Caroline
                    Farrior

Competitive games plants play


The success of a strategy for an individual plant is determined by its environment as well as the strategies that other plants play. 

Competition with other plants is intense in a forest, driving allocation to wood and fine roots at levels that decrease a plant's fitness in isolation.

With simplified models of plant physiology we investigated how nitrogen and water availability influence the maximally competitive allocation among the three basic plant tissues - leaves, wood, and fine roots. 

For water-limited plants this has the unintuitive effect of making plant's growth rate independent of water availability during water limitation. 

Predictions from both models, competition for nitrogen and light and competition for water and light, compare well with data from forests across the globe.

Currently we are exploring interactions of these strategies with other important plant traits and resource limitation combinations.
model
                      treeCartoon of a model tree depicting the simplified physiology.

Trees are made out of leaves (green),  woody biomass (grey), and fine roots (brown).  Roots take up water (blue) and nitrogen (red).  Leaf photosynthesis is a function of light availability (yellow), which drops off exponentially through the crown of a tree due to self shading.  If
                                                                    water is limiting carbon assimilation is proportional to water uptake. If nitrogen is limiting the number of leaf layers is limited.


Evolutionarily stable strategy analysis 
Dominant strategy is predicted to be the strategy that, when in monoculture, resists invasion by all other strategies
(if such a strategy exists).

ESS

References                                                                                                                                                                                                               

Farrior, CE, R Dybzinski, SA Levin, and SW Pacala. 2013. 
Competition for water and light in closed-canopy forests: a tractable model of carbon allocation with implications for carbon sinks The American Naturalist 181(3): 314-330.
 

Dybzinski, R, C Farrior, A Wolf, PB Reich, and SW Pacala. 2011.  Evolutionarily stable strategy carbon allocation to foliage, wood, and fine roots in trees competing for light and nitrogen: An analytically tractable, individual-based model and quantitative comparisons to data.  The American Naturalist 177(2):153-166.


Sequential resource limitation in plant communities


Simple resource addition experiments often have surprising results. Why would the same plants respond to more than one resource?  Why would plants respond to additions of a resource by increasing their investment in taking it up?
We added nitrogen and water in a factorial design to diverse prairie plots at Cedar Creek and found such confusing responses. 

Plants responded to both water and nitrogen additions.  With water additions, plants increased their fine-root biomass without concurrent increases in aboveground biomass. Further, this response only occured at low nitrogen levels, making it seem like the high nitrogen plots were water saturated, while the low nitrogen plots were not.

We found explanations for these responses through a comparison with theoretical predictions for competitive dominant allocation strategies of plants in competition for water, nitrogen, and space.

Cedar Creek diverse prairie plots where water and nitrogen were added in a factorial design.








(below)
Separating leaf biomass from structural and reproductive biomass.



Plant biomass responses to resource additions


We found plants are likely sequentially limited by nitrogen and water.  They are saturated with water during periods immediately following rain and limited by water during hot summer days in between storms.  During the periods of water saturation, plant photosynthesis is limited by the photosynthetic machinery it has, powered by nitrogen uptake.  Plants increase fine-root biomass when water is added because competition for this nitrogen becomes more important.  As nitrogen becomes more available, however, the effect is less significant.
References                                                                                                                                                                                                              

Farrior, CE, D Tilman, R Dybzinski, PB Reich, SA Levin, and SW Pacala, 2013.  Resource limitation in a competitive context determines complex plant responses to experimental resource additions. Ecology 94(11): 2505-2517.


Old-growth forest current and future carbon sinks


As the concentration of CO2 in the atmosphere rises, plant leaves become more productive.  But, sometimes this enhanced productivity does not translate into enhanced carbon storage of the whole forest. 

This increased productivity may not be allocated evenly among plant tissues and further plants may respond to the new environment by changing allocation strategy altogether.  These tissues have different residence times within the living biomass and the system.  A unit of carbon allocated to wood stays in the tree on the order of 100 years, while a unit of carbon allocated to fine roots will be shed by the plant and begin to decompose after only 2 years.  Thus changes in allocation can have significant impacts on carbon storage and the significance of plants for carbon sinks.

We are investigating the ways that resource limitation and CO2 fertilization interact with plant allocation strategies to build a mechanistic understanding of current carbon sinks in order to better predict their future.
 




Figure 4 of Farrior et al. 2013 AmNat. 

(A)  Effects of enhanced leaf-level water use efficiency during water-limitation on carbon storage for constant allocation (straight line) and competitive allocation (dashed line).
(B) Effects of enhanced leaf-level maximum photosynthetic rate on carbon storage.

References                                                                                                                                                                                                              
Papers in review, and


Farrior, CE, R Dybzinski, SA Levin, and SW Pacala. 2013.  Competition for water and light in closed-canopy forests: a tractable model of carbon allocation with implications for carbon sinks The American Naturalist 181(3): 314-330.
 


Allocation strategies in forests across a variety of rainfall regimes in the US


In an effort to understand the role of competition and resource availability in forest allocation patterns, I set up 22 plots across different rainfall regimes.

Within these forests, I have taken data on growth, leaf area index, and fine root biomass as well as net-nitrogen mineralization rate and rainfall.  

Sample processing of the final season is still underway, but preliminary results and observations have already motivated much of my work.










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