Evolution is in the program’s name, and this course draws the site’s evolutionary library into a graduate course on evolutionary epidemiology. Students learn how selection shapes pathogen virulence, drug resistance, and emergence, connecting within-host dynamics to between-host selection. The course uses the virulence–transmission trade-off, the Price equation, adaptive dynamics, and phylodynamics to reason about how pathogens evolve.
The course syllabus is shown below.
Draft syllabus. This is a scaffold for the concentration. Dates, meeting times, and specific assignments will be finalized before the semester begins.
Course title and instructors
Title: Evolutionary Epidemiology and the Evolution of Virulence
Course Number: BIO 6xx (proposed; with an undergraduate cross-list where
noted)
Semester: TBD
Credit Hours: 3
Meeting Time: TBD
Prerequisites: Infectious Disease Ecology;
population genetics recommended (BIO 315)
Course Director: Michael E. DeWitt, MS
Email: medewitt@wakehealth.edu or dewime23@wfu.edu
Course description
Pathogens evolve on the timescale of an outbreak, and that evolution shapes how dangerous they become and whether interventions hold. This course teaches how selection acts on pathogens. Students derive the virulence–transmission trade-off and predict how interventions shift optimal virulence, use the Price equation and adaptive-dynamics reasoning to analyze pathogen evolution, and connect within-host dynamics to between-host selection in a multi-scale view. We cover evolutionary game theory and the evolution of cooperation among pathogens, coalescent theory and phylodynamics, molecular clocks, the evolution of drug resistance, and evolutionary emergence of new pathogens. Students learn to interpret pathogen phylogenies and molecular-clock estimates in an epidemiologic context.
Learning outcomes
Upon successful completion of this course, students will be able to:
- Derive the virulence–transmission trade-off and predict how interventions shift optimal virulence
- Use the Price equation and adaptive-dynamics reasoning to analyze pathogen evolution
- Connect within-host dynamics to between-host selection in a multi-scale view
- Interpret pathogen phylogenies and molecular-clock estimates in an epidemiologic context
- Explain the evolution of drug resistance and its epidemiologic consequences
- Reason about the conditions for evolutionary emergence of new pathogens
Textbook and other resources
There is no single required textbook. Recommended references and readings include:
- Gandon S, Day T, et al. Selected reviews on the evolution of virulence.
- Frank SA. Foundations of Social Evolution. Princeton, 1998.
- Grenfell BT, et al. Unifying the epidemiological and evolutionary dynamics of pathogens. Science 2004;303:327–32. https://doi.org/10.1126/science.1090727
- Volz EM, Koelle K, Bedford T. Viral phylodynamics. PLoS Comput Biol 2013;9:e1002947. https://doi.org/10.1371/journal.pcbi.1002947
Additional primary literature will be assigned throughout the semester.
Site resources
This course draws on the following IDEEE reference pages:
- Evolution of virulence
- The Price equation
- Evolutionary game theory
- Evolution of cooperation
- Selection in population genetics
- Coalescent theory
- The molecular clock
- dN/dS
- Infectious Disease Ecology
Planned reference pages on within-host viral dynamics, nested (multi-scale) models, evolutionary emergence, phylodynamics, adaptive dynamics, and resistance evolution will be added and assigned as they come online.
Course structure and schedule
This course meets over 15 weeks and combines lecture with literature discussion. The schedule below is a draft outline of topics.
| Week | Topic |
|---|---|
| 1 | Introduction to evolutionary epidemiology |
| 2 | Fitness, R0, and selection on pathogens |
| 3 | The virulence–transmission trade-off |
| 4 | Interventions and the evolution of virulence |
| 5 | The Price equation and selection |
| 6 | Adaptive dynamics |
| 7 | Evolutionary game theory and pathogen cooperation |
| 8 | Within-host dynamics |
| 9 | Multi-scale (nested) models |
| 10 | Coalescent theory |
| 11 | Phylodynamics and molecular clocks |
| 12 | Reading pathogen phylogenies |
| 13 | Evolution of drug resistance |
| 14 | Evolutionary emergence of new pathogens |
| 15 | Student project presentations and wrap-up |
Note: Specific dates will be provided at the beginning of the semester. Topics may be adjusted based on class progress and student interests.
Grades and assignments
| Activity | Weight |
|---|---|
| Participation and literature discussion | 20% |
| Reading responses | 20% |
| Problem sets | 30% |
| Final project | 30% |
Final project: Students will analyze the evolution of a pathogen system of their choosing, applying trade-off theory, the Price equation, or phylodynamic reasoning, and synthesizing primary literature into a written report and presentation.
Course policies
Attendance: Regular attendance is expected, particularly for discussion sessions. Please alert the instructor if you are unable to attend for any reason.
Late/Makeup work: Assignments are due on the dates provided. We recognize that extenuating circumstances arise, and assignments may be submitted up to 2 days late without penalty. If you need an extension, contact the instructor as soon as possible and before the due date.
Artificial intelligence: Artificial intelligence tools and large language models such as ChatGPT, Claude, and Gemini are now part of the academic and professional landscape and we encourage you to find ways to use them to enhance your learning. However, if you use these tools, you must cite your sources and provide a detailed description of the tools you used to complete the assignment. In no way can these tools take the place of your own work and understanding of the material. They should be used to supplement your learning, not replace it. You are ultimately responsible for your work including content and the use of valid citations and references. Using these tools without proper attribution is plagiarism and will be treated as such.
Department/School/University policies
Academic Integrity: Wake Forest University is committed to a culture of academic integrity. As a part of this community, you share the responsibility for creating a place of honesty, intellectual curiosity, and individual accountability. As you committed to with your honor pledge signature, you agree “not to deceive any member of the community; not to steal, cheat, or plagiarize on academic work; and not to engage in any other form of academic misconduct.” If you have questions about documenting your work, working with external sources, or working with peers on assigned work, consult with me as soon as possible. Instances of academic dishonesty will be referred to the Honor and Ethics Council.
Accessibility: Wake Forest University provides reasonable accommodations to students with disabilities. If you are in need of an accommodation, please contact me privately as early in the term as possible. Retroactive accommodations will not be provided. Students requiring accommodations must also consult the Center for Learning, Access, and Student Success (118 Reynolda Hall, 336-758-5929, class.wfu.edu).
Accommodations for Religious or Spiritual Practices: Wake Forest University benefits from the multitude of faiths and spiritual identities held by members of our learning community. Should you need accommodations this semester, email me as soon as possible to ensure we have time to develop equitable alternatives.
Class recordings: In case any class recordings are provided, they are reserved only for students in this class for educational purposes and are protected under FERPA. The recordings should not be shared outside the class in any form.
Syllabus change notice
This syllabus and the dates herein are subject to change.