Studying evolutionary processes using models

In working with this tutorial, you should be learning about the following questions:

What biological phenomena can be studied with models?

How are models used to test hypotheses?

What are the procedures needed to use Populusâ?

How should results be compiled?

How should results be presented?

 

The word “theory” has very different meanings.  Some people use it in a negative sense (the flat earth theory).  It can describe an idea that is reasonable, but unproven (the theory that overgrazing causes a catastrophic shift of short-grass prairie to desert).  It can describe a set of concepts in science that are highly explanatory, integrative, and essentially proven (the theory of relativity, the theory of natural selection).  And finally, sets of mathematical equations derived to represent real-world processes comprise a “theory” about how those processes actually work.  In this tutorial, you will be using “theory” in the sense of using equations to model the real world.

 

What biological phenomena can be studied with models?

 

Just about any process can be approached with a mathematical model.  One needs to be able to identify objects (such as genotypes), have a way to quantify (such as counts), and have reasonable guesses about factors that influence changes in the objects (such as mortality, reproductive success).  If we want to understand how the relative abundance of different genotypes changes over generations, we can construct equations that represent the abundance of each genotype and the fitness of each genotype. 

 

Of course, a mathematical model must simplify the complexity of the real world.  The basic question is, can we capture the essence of biological process with our simpler model?  The model provides predictions about how organisms might react to differences in conditions (such as different fitnesses), but these predictions must be tested against the real world.  If the models are too simple to accurately predict experimental outcomes or observations in the real world, then we can increase reality (?) to the simple equations by adding more complex interactions (e.g., allow fitness to change with the abundance of each genotype). 

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How are models used to test hypotheses?

 

As indicated above, we start with a set of equations and parameters.  Theoretical biologists (e.g., biologists that develop mathematical theories – not “hypothetical biologists”) apply two approaches to see what predictions are derived from those equations and parameters.  The analytical approach solves sets of equations to derive predictions about the state of the system after it reaches equilibrium.  Much of population genetics is analytical.  The simulation approach calculates changes in the system with each cycle of time (e.g., each generation of survival and reproduction), to provide numerical patterns of change.  Obviously, computers are quite handy for the simulation approach.  The analytical approach has great advantages in being more general in its predictions, while the simulation approach allows for more complexity and more quantitative predictions.  We will be using the simulation approach in this tutorial.

 

The software that you saw used in class (Populusâ) is available for free for educational use.  The program has models for many different ecological and evolutionary processes; we are using just a small set of these models (but feel free to play around with models for population growth, competition, and predation).  In the autosomal selection model, Populusâ uses the simple equations that are given in the text and in class to calculate changes in genotype frequency, given particular fitnesses for each genotype.  You will set the parameters of genotype fitness.  In doing so, you can simulate no selection, directional selection, stabilizing selection and diversifying selection.  You also can simulate different forms of developmental influence such as dominance, co-dominance, or intermediate dominance.   After recalculating genotype frequencies over many generations (you can control the number of “iterations”), the software provides clear pictures of how the system changes, using different ways of looking at the system and it also is possible to examine numerical results.


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What are the procedures needed to use Populusâ?

  1. Start Populusâ by double-clicking on the Populusâ icon on the computer desktop.  The software is installed on all the computers in the computer lab in the basement of Fischman Hall.  You can download the program if you have a Windows operating system and Microsoft Explorer (see the course web page for instructions).
  2. Click in the “Model” box in the left side of the menu bar; select “autosomal selection.”
  3. In Plot Options, keep “p vs t”; later, you can change this to examine shifts in genotype frequencies.  “p” is, of course, the frequency of the model “A” allele. 
  4. In the “Fitness/Selection Coefficients” box, make sure the “fitness” tab is selected.  Later, you can select the “selection” tab, in which you examine the effects of different degrees of heritability.
  5. Set genotype fitnesses according to the hypothesis you wish to test (see below).
  6. Under “Initial Conditions” highlight the button for “Six Initial Frequencies.”

 


7. Now you are ready to run the simulation:  Click on the “View” button (green arrows).   The following graph will appear:

 

 

 

 

Each color represents the predicted change in the frequency of “A” over 200 generations, given different starting frequencies.  Note that the model predicts that A will become the only allele eventually (at equilibrium) no matter what the starting frequency. 

 

Using options at the top of the graph screen, you can save your results to a new file.  You can save the graph as a picture, or you can save a text file with the data used to generate the graph.  Be careful to note where you save files (you should use your H drive, not the local C drive). 

 

Also, you should take notes on the important predictions of each run of the model.  For example, if you reduce the heterozygote and aa homozygote fitnesses, how does the time to reach equilibrium change?

8. Now you can go back to the starting screen and alter starting conditions to evaluate different forms of selection.

 

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How should the results be compiled and presented?

First, establish the question that you are exploring.

Second, identify the response variable of interest - how do you measure the outcome?

Third, determine the parameters to change and the range of values you wish to explore.

Fourth, conduct a series of simulations, recording the responses.

Using a graph or table, organize the input and response data.

Finally, evalute the relationships and patterns.

 

Here are some questions:

If an allele is lethal but recessive, how fast can selection remove it from a population?

          For this question, you would set aa fitness to zero, AA and Aa to 1.0.

          Run the program, and determine the number of generations to fixation.

                   You may want to use “Options” on the graph menu to set up a grid.

                   Write the number of generations as a function of starting frequency.

          Report your results in the form of a short table.     

How is the rate of elimination of a deleterious allele affected by the selection differential?

          For this question, you will need to vary “aa” fitness (fitness = 1 – s).

          Choose a single starting frequency.

          Run the simulation and note the time to fixation.

          Reset the simulation for a different fitness, and note the new time to fixation.

          You will need to set up a table with a column for “aa” fitness and a column for

                   corresponding time to fixation.

          An Excel graph would be a good way to present this relationship.

Some other questions:

Under what combinations of genotype fitness will genetic diversity be maintained?

What happens when the heterozygote has the lowest fitness?


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