domingo, 10 de julio de 2016

eMotion, an R function for evolutionary motion of character values

Hi all!

 Some time ago, I was studying an article from Tamara Münkemüller et al. 2015: "Phylogenetic niche conservatism – common pitfalls and ways forward". It´s a nice study where the ability of current tools in order to identify the evolutionary model that drives the changes in a trait (in this case, an ecological suitability) is discussed.
 They simulate the evolution of a trait under several evolutionary processes and then, try to apply the tools that are commonly used to test what kind of process is behind the evolution of the trait, allowing us to infer several patterns or processes as phylogenetic niche conservatism. Their results are really interesting, and I recommend you to take a look to the whole research!

However, I would like to focus in one of the main pictures of the article:



 Looks really nice! As a beginner, it was very difficult to me to understand this awesome figure. From my biological background, to study all this mathematical stuff requires a lot of time that I should invert in laboratory or field work... but it´s also very important to know the underlying statistical background of all the analysis that I would like to apply to my data in the future! So I decided to try to replicate some of those boxes by myself using R language. Many times, it´s the best way to learn!

 The figure shows trait evolution over time under Ornstein–Uhlenbeck 1 parameter in the left column, Ornstein–Uhlenbeck evolutionary at middle column, and Ornstein–Uhlenbeck extended (multiple optimum) at right column, with different selection strength in each row. This means that the first row is actually a Brownian Motion process, since alpha (selection strength) is 0.

 The graphic is the result of applying the OU process equation:


x(t+dt)= x(t)+ alpha(theta(t)-x(t))dt + (sigma dt E)

where , x(t) is the trait value at time=t, alpha is the selection strength with which a specific trait value (optimum) is wanted, theta is the optimal value of the trait in time=t, sigma is Brownian motion rate (the amount of random trait variability per time) and E is a normal distribution(0,1).

 Following this equation, I developed a simple and intuitive R function that simulate an OU evolutionary process for a hypothetical character in two species: eMotion (from "evolutionary motion")

 You can provide each parameter of the OU equation for the two species, and then you should define the number of generations (time) and replicates (number of simulations per species).

 Here you have the code to load the function into R console:

 eMotion <- function (sigma, theta, alpha, sigma2, theta2, alpha2, generations, n) {  
   plot(seq(1:generations),seq(1:generations), ylim=c(-200,200), type="n",   
       xlab="generations", ylab="trait value", main=paste("Sp1- s=", sigma,  
       " th=", theta, " a=", alpha,"  Sp2- s=", sigma2, " th=", theta2,   
       " a=", alpha2, sep=""))  
   trait_a=numeric()  
   simulations= list(1:n)  
   for (j in 1:n){  
     a=0  
     for (i in 1:generations) {   
       a= a + (sigma * rnorm(1)) + (alpha * ( theta - a))  
       trait_a[i]=a  
     }  
     aa=data.frame(trait_a)  
     simulations[j]=aa  
   }  
   for(k in 1:n){  
     bb=data.frame(simulations[k])  
     lines(bb, col="red")  
   }  
   #######  
   trait_a=numeric()  
   simulations= list(1:n)  
   for (j in 1:n){  
     a=0  
     for (i in 1:generations) {   
       a= a + (sigma2 * rnorm(1)) + (alpha2 * ( theta2 - a))  
       trait_a[i]=a  
     }  
     aa=data.frame(trait_a)  
     simulations[j]=aa  
   }  
   for(k in 1:n){  
     bb=data.frame(simulations[k])  
     lines(bb, col="blue")  
   }  
 }  

 Once that you have loaded the function, you can use it to create simple plots to observe the effect of each parameter of OU equation in resultant graphic. For instance, here we describe an OU process of a trait in two species. The species 1 (red) follows an OU process of character evolution, with a sigma value (Brownian motion rate) of 2 and an optimum in 100, while species 2 (blue) follows an OU model with a sigma value of 0.5 and an optimum in -100. In both cases, alpha is 0.002.

eMotion(sigma, theta, alpha, sigma2, theta2, alpha2, generations, n)

 eMotion(2, 100, 0.002, 0.5,-100, 0.002, 10000, 20)  


The function automatically generates this simple plot, where you can observe that species 1 (red), is looking for a trait value around 100 (optimum), while species 2 (blue) is trying to get an optimum near to -100. Although both species have the same alpha (selection strength), the red one has a Brownian rate (sigma) much higher than the blue one. That explains why species 1 (red) has a higher range of variation than species 2 (blue), where all values are always near to the optimum (-100).

 You can use the function to explore other options in parameter values, which could be useful in order to teach about evolutionary processes or to understand the Ornstein–Uhlenbeck equation. Here you have one more example, enjoy!

 eMotion(1, 0, 0, 1,0, 0.01, 10000, 20)  

lunes, 27 de junio de 2016

nichEvolve documentation and code


Here you have some documentation about the nichEvolve model and a link to download the NetLogo code. I will write soon some practical experiments to test with the model... enjoy!
Link:  nichEvolve.nlogo

*(Don't you have NetLogo installed in your computer? NetLogo is a nice tool to implement simple agent-based models, and very useful to introduce yourself in the world of dynamic models!) Download from here: NetLogo

 WHAT IS IT?

Here is a simple approach to explain how different models of trait evolution can work over one or two different species. Two available models of trait evolution are available in the general framework of Ornstein-Uhlenbeck process, with the possibility of adapting it into a simpler Brownian Motion process by changing the settings.
The model could be usefull as a virtual laboratory to teach about this concepts.

author: Javier Fernandez-Lopez  jflopez.bio@gmail.com

HOW IT WORKS

The model represent the evolution of a single ecological trait that could be defined as "ecological suitability", and could be understood for instance, as a specific requirement of temperature or humidity. The program allows an investigator to observe the effect of changes in each parameter of the Ornstein-Uhlenbeck equation

x(t+dt)= x(t)+ alpha(theta(t)-x(t))dt + (sigma dt E)

which define the evolution through time of a trait, where x(t) is the trait value at time=t, sigma is Brownian motion rate (the amount of random variability of the trait per time), theta is the optimal value of the trait in time=t, and alpha is the selection strengh with which a specific trait value is wanted.

 By changing equation parameters, the researcher can check the evolution of the trait value over time in the graphic box at the right side of the model. The "optimum" boxes show as well the theoretical value of the trait value. If speciation switch is "off", the model represents just one species. When speciation is turned "on", the species is split (red and blue) and the trait evolution occur independently. Now, theta parameter (OU-optimum) could differ for each species.

 Left side of the model shows a espatialy-splicit representation of the model. 200 mobile agents of each colour are shown over a grid that simules different ecological conditions. Remember that if speciation is "off", both colours represent the same species with identical ecological requirements. When the simulation starts, each agent try to find a patch around him that best match with his ecological requirements (theorical optimun niche values). The "realized" boxes show the mean of the values of the environmental conditions that the mobile agents have found by tracking the surronding environment. You can also compare the theorical optimum with the realized niche of mobile agents.

 Following KISS philosophy (Keep It Simple, Stupid!) we have not included some processes as extinction when theoretical and realized niches doesn't match, or reproduction of mobile agents.

 Finally, we have included as an experiment the option of "climate-change". When this switch is "on", it only affects to the spatially explicit representation of the model. The environment value of each patch increases by time, changing the niche availability for the mobile agents and affecting (or not) their distribution. The background process of trait evolution is not affected by "climate-change".

HOW TO USE IT

Click "Setup" to initialize the model. Then, you can use "Step" to see how the model runs step by step, or use "Go" button to run the model in a continuous way.

1) Start with a simple Brownian Motion model for a single species (speciation "off"). For this, alpha slide should be 0, theta's values are not important, since alpha (optimum selection strengh) is zero. You could test for changes in graphical box when sigma values are too large or too small.

2) Once that you know the effect of the Brownian motion rate (sigma), you could turn on speciation switch, to split the species. Now you can observe how both species are evolving independently, and, if you are lucky, you could see how their distributions differs from each other in the 2D display.

3) Now you can explore OU models increasing the value of alpha. You can try first using the same theta value for both species and changing it latter. You can also observe how interact Brownian motion rate (sigma) with optimum slection strengh (alpha)

4) Finally you could check the behaivour of mobile agents when climate-change is "on"


THINGS TO NOTICE

 By observing the patterns drawn in graphical box by different combinations of each parameter values, you could think about how difficult it is to guess the actual background process that drives a specific trait evolution if we only know the final state of the values.  


HOW TO CITE

Fernandez-Lopez, J. 2016. nichEvolve: a niche evolution model



domingo, 26 de junio de 2016

nichEvolve, a niche evolution model with NetLogo


 During the last year I was reading some articles about niche conservatism/evolution. There are a lot of theories and methodologies to deal with this kind of questions, but sometimes it's hard to understand all this mathematical and theoretical concepts about models of evolution: Brownian motion models, Ornstein-Uhlenbeck processes, etc. Following KISS philosophy (Keep It Simple, Stupid!) I wrote some code using the wonderful tool NetLogo to develop a very simple model that helps to understand some simple processes related with ecological traits evolution...

 The model implements two usual processes used to describe the evolution of trait values: Brownian motion and Ornstein-Uhlenbeck. While Brownian motion process represents random evolution of a trait value, depending only on Brownian rate (amount of change allowed per time), Ornstein-Uhlenbeck process means directional evolution towards an optimum with a specific selection strength. In the model you can explore diverse combinations of each parameter and observe the results also in a spatial explicit display with mobile agents that track environmental conditions trying to match their theoretical requirements.


 Code will be uploaded soon, so you could modify it and report any suggestion or bug.

 Cheers!