Mycelium in the PRV

1. Mycelium in the PRV

2. I. Evolution of Land Plants • Land plants arose from mutualistic relationships between fungi and green algae • Evidence: • Over 90% of land plants have symbioses with mycorrhizal fungi • Fungi: aid in water and nutrient uptake, • Plants provide fungi with sugars due to photosynthetic ability • Incorporation of fungi into seeds • VAM associations revealed in palaeobotanical study • Genes required for mycorrhizal associations present in ancestors of land plants (Wang et al., 2009) (Stamets, 2005) (Margulis & Fester Eds., 1991) (Brundrett, 2002)

3. http://mycorrhizas.info/evolution/root-evolution.gif

4. II. Types of Fungi • Saprophytic: decomposers (soil builders) Example in Potomac River Valley: (Stamets, 2005) • Morels (black, yellow) can be saprophytic and also mycorrhizal (Kuo, 2007) http://michiganmushroomhunters.org/Images/morels/Black%20morel.jpg

5. II. Types of Fungi • Parasitic: inflict cankers/lesions on trees Example: Phytophthora ramorum - causes sudden oak disease Link to PRV: Oaks in PRV are susceptible to this fungal pathogen (Stamets, 2005) (Virginia Bioinformatics Institute, 2004) http://cisr.ucr.edu/sudden_oak_death.html

6. II. Types of Fungi • Mycorrhizal: form mutualisms with plants • Ectomycorrhizal-form sheaths around the roots of partner plants • Endomycorrhizal-invade interior root cells of host plants (also called vesicular arbuscular mycorrhizae, VAM (Stamets, 2005) (Margulis & Fester Eds., 1991) http://www.palaeos.com/Plants/Lists/Glossary/Images/Endomycorrhizae.gif

7. II. Types of Fungi • Benefits of mycorrhizal associations: • Increased length and surface area for absorption • Cool fact: Absorption capacity of mycorrhizal fungi may be 10-100 times greater than SA of leaves in a forest (Stamets, 2005) http://www3.ntu.edu.sg/home/mvvkulish/Image21.jpg http://www.technion.ac.il/~mdcourse/274203/slides/Digestive%20tract/17-Intestinal%20villi%20Jejunum-A.jpg http://course1.winona.edu/sberg/IMAGES/mito3.gif

8. II. Types of Fungi (Johnson et al., 2006)

9. II. Types of Fungi 2. Nutrient sharing (one mushroom species can connect many acres of a forest in a continuous network of cells) • Link to coevolution: • Possibility of fungi providing more nutrients to a tree with which it forms a better mutualistic association (positive feedback) (Stamets, 2005) (Johnson et al., 2006)

10. II. Types of Fungi Ectomycorrhizal Shaded Douglas Fir Western Red Cedar Nutrient Sharing Paper Birch Endomycorrhizal (Stamets, 2005)

11. II. Types of Fungi • Example of nutrient sharing: • Douglas fir and paper birch connected by one type of ectomycorrhizal fungi • Western Red Cedar in a mutualistic relationship with a different type of mycorrhizae (a VAM) • Differential shading led to change in nutrient flow (from birch to fir) • Amount of carbon transferred is directly related to amount of shading • Possibly helps saplings survive under canopy? (Link to dogwoods in PRV coming later!) (Stamets, 2005)

12. II. Types of Fungi • Benefits continued… 3. Resistance to pests 4. Tolerance of extreme conditions (Bouchez & Roncho, 2008) (Johnson et al., 2006)

13. II. Types of Fungi • Benefits continued… 5. Soil aggregation 6. Reduced erosion (Johnson et al., 2006)

14. Benefits on different ecological scales: II. Types of Fungi (Johnson et al., 2006)

15. II. Types of Fungi • Endophytic: mutualistic fungi that live inside plants • Ex: Endophytic fungi in grasses produce mycotoxins that offer protection from grazers (Faeth, 2002) http://www.ifsqn.com/images/cow.gif

16. III. Plants of the Potomac River Gorge and Surrounding Area • Corridor of biodiversity • Variety of seeds brought to PRV due to diverse ecosystems in watershed • Gorge is surrounded by unique rock formations called “bedrock terraces” • 1,400+ different kinds of plants grow in the Potomac Valley Gorge (Cohen, 2005)

17. III. Plants of the Potomac River Gorge and Surrounding Area • Examples of herbaceous plants include wild columbine, golden rod lyre-leaved rock cress, Virginia blue bells, Dutchmen’s breeches, jewelweed and wild ramps (leeks) • Examples of woody plants include white ash, red maple, sycamore, sugar maples, red & white oaks, hickory and flowering dogwoods • Examples of woody shrub-like plants include hop hornbeam and bladdernut (Cohen, 2005)

18. III. Plants of the Potomac River Gorge and Surrounding Area • What type of plant is this? (Game!) Hickory! (Cohen, 2005) http://www.dcr.virginia.gov/natural_heritage/ncPIf.shtml

19. III. Plants of the Potomac River Gorge and Surrounding Area • What type of plant is this? (Game!) Red Maple! (Cohen, 2005) http://www.dcr.virginia.gov/natural_heritage/ncPIf.shtml

20. III. Plants of the Potomac River Gorge and Surrounding Area • What type of plant is this? (Game!) Switch grass! (Cohen, 2005) http://www.dcr.virginia.gov/natural_heritage/ncPIf.shtml

21. III. Plants of the Potomac River Gorge and Surrounding Area • What type of plant is this? (Game!) Sycamore! (Cohen, 2005) http://www.dcr.virginia.gov/natural_heritage/ncPIf.shtml

22. III. Plants of the Potomac River Gorge and Surrounding Area • What type of plant is this? (Game!) Silver Maple! (Cohen, 2005) http://www.dcr.virginia.gov/natural_heritage/ncPIf.shtml

23. III. Plants of the Potomac River Gorge and Surrounding Area • What type of plant is this? (Game!) Western Sunflower! (Cohen, 2005) http://www.dcr.virginia.gov/natural_heritage/ncPIf.shtml

24. III. Plants of the Potomac River Gorge and Surrounding Area • What type of plant is this? (Game!) White Ash! (Cohen, 2005) http://www.dcr.virginia.gov/natural_heritage/ncPIf.shtml

25. III. Plants of the Potomac River Gorge and Surrounding Area • What type of plant is this? (Game!) White oak! (Cohen, 2005) http://www.dcr.virginia.gov/natural_heritage/ncPIf.shtml

26. III. Plants of the Potomac River Gorge and Surrounding Area • What type of plant is this? (Game!) Virginia Bluebells! (Cohen, 2005) http://www.dcr.virginia.gov/natural_heritage/ncPIf.shtml

27. IV. PRV Mutualisms • Mycorrhizae and tree mutualisms: • Cantharellus cibarius (ectomycorrhizal) and Quercus rubra (Northern red oak) • Fraxinus americana (white ash) and morels (many different species) (Stamets, 2005) http://www.dcr.virginia.gov/natural_heritage/ncTIg.shtml

28. IV. PRV Mutualisms • Flowering Dogwood (Cornus florida) & Fungi: • Mycorrhizal fungi allocate nutrients to dogwoods (an understory tree) from canopy trees • Pathogenic fungi (Dogwood anthracnose or Discula destructiva) (Kuo, 2007) http://www.dcr.virginia.gov/natural_heritage/ncTIIIc.shtml (Carr & Banas, 2000)

29. IV. PRV Mutualisms • Mycorrhizae and herbaceous plants: • Wild leeks (also called ramps) form mutualistic relationships with mycelium which promote leek health by: • Bioremediation • Increasing surface area for water uptake • Changing physical and chemical properties of soil (Leyval & Binet, 1998) http://greayer.com/studiog/wp-content/uploads/2009/04/mosaic1602239.jpg

30. IV. PRV Mutualisms • Fungal and animal mutualisms: • Gyrodon merulioides-a fungi found near white ash • Forms a mutualistic relationship with Meliarhizophagus fraxinifolii (leafcurl ash aphid) • Fungi: form structures that surround/protect aphid • Aphids: provide honeydew to fungi (Kuo, 2007) http://mycorrhizas.info/methods/ash-bolete-aphid.jpg

31. IV. PRV Mutualisms • Fungal and animal mutualisms: • Marsh periwinkle and fungi: • Snail eats fungi and then defecates on marsh grass • Fungi break down marsh grass, snail returns and eats fungi and digested marsh grass (Silliman & Newell, 2003) http://www.frauleindi.com/images/HHNature/OurSaltMarsh-MarshPeriwinkles.jpg

32. V. Fungal Associations and History • Example: Salem witch trials • Thought questions: • Did mycorrhizal interactions affect the formation of civilizations (dependence on agriculture)? If so, how? • How will/does the use of fungicides affect agriculture? • How can we apply our knowledge of fungal mutualisms to enhance crop production? (Le Couteur & Burreson, 2003)

33. V. Fungal Associations and History • How can we apply our knowledge of fungal mutualisms to enhance crop production? • Example 1: Insect control • Production of mycotoxins by endophytes can be used to kill insects. • Link to PRV: Could lead to decreased use of insecticides. (Stamets, 2005)

34. V. Fungal Associations and History • How can we apply our knowledge of fungal mutualisms to enhance crop production? • Example 2: Microbial Control • Application of endophytes to cocoa trees led to a decline in leaf necrosis (possible antimicrobial activity?) • Application of an endophyte (called Piriformospora indica) to wheat plants caused an increase in biomass and seed production by protecting roots from pathogenic microorganisms • Link to PRV: Could be biodefensive (antimicrobial) against fungus that causes sudden oak disease or the fungus that affects the dogwoods. (Stamets, 2005)

35. V. Fungal Associations and History • How can we apply our knowledge of fungal mutualisms to enhance crop production? • Example 3: Heat and Drought Tolerance • Dusting wheat with extremophile endophytes from Yellowstone conferred increased drought resistance • Barley associated with mycorrhizae grew better under poor conditions (drought, addition of xenobiotics) than barley without fungal associations • Link to PRV: Could be used in fields surrounding the PRV which could promote increased yield due to tolerance of drought conditions and increased ability to produce biomass without fertilizers. (Stamets, 2005) (Khalvati et. al, 2009)

36. VI. Conserving the PRV • Future and current threats to biodiversity of Potomac River Gorge plant species include (in order of impact): • 1. Development • 2. Invasion of non-native species (currently 273 exotic species grow within the gorge including English ivy and Japanese honeysuckle) • 3. Overabundance of foraging deer (78 per square mile in VA, 40+ is considered too much) • 4. Too many visitors (over 3.5 million each year) (Cohen, 2005)

37. VI. Conserving the PRV • Possible methods to conserve the PRV using fungi as a means of ecological restoration and facilitation: • Mycorestoratin through: • Mycofiltration (purifying water) • Mycoforestry (ecoforestry policy) • Mycoremediation (denaturing toxic wastes) • Mycopesticides (controlling insect pests) (Stamets, 2005)

38. VI. Conserving the PRV • Problem 1: Fossil fuel emissions • Fossil fuels release polyaromatic hydrocarbons (PAHs) • One experiment found that increasing amounts of PAHs in the soil decreased mycorrhizal colonization of plant roots • The same experiment found that plants with mycorrhizal associations were able to survive and grow in soils with higher amounts of PAH than plants without these associations (Leyval & Binet, 1998) (Bouchez et. al, 1995)

39. VI. Conserving the PRV • Possible reasons for these findings include: • Mycorrhizae bioremediate through bacterial recruitment and improving soil conditions • Bacteria that are recruited can use PAHs as a source of carbon and thus break them down and detoxify them • Link to PRV: Leeks (wild ramps in the PRV) were used in this study and could help detoxify fossil fuel emissions from major highways (Leyval & Binet, 1998) (Bouchez et. al, 1995)

40. (Whelan & Rock, 2006)

41. VI. Conserving the PRV • Problem 2: Input of nitrates into Chesapeake Bay: • One experiment found that Paxillus involutus, a mycelium associated with Picea abies (Norway spruce) and (Betula pendula) silver birch increased nitrate assimilation (although it was affected by pH) • Are there any mutualisms in the PRV that do this? (Andersson et. al, 1994)

42. VII. Link to Final Product: • How can we use mycorrhizae’s ability to bioremediate to decrease levels of pollutants at lower points in the PRV watershed in order to protect other kinds of symbioses found in the PRV?

43. References • Beltrano, J. &  Ronco, M.G. (2008). Improved tolerance of wheat plants (Triticum aestivum L.) to drought stress and rewatering by the arbuscular mycorrhizal fungus Glomus claroideum: effect on growth and cell membrane stability. Braz. J. Plant Physiol. [online]. 20(1), 29-37 . Retrieved from: <http://www.scielo.br/scielo.php?script=sci_arttext&pid=S1677- 04202008000100004&lng=en&nrm=iso>. ISSN 1677-0420.  doi: 10.1590/S1677-04202008000100004. • Bouchez, M., Blanchet, D., and Vandercasteele, J.P. (1995). “Degradation of polycyclic aromatic hydrocarbons by pure strains and by defined strain associations: inhibition phenomena and cometabolism.” Applied Microbiology and Biotechnology. 43.1: 156-164. • Brundrett, M. C. (2002). Coevolution of roots and mycorrhizas of land plants. New Phytologist, 154(2), 275-304. • Carr, D. E., & Banas, L. E. (2000). Dogwood Anthracnose (Discula Destructiva): Effects of and Consequences for Host (Cornus Florida) Demography. American Midland Naturalist, 143(1), 169-177.    • Cohen, J.P. (2005). “A wild river runs through Washington.” Smithsonian ZooGoer 34(6). Retrieved from: http://nationalzoo.si.edu/Publications/ZooGoer/2005/6/potomac.cfm

44. References • Ek, H., Andersson, S., Arnebrant, K., & Söderström, B. (1994). Growth and Assimilation of NH4 + and NO3 - by Paxillus involutus in Association with Betula pendula and Picea abies as Affected by Substrate pH. New Phytologist, 128(4), 629-637.   • Faeth, S. H. (2002). Are endophytic fungi defensive plant mutualists? Oikos, 98(1), 25-36.   • Johnson, N. C., Jason D. Hoeksema, Bever, J. D., Chaudhary, V. B., Gehring, C., Klironomos, J., et al. (2006). “From Lilliput to Brobdingnag: Extending Models of Mycorrhizal Function across Scales.” BioScience, 56(11), 889-900.   • Khalvati, M., Bernadett, B., Dupigny, A. and Schroder, P. (2009). “Arbuscular mycorrhizal association is beneficial for growth and detoxification of xenobiotics of barley under drought stress.” Journal of Soils and Sediments JSS. 1.1: 54- 64. • Kuo, M. (2007). Mushroom Expert.com. Retrieved from: http://www.mushroomexpert.com/trees/index.html • Le Couteur P & Burreson J. (2003). Napoleon’s buttons: how 17 molecules changed history. New York(NY): Jeremy P. Tarcher/Putnam.

45. References • Leyval, C., and P. Binet. (1998). "Effect of polyaromatic hydrocarbons in soil on arbuscular mycorrhizal plants." Journal of Environmental Quality 27.2: 402-7. Applied Science Full Text. Web. 13 Feb. 2010. • Margulis, L., and R. Fester (Eds.). (1991). Symbiosis as a source of evolutionary innovation : speciation and morphogenesis. MIT Press, Cambridge Mass. Retrieved from: http://books.google.com/books?id=3sKzeiHUIUQC&printsec=frontcover#v=onepage&q= &f=false • Silliman, B. R., & Newell, S. Y. (2003). Fungal Farming in a Snail. Proceedings of the National Academy of Sciences of the United States of America, 100(26), 15643-15648.   • Stamets, P. (2005). Mycelium running : how mushrooms can help save the world. Ten Speed Press, Berkeley Calif. Retrieved from: http://books.google.com/books?id=NPI8_omzvsC&dq=mycelium+running&printsec=front cover&source=bn&hl=en&ei=2PN5S93XOZCqNpuayLQH&sa=X&oi=book_result&ct=res ult&resnu m=6&ved=0CCIQ6AEwBQ#v=onepage&q=&f=false • Virginia Bioinformatics Institute. (2004). “Researcher from VBI advances fight against sudden oak death disease.” Retrieved from: https://www.vbi.vt.edu/public_relations/press_releases/sudden_oak_death_disease • Wang, B., Yeun, L. H., Xue, J., Liu, Y., An, J., & Qiu, Y. (2009). Presence of three mycorrhizal genes in the common ancestor of land plants suggests a key role of mycorrhizas in the colonization of land by plants. New Phytologist, 9999(9999). Retrieved from: http://dx.doi.org/10.1111/j.1469- 8137.2009.03137.x.   • Whelan A. & Rock, S. (2006). Phytotechnology Expediates Removal of Oil Waste from Shoreline Sediment.” EPA: Technology News and Trends. Retrieved from: http://www.clu in.org/products/newsltrs/tnandt/view.cfm?issue=0506.cfm#4