1. Soil Ecology and Tree Health: Implications for Management of Urban Forests and Ornamental Landscapes Dan Herms Department of Entomology The Ohio State University Ohio Agricultural Research and Development Center herms.2@osu.edu

2. Acknowledgements: Students and Post-Docs Jim Blodgett, Rodrigo Chorbadjian, Carolyn Glynn, Bethan Hale, Nate Kleczewski, Joe LaForest, John Lloyd, Marie Egawa Collaborators Enrico Bonello, Robert Hansen, Harry Hoitink, Bill Mattson, Ben Stinner Funding Sources: TREE Fund USDA National Urban Community Forestry Advisory Council

3. The Ornamental Landscape as an Ecosystem: Implications for Pest Management Herms et al. (1984) J. Arboriculture 10:303-307. “Understanding the ecological interactions between the biotic and abiotic factors within a landscape enables more effective management of pests.”

4. Objective: Understand how trees allocate their resources in different environments, and the implications for the care of trees. Approach: Develop framework based on carbon allocation that can be used to predict tree behavior in different environments. Conduct experiments to test this framework.

5. Framework: carbon allocation patterns of trees Herms, D.A. 2002. Effects of fertilization on insect resistance of woody ornamental plants: reassessing an entrenched paradigm. Environmental Entomology 31:923-933. Herms, D.A. 2001. Resource allocation trade-offs in trees. Arborists News 10(5):41-47. Herms, D.A. 2001. Fertilization and pest control. Tree Care Industry 12(5):8-10,12,14. Herms, D.A. 1998. Understanding tree responses to abiotic and biotic stress complexes. Arborist News 7(1):9-15. Herms, D.A., and W.J. Mattson. 1997. Trees, stress, and pests. pp. 13-25. In J.E. Lloyd, ed. Plant Health Care for Woody Ornamentals, International Society of Arboriculture, Savoy, IL. Herms, D.A., and W.J. Mattson. 1992. The dilemma of plants: to grow or defend. Quarterly Review of Biology 67(3):283-335.

6. Different patterns of resource allocation and acquisition work in different environments

8. Concepts to emphasize: resource acquisition vs. resource allocation carbon budgets and allocation tradeoffs integration of above- and below-ground growth acclimation to stressful environments

9. Resource acquisition vs. allocation (Income vs. budgeting)

10. Whole plant carbon budget: photosynthesis rate per unit leaf area total leaf area

11. Allocation tradeoffs: plants have limited resources to support: growth maintenance reproduction storage defense

12. The chemical arsenal of plants: defense and stress tolerance Tannins Phenolic glycosides Terpenes Alkaloids Cyanogenic compounds Defensive proteins

13. Resource allocation patterns In faster growing plants: high allocation to total leaf area high photosynthesis rate lower allocation to root growth lower levels of defensive compounds In slower growing plants: lower allocation to total leaf area high photosynthesis rate higher allocation to root growth higher levels of defensive compounds

14. Computer-controlled fertigation system to study responses of willow to nutrient availability Glynn et al. (2007) New Phytologist 176:623-634

18. Nutrient availability and carbon acquisition: no effect on photosynthesis rate / leaf area increased total leaf area

19. Source / Sink Interactions: carbon moves from sources to sinks via phloem transport

23. Mechanisms of photosynthetic acclimation: Nitrogen allocation and specific leaf mass Root:shoot ratios

26. Nitrogen deficiency does not cause chlorosis in plants that have had time to acclimate to their environment. Harris, R. W. 1992. Root- shoot ratios. J. Arboriculture 18: 39-42

28. Stable root:shoot ratios between days 40-85 consistent with equilibrium patterns of resource allocation

29. Soil fertility and insect resistance: “Properly fertilized trees are better able to ward off both insect and disease damage." “Fertilizing landscape plants promotes their general health and vitality, making them more resistant to insect and disease attack." "Fertilization promotes vigorous growth, disease, and insect resistance, and stress tolerance."

30. Fertilization decreased the insect resistance of woody plants in almost every study. No study showed increased resistance. Herms, D.A. 2002. Effects of fertilization on insect resistance of woody ornamental plants: reassessing an entrenched paradigm. Environmental Entomology 31:923-933.

31. Field Studies: Effects of fertilization on paper birch and red pine

32. Fertilizer Treatment (ANSI standard): Rate:4.1 lb N / 1000 ft 2 / yr 200 kg N / ha / yr 178 lb N / acre / yr Formulation: 18:5:4 NPK (56 % N slow release) Timing: early May and mid-Sept (split application)

36. Fertilization increased growth of Sphaeropsis tip blight lesions by 50% Blodgett et al. 2005. Forest Ecology and Management 208:273-382.

37. Effects of nursery fertility regime on crabapple following transplanting 1998: transplanted to low maintenance landscape. 1997: three fertility treatments in container nursery (Lloyd, J.E., et al. 2006. HortScience 41:442-445)

41. Greater impact of drought stress on photosynthesis of high fertility plants.

42. Fertilization and stress tolerance: decreased root:shoot ratio increased water requirements decreased secondary metabolites

43. Fertilization decreased drought stress tolerance: Red oak, chestnut oak (Kleiner et al. 1992) American elm (Walters and Reich 1989) Monterey pine (Linder et al. 1987) Red pine (Miller and Timmer 1994) Loblolly pine (Green et al. 1994) Scots pine (Nilsen 1990) Norway spruce (Nilsen 1995)

44. Nutrient cycling in a forest: the ultimate slow release fertilizer

45. Disrupted nutrient cycles in constructed landscapes

46. Soil quality: the central role of organic matter (SOM) Key determinant of soil structure: oxygen, drainage, water / nutrient holding capacity. Source of essential nutrients for plants. Foundation of soil food web. Continuously depleted and replenished.

47. In an average cup of healthy soil: Bacteria: 200 billion Fungi: 60 miles of hyphae Protozoa: 20 million Nematodes: 100,000 Arthropods: 50,000 The living soil: From: S. Frey, Ohio State University

48. Most labile N is tied up by microbes

49. Nutrient Flow: The Central Role of Microbes Organic Matter Soil Microbes Plants

50. Mineral N (NH 4,, NO 3 ) Immobilization Organic N Organic Matter (Mulch) Mineralization Plant Uptake Decomposition Microbial Uptake Nutrient Cycling in Ornamental Landscapes Fertilizer Microbial Turnover

51. Key principles of nutrient cycling theory: Microbes are C limited. Plants are N limited. Microbes out-compete plants for N. High C:N organic matter: greater proportion of N immobilized by microbes. Low C:N organic matter: greater proportion of N released (mineralized) by microbes.

53. Objective: Establish general principles for predicting effects of diverse sources of organic matter on soil fertility and plant health.

54. How does mulch affect nutrient availability? : 1. Is the carbon available? Stability of OM 2. Who gets the nitrogen? C:N ratio of OM

55. Availability of C for microbes: rate of decomposition Slow Inorganic mulch (stone, shredded tires) Softwood bark (mature trees) Softwood bark (immature trees) Hardwood bark Ground wood Wood chips Composted yard waste Sawdust Composted Manure Fast

56. N available for plants determined by net balance between: N mineralization by microbes. N immobilization by microbes.

57. C:N Ratio of OM and Nutrient Availability: C:N ratio > 30:1 Microbes N-limited, scavenge N from soil Available N tied up by microbes N available for plants decreases C:N ratio < 30:1 N exceeds microbial requirements N release rates increase N available for plants increases

58. Material C:N Ratio Recycled pallets125:1 Ground pine bark105:1 Fresh wood chips 95:1 Hardwood bark 70:1 Fresh wood chips w/ foliage 65:1 Pine straw 64:1 Freshly senesced leaves 55:1 Composted wood chips 40:1 Composted yard waste 17:1 Composted manure 12:1

59. Case study: effects of mulch on soil microbes, nutrient cycling, and plant health. Recycled organic wastes: Composted yard waste (C:N = 17:1) Ground pallets (C:N = 125:1)

60. Recyled organic waste as mulch

61. Experimental Mulches Ground Wood Pallets C:N ratio = 125:1 Composted Yard Trimmings C:N ratio = 17:1

62. Composted mulch Ground wood pallets

63. Experimental approach:

64. Three Mulch Treatments: 1. Composted yard waste (C:N ratio = 17:1) 2. Ground wood pallets (C:N ratio = 125:1) 3. Bare soil control Each with and without fertilization (18-5-4 NPK, 3 lbs N / 1000 ft 2 / yr)

68. Mulch effects on tree growth

72. Nitrate as signaling molecule: gene expression and regulation of carbon allocation in Arabidopsis: High soil nitrate: Up regulation of genes for shoot growth, protein synthesis. Down regulation of genes for secondary metabolism, root growth. Low soil nitrate: Down regulation of genes for shoot growth, protein synthesis. Up regulation of genes for secondary metabolism, root growth. Scheible, et al. 2004. Plant Physiology 136:2483-2499. Zhang and Forde. 2000. Journal of Experimental Botany 51: 51-59.

73. Hypothesis: trees are adapted to the nutrient fluxes and signals associated with gradual decomposition of leaf litter, including low nitrate levels and high organic N sources. Can trees be tricked into maladaptive allocation patterns?

74. Fall Webworm Japanese Beetle

75. Trophic cascade from microbes through plants to insect herbivores: Organic Matter Microbe Effects on Nutrient Availability Plant Growth and Defense Plant-Feeding Insects

78. Conclusions: 1. Both mulches increased: soil organic matter microbial biomass and activity 2. Yard waste increased, ground wood decreased: nutrient availability plant growth susceptibility to insects

79. How can mulch applied to the soil surface affect nutrient availability below? 1. Soil homogenization by abiotic and biotic forces (rapid increase in SOM in mulched plots) 2. Subterranean foraging by hyphae of fungi that have colonized the mulch. 3. It just does.

80. Consistent with hypotheses: 1. Soil microbes are carbon-limited. 2. Plants are nitrogen limited. 3. Microbes out-compete plants for nitrogen. 4. Competition for N mediated by C:N ratio of OM. 5. Trade-off between growth and defense in plants.

81. Prescription mulching: Low C:N mulch (e.g. composted yard trimmings): degraded soils increased plant growth new landscapes High C:N mulch (e.g. recycled pallets): slow to moderate growth established plantings

82. Mulch volcanoes are not good for trees!

83. If you must make volcanoes, at least keep the mulch in the bags

84. Ecological Interactions in Sub-Soil

85. Sub Soil Top Soil Organic Matter (%) 0.75 2.24 Clay (%) 24 17 Total N (ppm) 5601790 Nitrate N (ppm) 8 161 Phosphorus (ppm) 8 50 Comparison of Sub-Soil and Top Soil Plots:

86. In subsoil: Fertilization increased growth and decreased phenolic compounds. In topsoil: Fertilization had no effect on growth or phenolics.

87. Fertilization effects on fall webworm

88. Mycorrhizae research: 1. Root colonization in subsoil 2. Effects of fertilizer 3. Interactions between native and commercial mycorrhizae Nate Kleczewski

89. Allocation to mycorrhizae Benefits: phosphorus acquisition organic nitrogen uptake increased drought tolerance increased resistance to root disease Costs: up to 40% of carbon assimilated by the plant.

90. Symbiosis (living together): mutualism parasitism Under some conditions, mycorrhizal fungi act as parasites, taking more than they give.

91. Plants can suppress mycorrhizae when costs exceed benefits: High nutrient availability (should mycorrhizal spores be applied with fertilizer?) Low carbon availability (e.g. shade, defoliation)

92. Questions regarding commercial mycorrhizae: Will they establish? Will they compete with native mycorrhizal fungi? Will they enhance plant growth and survival?

93. * Estimated number of viable propagules Key Findings: 1. Only native EMF were detected. 2. No difference between top-soil and sub-soil. *

94. * Key Findings: 3. High fertility suppressed mycorrhizae.

95. Key Findings: 1. Only native EMF were detected. 2. No difference between top-soil and sub-soil. 3. High fertility suppressed mycorrhizae.

96. Conclusions: Increased soil fertility: Increases growth Decreases chemical defenses Decreases root:shoot ratio Can decrease pest resistance Can decreases drought stress tolerance

97. Effects are independent of nutrient source, form, or timing: container fertigation conventional fertilization (ANSI guidelines) stored nutrients obtained the previous year mulch effects on soil microbes and nutrient cycling inherently fertile vs. infertile soil.

98. This doesn’t mean: fertilization is bad. fertilization will increase pest problems in landscapes (these studies haven’t been done yet). This does mean: The data do not support the conventional wisdom that fertilization increases pest resistance.

99. The Natural Tree Environment: nutrient limited soils frequent episodes of drought stress insects and pathogens The Natural Tree Response: high root:shoot ratios high levels of storage carbohydrates high levels of defensive chemicals moderate growth