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Improving Model of Interaction
               Networks

              Konstans Wells & Bob O'Hara



                              Blogged at
http://blogs.nature.com/boboh/2012/03/29/doing_stuff_with_ecological_network
                                      s
Typical Data


                         Fi lu    Fi th     Pr af    Fi su    Ma bu
Commom Bulbul            128      110        70       30       27
Blue Monkey               19       35         2       28       36
Red tailed Monkey         11       19         0       52       29
Voilet backed starling     1        0       190        0        0
Blackcap                  64       23         3        8        9




  http://www.nceas.ucsb.edu/interactionweb/html/schleuning-et-al-2010.html
What's wrong?
Network statistics are messy


          Q=
              1
                     ( ki k j
                                 )
                ∑ Aij − 2m δ(c i , c j )
             2m i , j


Derived for known networks
  behaviour when uncertain is difficult to understand
Statistical Problems
How do we estimate
 sampling error?
Are the zeroes real?
What if the sampling is
 not representative?
A Better Approach




Model the data
Building the Model I

                       Fi lu   Fi th   Pr af   Fi su   Ma bu
 Common Bulbul 1        λ11     λ21    λ131     λ41     λ51
 Common Bulbul 2        λ12     λ22    λ132     λ42     λ52

 Red tailed Monkey 1    λ13     λ23     λ33     λ43     λ53

 Red tailed Monkey 2    λ14     λ24     λ34     λ44     λ54

 Red tailed Monkey 3    λ15     λ25     λ35     λ45     λ55



Start with mean rates of interaction per individual
Building the Model II

                         Fi lu   Fi th   Pr af   Fi su   Ma bu
   Common Bulbul 1        λ11     λ21    λ131     λ41     λ51
   Common Bulbul 2        λ12     λ22    λ132     λ42     λ52

   Red tailed Monkey 1    λ13     λ23     λ33     0       λ53

   Red tailed Monkey 2    λ14     0       λ34     λ44     λ54

   Red tailed Monkey 3    λ15     λ25     λ35     λ45     λ55



Can set some means to zero
What is λij?

Individual rate of visitation


    No. of visits ~ Poisson(λij)



We can model this further
Individual To Species
Species-level rate of interaction is


          Λc ,r=     ∑                            ̄
                                  λ i , j =nc m r λ c , r
                   i , j ∈c , r
Individual To Species
Species-level rate of interaction is


          Λc ,r=     ∑                            ̄
                                  λ i , j =nc m r λ c , r
                   i , j ∈c , r

              Abundances of
              resource &
              consumer
Individual To Species
Species-level rate of interaction is


          Λc ,r=     ∑                            ̄
                                  λ i , j =nc m r λ c , r
                   i , j ∈c , r

              Abundances of                           Mean individual-
              resource &                               level
              consumer                                 preferences

  Extracts abundance effects from preferences
In practice...
We might observe trees, and not be able to
 distinguish individuals visiting them
  We have several resources, but lump consumers
   together
     nc = 1

  If we estimate nc, we can get back to λij
Further modelling
Log-linear:

        log(λ ij )=β(r i , c j )+ γ (i , j)

        β(r i , c j )=ρ(r i )+ χ (c j )+ ι (r i , c j )
                Resource + Species + Interaction


 Separates out “palatability” and “hungriness” from
  specificity
Better Measures
  Modularity

Q=
    1
   2m i , j (  ki k j
                           )    1
      ∑ A ij − 2m δ(ci , c j )= 2 ∑ ( pij − pi⋅ p⋅j ) δ(c i , c j )
                                  i, j

 “the fraction of edges that fall within communities minus the
   expected value of the same quantity if edges are assigned at
   random, conditional on the given community memberships
   and the degrees of vertices.”
           But

             logit p ij −logit pi⋅ p⋅j =ι (r i , c j )
Fitting the Model
Simplest: log-linear model


    glm(Count ~ Resource*Consumer,
                   family=poisson())


Assumes no over-dispersion
Can model further, e.g. add resource-specific
 covariates
More complicated models
If we have several individuals of resource and
   consumer:


glm(Count ~ Resource*Consumer +
               Res.Ind*Con.Ind + offset(Time),
 family=poisson()


Now Res.Ind:Con.Ind is over-dispersion
Could use random effects
Adding Zeroes
Where the data is a 0, λij is estimated as low
  Can't tell where the “true” zeroes are
So, use a zero-inflated Poisson distribution
  adds zeroes
But they are uncertain
How well does our model perform?
Simulation study
12 resource species, 9 consumers
   Effects: Each cell a 3 x 3 matrix
               Generalist     Opportunist   Specialist
Generalist         0.75           0.25          0
Opportunist        0.2            0.01          0
Specialist         0.01                0       0.2
Erratic            0.05           0.01          0
Sample Sizes
Balanced:
  3,5,10,15,20 individuals of each species
Unbalanced:
  5 individuals of each resource species
  5, 10, 15, 20 individuals of generalist consumers
  3 individuals of other consumers
Balanced Results
Unbalanced Results
Thoughts
Model links more closely to actual mechanisms
  more interpretable
Can build models for specific questions
  modularity
Can build hierarchical models, to combine
 several networks
  meta-regression
What More?




You tell us

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Interaction networks

  • 1. Improving Model of Interaction Networks Konstans Wells & Bob O'Hara Blogged at http://blogs.nature.com/boboh/2012/03/29/doing_stuff_with_ecological_network s
  • 2. Typical Data Fi lu Fi th Pr af Fi su Ma bu Commom Bulbul 128 110 70 30 27 Blue Monkey 19 35 2 28 36 Red tailed Monkey 11 19 0 52 29 Voilet backed starling 1 0 190 0 0 Blackcap 64 23 3 8 9 http://www.nceas.ucsb.edu/interactionweb/html/schleuning-et-al-2010.html
  • 3. What's wrong? Network statistics are messy Q= 1 ( ki k j ) ∑ Aij − 2m δ(c i , c j ) 2m i , j Derived for known networks behaviour when uncertain is difficult to understand
  • 4. Statistical Problems How do we estimate sampling error? Are the zeroes real? What if the sampling is not representative?
  • 6. Building the Model I Fi lu Fi th Pr af Fi su Ma bu Common Bulbul 1 λ11 λ21 λ131 λ41 λ51 Common Bulbul 2 λ12 λ22 λ132 λ42 λ52 Red tailed Monkey 1 λ13 λ23 λ33 λ43 λ53 Red tailed Monkey 2 λ14 λ24 λ34 λ44 λ54 Red tailed Monkey 3 λ15 λ25 λ35 λ45 λ55 Start with mean rates of interaction per individual
  • 7. Building the Model II Fi lu Fi th Pr af Fi su Ma bu Common Bulbul 1 λ11 λ21 λ131 λ41 λ51 Common Bulbul 2 λ12 λ22 λ132 λ42 λ52 Red tailed Monkey 1 λ13 λ23 λ33 0 λ53 Red tailed Monkey 2 λ14 0 λ34 λ44 λ54 Red tailed Monkey 3 λ15 λ25 λ35 λ45 λ55 Can set some means to zero
  • 8. What is λij? Individual rate of visitation No. of visits ~ Poisson(λij) We can model this further
  • 9. Individual To Species Species-level rate of interaction is Λc ,r= ∑ ̄ λ i , j =nc m r λ c , r i , j ∈c , r
  • 10. Individual To Species Species-level rate of interaction is Λc ,r= ∑ ̄ λ i , j =nc m r λ c , r i , j ∈c , r Abundances of resource & consumer
  • 11. Individual To Species Species-level rate of interaction is Λc ,r= ∑ ̄ λ i , j =nc m r λ c , r i , j ∈c , r Abundances of Mean individual- resource & level consumer preferences Extracts abundance effects from preferences
  • 12. In practice... We might observe trees, and not be able to distinguish individuals visiting them We have several resources, but lump consumers together nc = 1 If we estimate nc, we can get back to λij
  • 13. Further modelling Log-linear: log(λ ij )=β(r i , c j )+ γ (i , j) β(r i , c j )=ρ(r i )+ χ (c j )+ ι (r i , c j ) Resource + Species + Interaction Separates out “palatability” and “hungriness” from specificity
  • 14. Better Measures Modularity Q= 1 2m i , j ( ki k j ) 1 ∑ A ij − 2m δ(ci , c j )= 2 ∑ ( pij − pi⋅ p⋅j ) δ(c i , c j ) i, j “the fraction of edges that fall within communities minus the expected value of the same quantity if edges are assigned at random, conditional on the given community memberships and the degrees of vertices.” But logit p ij −logit pi⋅ p⋅j =ι (r i , c j )
  • 15. Fitting the Model Simplest: log-linear model glm(Count ~ Resource*Consumer, family=poisson()) Assumes no over-dispersion Can model further, e.g. add resource-specific covariates
  • 16. More complicated models If we have several individuals of resource and consumer: glm(Count ~ Resource*Consumer + Res.Ind*Con.Ind + offset(Time), family=poisson() Now Res.Ind:Con.Ind is over-dispersion Could use random effects
  • 17. Adding Zeroes Where the data is a 0, λij is estimated as low Can't tell where the “true” zeroes are So, use a zero-inflated Poisson distribution adds zeroes But they are uncertain
  • 18. How well does our model perform? Simulation study 12 resource species, 9 consumers Effects: Each cell a 3 x 3 matrix Generalist Opportunist Specialist Generalist 0.75 0.25 0 Opportunist 0.2 0.01 0 Specialist 0.01 0 0.2 Erratic 0.05 0.01 0
  • 19. Sample Sizes Balanced: 3,5,10,15,20 individuals of each species Unbalanced: 5 individuals of each resource species 5, 10, 15, 20 individuals of generalist consumers 3 individuals of other consumers
  • 22. Thoughts Model links more closely to actual mechanisms more interpretable Can build models for specific questions modularity Can build hierarchical models, to combine several networks meta-regression

Notes de l'éditeur

  1. Agent-based interaction models: estimating per individual interaction strength and covariates before simplifying data into per species ecological networks