1. Measuring complexity in soil ecosystems Monika Gorzelak November 24 th 2014

2. Objectives How can I ask a CAS question in my research? Soil is complex Should behave as a complex adaptive system Explore whether that is measurable in my experiments/research

4. Outline Revisit “response diversity” Historical context Tilman Cedar Creek Current theory Introduction to mycorrhizal networks My experiment(s) + CAS

6. Response diversity Definition Diversity of responses to environmental change among species that contribute to the same ecosystem function Diversity within functional groups is important to the adaptive capacity of ecosystems; not just species richness

7. Concepts of diversity in plant ecosystems Tilman 2001 Hector 1999 Large scale field experiments that manipulated diversity and measured fitness outcomes Debate

8. Cedar Creek Long term experiment Temperate grasslands, in Minnesota 16 species, chosen at random into plots in increasing species richness Measure productivity (biomass) Increased diversity = increased productivity

9. BIODEPTH Europe experiment Same design, 32 plants 8 sites across Europe Hector et al. (2002) Same results: increased diversity results in increased productivity (biomass)

10. (Loreau, 2010)

11. Mechanisms Complementarity effect: functional niche complementarity, permanent association of species leads to better collective performance Selection effect: trait variation as initial effect upon which selective process promotes dominance of a particular species Both at work (Loreau et al. 2001); complementarity more likely (Cardinal et al. 2007)

12. What functions? Root depth and architecture (Dimitrakopoulos and Schmid 2004) Nutrient use efficiency (van Ruijen and Berendse 2005) Increase input and retention of nitrogen (Fargione et al. 2007) ***grasslands only!

13. Bonfante and Anca 2009

14. More functions, more interactions

15. Mutualisms Nitrogen-fixing bacteria (classic example affecting plant diversity, fix nitrogen, facilitate co-existence of other species by generating a nutrient source) Mycorrhizas. Symbiosis between soil fungi and plant. Plant provides fixed carbon, soil fungus provides mineral nutrient uptake

16. Microscopic structure

17. Colonized root tips Potential for different types of connections because different species

19. Scales of complexity in soil Spatial – Roots create heterogeneous structure* – Networks Temporal – Within a trophic level – Between trophic levels (different time scales apply)

20. Root architecture

21. Networks (Beiler et al. 2010)

22. Scale-free network (Beiler et al. 2010)

23. Temporal (within trophic level) Behaviour* (within life span, quick changes) Adaptation (positive/negative feedbacks) – Expression of genes to exploit resources – Improve fitness – Adaptive niche construction (Callahan 2014), vs. homogeneous medium environmental modification Evolution (“permanent” changes to genome) – Longer term

24. Growth curve example Medium term (mal) adaptation

25. Adaptive niche construction “Opposite” of a growth curve experiment More likely in a heterogeneous environment Modification of a local environment to confer fitness advantage, passing the local improved environment onto offspring Recently demonstrated in a lab setting using bacteria (easier to do with bacteria because they have short generation times)

26. Soil trophic levels 1 st level: primary producers (plants) 2 nd level: decomposers, mutualists, pathogens 3 rd level: shredders, predators, grazers 4 th level: higher level predators (carni) 5 th level: even higher level predators (?) Also: food webs, organisms can play at different levels (nematodes eat fungi and are eaten by other fungi...next slide)

27. Fungi eat nematodes! © George L. Barron

28. Temporal scales across trophic levels Plants interact with mycorrhizas (2 trophic levels) Bacteria interact with both Reaction times differ between these 3 components (scale: years, months, days) (Food webs: there are other players, I’m ignoring them)

29. Seasonal changes in fungi (Pickles et al. 2010)

30. My project: what we know Douglas-fir connect with mycorrhizas which form networks Douglas-fir can transfer resources and signals through its mycorhizal network Douglas-fir preferentially transfers carbon to kin Douglas-fir transfers defense signals via the mycorrhizal network in response to herbivory

31. My project: what we want to know Do Douglas-fir transfer defense signals preferentially to kin in response to herbivory? Need to look at 3 factors: – Mycorrhizal network (yes and no) – Kin vs stranger – Herbivory (insect, manual defoliation, no)

32. Treatments MN=mycorrhizal network, using mesh bags that allow fungi to pass, but not roots Kin vs stranger defoliation insect control MN no-MN no mesh Kin vs stranger No-MN=mesh bags exclude roots and fungi, eliminating the mycorrhizal network, while allowing the passage of water and bulk soil flow including nutrients No-MN=mesh bags exclude roots and fungi, eliminating the mycorrhizal network, while allowing the passage of water and bulk soil flow including nutrients No mesh=control for unknown impacts of including the mesh**changes to spatial heterogeneity I hope to be able to detect by considering complexity of the soil in the design and trying to quantify it under these conditions No mesh=control for unknown impacts of including the mesh**changes to spatial heterogeneity I hope to be able to detect by considering complexity of the soil in the design and trying to quantify it under these conditions

33. Design: Two recipient D = donor Rs = recipient, stranger Rs = recipient, stranger Rs = recipient, kin Rs = recipient, kin

34. Design: Tripartite D = donor Rs = recipient, stranger Rs = recipient, stranger Rs = recipient, kin Rs = recipient, kin

35. Thoughts on design Tripartite is more complex Should I expect different results? Its essentially the same experiment... Dilution of label added to Donor Change to the spatial heterogeneity of the soil structure (can I see this change by monitoring the bacteria...if they change, will they impact the system and possibly alter responses to treatments?)

38. Complexity in soil: Relevant concepts Spatial heterogeneity causes adaptive radiation in bacteria (increase diversification) Priority effects (first bacteria arriving takes priority, impacts habitat in the future) Time-scale differential between plants, fungi, and bacteria

39. Apply treatments: Herbivory, mycorrhizal networks Bacteria Productivity (measure biomass) Diversity species diversity Functional diversity Time-scale comparisons Bacteria at start Bacteria at finish Productivity (measure biomass) Diversity species diversity Functional diversity Time-scale comparisons Bacteria at start Bacteria at finish

40. Diversity and Productivity Aboveground plant diversity unchanged Soil spatial structure is altered, allowing for adaptive radiation Parallel to Tilman: will productivity increase with increased diversity (in bacteria rather than plants?) Plant stress response have different effects on bacterial community systems (is high diversity system more resilient?) Will kin vs stranger change the diversity/productivity of bacteria in the system?

41. Questions Does the bacterial community change in response to – the presence/absence of a mycorrhizal network? – herbivory (with and without a mycorrhizal network)? – Density of seedlings (and therefore changes to root architecture)? – Can these changes be seen temporally over the duration of the experiment? – If layered on top of the original experiment, can these measures inform signal transfer through a MN?

42. Thoughts on measuring complexity? What should I add? Is there a better measure of soil complexity (bacteria will be most responsive)? Change the experiment to include explicit measures of complexity Do a separate experiment to tease out the factors – Consider no plant pot, single plant pots etc

43. Functional diversity in bacteria Too much going on in the design? Simplify with a lab experiment? mimic Tilman experiment, but upside down. Keep above-ground constant, inoculate the soil with known species, use fake roots, fake soil (simplify the system), look for productivity increases and changes in diversity

44. The end.