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Updated: Feb 25, 2026

A Quantitative Fitness Analysis Workflow
Published on: August 13, 2012
Defining fitness in an uncertain world
Paul Crewe1, Richard Gratwick1, Alan Grafen2,3
1St John's College, University of Oxford, Oxford, OX1 3JP, UK.
This study simplifies natural selection models by integrating Fisher's fitness definition with reproductive values in dynamic populations. It reveals natural selection acts via expected fitness, offering a clearer understanding for evolutionary biology research.
Area of Science:
- Evolutionary Biology
- Population Genetics
- Mathematical Biology
Background:
- Fisher's fundamental theorem of natural selection is crucial for understanding evolution.
- Existing models struggle with complex population dynamics like random environments and class structure.
- Reproductive value definitions require re-evaluation in fluctuating population settings.
Purpose of the Study:
- To unify Fisher's fitness definition with reproductive values in class-structured populations.
- To develop generalizable equations for natural selection under environmental and demographic uncertainty.
- To explore the axiomatic foundations of reproductive value.
Main Methods:
- Combined Fisher's definition of fitness with context-specific reproductive values.
- Developed generalized versions of the Price Equation and Fisher's fundamental theorem.
- Incorporated arbitrary population classes, including continuous variables and varying ploidy (haploid, diploid, haplodiploid).
Main Results:
- Natural selection simplifies to acting solely through the arithmetic expectation of fitness across uncertainties.
- New equations generalize the Price Equation and fundamental theorem for fluctuating demography.
- The model accommodates diverse ploidy levels and arbitrary population structures.
Conclusions:
- The study provides a significantly simplified framework for understanding natural selection in complex populations.
- Results support the individual-as-maximising-agent analogy in finite populations with environmental and distributional fluctuations.
- The model offers potential for improved empirical measurement of fitness in natural populations.
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