers)


  1. It is a REQUIREMENT to participate and be punctual to all lectures and activities. If you will not be able to attend due to a mayor cause (sickness, death in the family, etc.) you are responsible of notifying the course coordinator PRIOR to the activity and to make arrangements for reposition of the missed activity. As a general rule, no excuses will be given unless there is an emergency and each case will be evaluated separately. If you are absent, you MUST present a formal excuse from the Office of the Assistant Dean of Student Affairs prior to the completion of the course.
  2. In case there is a special need to be absent, the procedures of the “Politica Institucional para Excusar Estudiantes de Actividades Evaluativas o Docentes” will be followed.
  3. Students are expected to demonstrate each and every one of the course objectives.
  4. Should knowledge become available that dishonesty regarding any particular examination has occurred; the course faculty reserves the right to cancel the examination before or after it has been administered and to require a repeat exam or completely eliminate the exam from the course evaluation.
  5. Dress code: All students must comply with the UPR School of Medicine Dress code, approved in 1996 and revised in 2007. (Posted at the SOM Web Page)


Angel A. Román-Franco MD
Distinguished Professor
University of Puerto Rico