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CAMB/GCB 513 Evolution in Cancer

Carlo C. Maley, Ph.D.

Lauren Merlo, Ph.D.

Cellular and Molecular Oncogenesis Program
The Wistar Institute
3601 Spruce St. Rm. 352B
Philadelphia, PA 19106


  1. To develop a conceptual framework for understanding cancer.
  2. To provide cancer biologists with a background in evolutionary biology and vice versa.
  3. To learn how to read, dissect, and evaluate a research paper.
  4. To develop the critical ability to generate and test biological hypotheses. 

Course Structure

Format: Graduate seminar with both student and faculty presentations and discussion.

Course unit value: 1

Contact hours per week and proposed days of the week: 4 (2hr seminars), TTh 11am-1pm in Blockley 1311 (you are welcome to bring your lunch). In addition, student presenters for each class will be required to meet/email with the instructors as part of their preparation.

Activity types: Each class consists of group discussions led by students responsible for the research articles under discussion for that day. All students are responsible for completing the daily short answer assignment.

Course materials: A reference textbook will be assigned for background reading on evolution (Mark Ridley, Evolution) and ecology (Begon, Townsend and Harper’s Ecology). Both textbooks are recommended but not required. Most of the class will be based on discussion of journal articles.

Enrollment limits and priority: Enrollment will be capped at 20 to facilitate the seminar format. Priority will be assigned as follows:

  1. Students who have taken CAMB 512 (Cancer Genetics).
  2. Students in the following programs: M.D. students, Cell Growth and Cancer, Genomics and Computational Biology, and Ecology, Evolution and Biodiversity
  3. Other BGS students

Evaluation of student performance


Students will be responsible for preparing and leading a discussion of approximately 2 papers during the semester (depending on class size). The preparation will entail background reading of the relevant literature, identification of the critical issues that should be discussed in class and any questions to stimulate debate if class discussion lags.

We will be reading and discussing one paper from the primary literature each class, often with a background paper. Each reading will be accompanied by 3-4 short answer questions designed to make sure students understood the paper and to stimulate their creativity in developing alternative explanations and experimental tests of their ideas. I will randomly collect these responses 1 out of 5 classes (though we reserve the right to collect answers at will). Answers will be collected at the beginning of class (no skipping class to finish an assignment or writing answers during class). The first time responses are collected they will only be graded with a check (whether or not the students did the work). We will then go over the process of how one can read that paper and generate intellectually satisfying responses to the questions. Subsequent collected responses will be graded with a check minus, check or check plus.

Students may work with others to discuss the papers and develop their ideas for the answers to questions. However, the answers must be written in your own words, and you must take intellectual responsibility for developing your critical thinking. If our friend is always the one to come up with the alternative hypotheses and the experiments to test them, you will not do well in class discussion or the final project.

Instances of plagiarism will result in a 0 for the short answer (or mini-grant) portion of the final grade and possible failure of the class.

The final project will be a 5 page mini-grant that consists of an introductory 1-2 paragraphs, 2-3 specific aims and experimental design for each aim. The grants should be designed to fill a gap of our knowledge of the evolution or ecology of neoplastic cells. At the approval of the instructor, students may write a review paper or do a project relating to evolution in cancer on the condition that it has a reasonable chance of getting published. References are not included in the 5 page limit.

Advice on grant writing can be found at:

Important dates:

  October 4 - Project outline due

  November 20 - Final project due        


The grading will be based on short answer questions and problem sets (35%), mini-grant (25%), discussion leadership (15%) and class participation (25%). Furthermore, the lowest homework grade will be dropped from the calculation of the final grade to allow for sick days, emergencies, etc.

The instructors will keep notes on who participated after each class. Students will get extra class participation points for coaxing their shy peers into the discussion. All criticisms of peers should be constructive and should be couched diplomatically so as to encourage further discussion. Criticisms of the literature may be as harsh as you like, unless the authors are in the room