On inference in ecology and evolutionary biology: the problem of multiple causes.

If one investigates a process that has several causes but assumes that it has only one cause, one risks ruling out important causal factors. Three mechanisms account for this mistake: either the significance of the single cause under test is masked by noise contributed by the unsuspected and uncontrolled factors, or the process appears only when two or more causes interact, or the process appears when there are present any of a number of sufficient causes which are not mutually exclusive. In ecology and evolutionary biology, experiments usually test single factor hypotheses, and many scientists apparently believe that hypotheses incorporating several factors are so much more difficult to test that to do so would not be practical. We discuss several areas in ecology and evolutionary biology in which the presupposition of simple causation has apparently impeded progress. We also examine a more mature field, the study of atherosclerosis, in which single factor studies did significantly delay progress towards understanding what now appears to be a multifactor process. The problem has three solutions: either factorial experiments, dynamic models that make quantitative predictions, response-surface methods, or all three. In choosing a definition for 'cause', we make a presupposition that profoundly influences subsequent observations and experimental designs. Alternative definitions of causation should be considered as contributing to potential cures for research problems.