Journal of Chemical Ecology

, Volume 16, Issue 4, pp 1111–1130 | Cite as

Seasonal patterns of juglone in soil beneathJuglans nigra (black walnut) and influence ofJ. nigra on understory vegetation

  • Bettina de Scisciolo
  • Donald J. Leopold
  • Daniel C. Walton


The allelopathic nature ofJ. nigra L. was investigated in several planted mixed hardwood stands located near Syracuse, New York. Concentrations of chloroform-extracted juglone from soil collected beneathJ. nigra was determined by thin-layer chromatography (TLC) and high-pressure liquid chromatography (HPLC). Soil juglone concentrations were corrected based on recovery of synthetic juglone added to soil. Soil juglone levels were high in the spring, decreased during the summer, and were high again in the fall. The quantification of juglone from soil by HPLC was found to be more accurate than by TLC. Regression analysis indicated that individual tree variation in soil juglone levels could not be explained by differences in soil moisture, pH, organic matter content, and texture. The results of juglone recovery experiments suggest that chloroform-extractable juglone does not persist in soil. Juglone degradation by microorganisms could only explain a portion of the juglone decline. Ordinations revealed that the herbaceous and woody vegetation beneathJ. nigra, in comparison to vegetation beneathAcer saccharum andQuercus rubra, is distinct in only one of the four stands studied. This vegetational difference did not appear to be a consequence of any strong allelopathic influences ofJ. nigra (Scheffe's method of contrast, chi-square analysis). The allelopathic nature of juglone under these field conditions is questionable.

Key words

Allelopathy black walnut Jugions nigra L. juglone 5-hydroxy-1,4-naphthoquinone 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Al-Naib, F.A.G., andRice, E.L. 1971. Allelopathic effects ofPlatanus occidentalis. Bull.Torrey Bot. Club 98:75–82.Google Scholar
  2. Beals, E.W. 1984. Bray-Curtis ordination: An effective strategy for analysis of multivariate ecological data.Adv. Ecol. Res. 14:1–55.Google Scholar
  3. Beatty, S.W. 1984. Influence of microtopography and canopy species on spatial patterns of forest understory plants.Ecology 65:1406–1419.Google Scholar
  4. Biswell, H.H. 1935. Effects of environment upon the root habits of certain deciduous forest trees.Bot. Gaz. 96:676–708.Google Scholar
  5. Boes, T.K. 1986. Allelopathy: Chemical interactions between plants.Am. Nurs. 163:67–72.Google Scholar
  6. Borner, H. 1960. Liberation of organic substances from higher plants and their role in the soil sickness problems.Bot. Rev. 26:393–424.Google Scholar
  7. Bratton, G.F. 1974. The role and allelopathic effects ofJuglans nigra in old field succession. MS thesis. Emporia Kansas State College, Emporia, Kansas.Google Scholar
  8. Bratton, S.P. 1976. Resource division in an understory herb community: Responses to temporal and microtopographic gradients.Am. Nat. 110:679–693.Google Scholar
  9. Brooks, M.G. 1951. Effects of black walnut trees and their products on other vegetation. Agricultural Extension State Bulletin No. 347, West Virginia University, Morgantown.Google Scholar
  10. Camp, R.F. 1986. Walnuts and white pine can be grown together successfully.Tree Plant. Notes U.S. Dept. Agric. For. Serv. 37:29–31.Google Scholar
  11. Chou, C.H., andKuo, Y.L. 1986. Allelopathic research of subtropical vegetation in Taiwan III. Allelopathic exclusion of understory byLeucaena leucocephala (Lam) de Wit.J. Chem. Ecol. 12:1431–1448.Google Scholar
  12. Chou, C.H., andYang, C.M. 1982. Allelopathic research of subtropical vegetation in Taiwan II. Comparative exclusion of understory byPhyllostachys edulis andCryptomeria japonica.J. Chem. Ecol. 8:1489–1507.Google Scholar
  13. Cline, S., andNeely, D. 1984. Relationship between juvenile-leaf resistance to anthracnose and the presence of juglone and hydrojuglone glucoside in black walnut.Phytopathology 74:185–188.Google Scholar
  14. Coder, K.D. 1983. Seasonal changes of juglone potential in leaves of black walnut (Juglans nigra L.).J. Chem. Ecol. 8:1203–1212.Google Scholar
  15. Collins, B.S., Ferrara, L.S., andMotto, H.L. 1984. Coincidence of spring herb distribution and flowering with tree bases in a New Jersey Piedmont forest.Bull. Torrey Bot. Club 111:301–306.Google Scholar
  16. Collins, S.L., andGood, R.E. 1986. Canopy-ground layer relationships of oak-pine forests in the New Jersey Pine Barrens.Am. Midl. Natl. 117:280–288.Google Scholar
  17. Cook, M.T. 1921. Wilting caused by walnut trees.Phytopathology 11:346.Google Scholar
  18. Crozier, C.R., andBoerner, R.E. 1984. Correlations of understory herb distribution patterns with microhabitats under different tree species in a mixed mesophytic forest.Oecologia (Berl.) 62:337–343.Google Scholar
  19. Crozier, C.R., andBoerner, R.E. 1986. Stemflow induced soil nutrient heterogeneity in a mixed mesophytic forest.Bartonia 52:1–8.Google Scholar
  20. Daglish, C. 1950. The determination and occurrence of a hydrojuglone glucoside in the walnut.Biochem. J. 47:458–462.Google Scholar
  21. Dalton, B.R., Blum, U., andWeed, S.B. 1983. Allelopathic substances in ecosystems: Effectiveness of sterile soil components in altering recovery of ferulic acid.J. Chem. Ecol. 9:1185–1201.Google Scholar
  22. Davidson, J. 1939. Some unanswered questions.North. Nut Grow. Assoc. Ann. Rep. 30:58–66.Google Scholar
  23. Davis, E.F. 1928. The toxic principle ofJuglans nigra as identified with synthetic juglone, and its toxic effects on tomato and alfalfa plants.Am. J. Bot. 15: 620.Google Scholar
  24. Dawson, J.O., andSeymour, P.E. 1983. Effects of juglone concentration on growth in vitro ofFrankia ArI3 andRhizobium japonicum strain 71.J. Chem. Ecol. 9:1175–1183.Google Scholar
  25. del Moral, R., andMuller, C.H. 1970. The allelopathic effects ofEucalyptus camaldulensis.Am. Midl. Nat. 83:254–282.Google Scholar
  26. De Scisciolo, B. 1988. The seasonal patterns of juglone in soil beneathJuglans nigra (black walnut) and the influence ofJ. nigra on understory vegetation. MS thesis. State University of New York College of Environmental Science and Forestry, Syracuse, New York.Google Scholar
  27. Everett, R.L., Sharrow, S.H., andMeeuwig, R.O. 1983. Pinyon-juniper woodland understory distribution patterns and species associations.Bull. Torrey Bot. Club 110:454–463.Google Scholar
  28. Fisher, R.F. 1978. Juglone inhibits pine growth under certain moisture regimes.Soil Sci. Soc. Am. J. 42:801–803.Google Scholar
  29. Friedham, E.A.H. 1934. Natural reversible oxidation-reduction systems as accessory catalysts in respiration: juglon and lawson.Biochem. J. 28:180–188.Google Scholar
  30. Fuerst, E.P., andPutnam, A.R. 1983. Separating the competitive and allelopathic components of interference: theoretical principles.J. Chem. Ecol. 9:937–944.Google Scholar
  31. Funk, D.T., Case, P.J., Rietveld, W.J., andPhares, R.E. 1979. Effects of juglone on the growth of coniferous seedlings.For. Sci. 25:452–454.Google Scholar
  32. Gabriel, W.J. 1975. Allelopathic effects of black walnut on white birches.J. For. 73:234–237.Google Scholar
  33. Gersper, P.L., andHolowaychuk, N. 1970. Effects of stemflow water on a Miami soil under a beech tree: II Chemical properties.Soil Sci. Soc. Am. Proc. 34:786–794.Google Scholar
  34. Gersper, P.L., andHolowaychuk, N. 1971. Some effects of stem flow from forest canopy trees on chemical properties of soils.Ecology 52:691–702.Google Scholar
  35. Gleason, H.A., andCronquist, A. 1963. Manual of vascular plants of northeastern United States and adjacent Canada. D. Van Nostrand, New York.Google Scholar
  36. Graves, C.H., Hedin, P.A., andLanghans, V.E. 1979. A survey of juglone levels among pecan, hickory, and walnut, pp. 103–111,in: Proc. 72nd Annu. Conv. Southeast Pecan Growers Assoc., Hilton Head, South Carolina.Google Scholar
  37. Greene, K.W. 1930. The toxic (?) effect of the black walnut.North. Nut Grow. Assoc. Proc. 1929:152–157.Google Scholar
  38. Gries, G.A. 1943. Juglone—the active agent in walnut toxicity.North. Nut Grow. Assoc. Annu. Rep. 34:52–55.Google Scholar
  39. Hedin, P.A., Langhans, V.E., andGraves, C.H. 1979. Identification of juglone in pecan as a possible factor of resistance toFusicladium effusum.J. Agric. Food Chem. 27:92–94.Google Scholar
  40. Hedin, P.A., Collum, D.H., Langhans, V.E., andGraves, C.H. 1980. Distribution of juglone and related compounds in pecan and their effect onFusicladium effusum.J. Agric. Food Chem. 28:340–342.Google Scholar
  41. Hicks, D.J. 1980. Intrastand distribution patterns of southern Appalachian cove forest herbaceous species.Am. Midl. Nat. 104:209–223.Google Scholar
  42. Holch, A.E. 1931. Development of roots and shoots of certain deciduous tree seedlings indifferent forest sites.Ecology 12:259–298.Google Scholar
  43. Kershaw, K.A., andLooney, J.H.H. 1985. Quantitative and Dynamic Plant Ecology, 3rd ed. Edward Arnold, London.Google Scholar
  44. Lee, K.C., andCampbell, R.W. 1969. Nature and occurrence of juglone inJuglans nigra L. Hortic. Sci. 4: 297–298.Google Scholar
  45. Lodhi, M.A.K. 1976. Role of allelopathy as expressed by dominating trees in a lowland forest in controlling the productivity and pattern of herbaceous growth.Am. J. Bot. 63:1–8.Google Scholar
  46. Lodhi, M.A.K. 1977. The influence and comparison of individual forest trees on soil properties and possible inhibition of nitrification due to intact vegetation.Am. J. Bot. 64:260–264.Google Scholar
  47. Lodhi, M.A.K., andRice, E.L. 1971. Allelopathic effects ofCeltis laevigata.Bull. Torrey Bot. Club 98:83–89.Google Scholar
  48. MaCuaniels, L.H., andMuenscher, W.C. 1941. Black walnut toxicity.North. Nut Grow. Assoc. Annu. Rep. 31:172–179.Google Scholar
  49. MaCdaniels, L.H., andPinnow, D.L. 1976. Walnut toxicity, an unsolved problem.North. Nut Grow. Assoc. Annu. Rep. 67:114–122.Google Scholar
  50. Maguire, D.A., andForman, R.T.T. 1983. Herb cover effects on tree seedling patterns in a mature hemlock-hardwood forest.Ecology 64:1367–1380.Google Scholar
  51. Marking, L.L. 1970. Juglone (5-hydroxy-1,4-naphthoquinone) as a fish toxicant.Trans. Am. Fish Soc. 99:510–514.Google Scholar
  52. Massey, A.B. 1925. Antagonism of the walnuts (Juglans nigra L. andJ. cinerea L.) in certain plant associations.Phytopathology 15:773–784.Google Scholar
  53. Mattoon, H.G. 1944. A commercial black walnut venture.North. Nut Grow. Assoc. Annu. Rep. 35:79–82.Google Scholar
  54. McCune, B. 1987. Multivariate Analysis on the PC-ORD system. A Software Documentation Report. Holcomb Research Institute, Butler University, Indianapolis, Indiana.Google Scholar
  55. Oleszek, W., andJurzysta, M. 1987. The allelopathic potential of alfalfa root medicagenic acid glycosides and their fate in soil environments.Plant Soil 98:67–80.Google Scholar
  56. Perry, S.F. 1967. Inhibition of respiration by juglone inPhaseolus andLycopersicon.Bull. Torrey Bot. Club 94:26–30.Google Scholar
  57. Pirone, P.P. 1938. The detrimental effect of walnut to rhododendrons and other ornamentals.Nurs. Dis. Notes 11:13–16.Google Scholar
  58. Ponder, F., Jr. 1987. Allelopathic interference of black walnut trees with nitrogen-fixing plants in mixed plantings, pp. 195–204,in G.R. Waller (ed.). Allelochemicals: Role in Forestry and Agriculture. ACS Symposium Series No. 330. American Chemical Society, Washington, D.C.Google Scholar
  59. Ponder, F., Jr., andTadros, S.H. 1985. Juglone concentration in soil beneath black walnut interplanted with nitrogen-fixing species.J. Chem. Ecol. 11:937–942.Google Scholar
  60. Prichard, R.P. 1941. Report on experimental planting Syracuse Forest Experiment Station. Bull. NYS College of Forestry Tech. Publ.57, Syracuse, New York. 43 pp.Google Scholar
  61. Reinking, O.A. 1943. Possible black walnut toxicity on tomato and cabbage.North. Nut Grow. Assoc. Annu. Rep. 34:56–58.Google Scholar
  62. Rettenmaier, H.J. 1985. The enzymatic bacterial decomposition of juglone. PhD thesis. University of Hohenheim, Hohenheim, West Germany. 88 pp.Google Scholar
  63. Rettenmaier, H.J., Kupas, E., andLingens, F. 1983. Degradation of juglone byPseudomonas putida J 1.FEMS Microbiol. Lett. 19:193–195.Google Scholar
  64. Rice, E.L. 1984. Allelopathy, 2nd ed. Academic Press, Orlando, Florida.Google Scholar
  65. Richards, L.A. 1969. Diagnosis and improvement of saline and alkali soils. Agric. Handb. No. 60, USDA. U.S. Government Printing Office, Washington, D.C.Google Scholar
  66. Rietveld, W.J. 1981. The significance of allelopathy in black walnut cultural systems.North. Nut Grow. Assoc. Annu. Rep. 72:117–134.Google Scholar
  67. Rietveld, W.J. 1983. Allelopathic effects of juglone on germination and growth of several herbaceous and woody species.J. Chem. Ecol. 9:295–308.Google Scholar
  68. Rietveld, W.J., Schlesinger, R.C., andKessler, K.J. 1983. Allelopathic effects of black walnut on European black alder coplanted as a nurse species.J. Chem. Ecol. 9:1119–1133.Google Scholar
  69. Riha, S.J., James, B.R., Senesac, G.P., andPallant, E. 1986. Spatial variability of soil pH and organic matter in forest plantations.Soil Sci. Soc. Am. J. 50:1347–1352.Google Scholar
  70. Schneiderhan, F.J. 1927. The black walnut (Juglans nigra L.) as a cause of the death of apple trees.Phytopathology 17:529–540.Google Scholar
  71. Schreiner, E.J. 1949. Can black walnut trees poison pines?Penn. Univ. Morris Arboretum Bull. 4:94–96.Google Scholar
  72. Sherman, R.R. 1971. An ecological evaluation of the allelopathic influence ofJugions nigra onLycopersicon esculentum. MS thesis. Michigan State University, East Lansing, Michigan.Google Scholar
  73. Smith, R.M. 1942. Some effects of black locusts and black walnuts on southeastern Ohio pastures.Soil Sci. 53:385–398.Google Scholar
  74. Sokal, R.R., andRohlf, F.J. 1981. Biometry, 2nd ed. W.H. Freeman, San Francisco.Google Scholar
  75. Strong, M.C. 1944. Walnut wilt of tomato.Mich. Agric. Exp. Sta. Q. Bull. 26:194–195.Google Scholar
  76. Thomson, R.H. 1971. Naturally Occurring Quinones. Academic Press. New York. pp. 2–3, 13–21.Google Scholar
  77. Tukey, H.B. 1966. Leaching of metabolites from above-ground parts and its implications.Bull. Torrey Bot. Club 93:385–401.Google Scholar
  78. Turner, D.P., andFranz, E.H. 1986. The influence of canopy dominants on understory vegetation patterns in an old-growth cedar-hemlock forest.Am. Midi. Nat. 116:387–393.Google Scholar
  79. U.S. Department of Agriculture Soil Conservation Service. 1972. Soil survey of Onondaga County.Google Scholar
  80. Voigt, O.K. 1960. Distribution of rainfall under forest stands.For. Sei. 6:2–10.Google Scholar
  81. von Althen, F.W. 1968. Incompatibility of black walnut and red pine.Can. For. Serv. Bi-momh. Res. Notes 24:19.Google Scholar
  82. Werner, P.A., andHarbeck, A.L. 1982. The pattern of tree seedling establishment relative to staghorn sumac cover in Michigan old fields.Am. Midi. Nat. 108:124–132.Google Scholar
  83. Wiant, H.V., Jr., andRamirez, M.A. 1974. Don't plant white pine near walnut!Tree Plant Notes U.S. Dept. Agric. For. Serv. 25:30.Google Scholar
  84. Wilde, S.A., Voigt, G.K., andIyer, J.G. 1972. Soil and Plant Analysis for Tree Culture. Oxford and IBH Publishing, New Delhi, India.Google Scholar
  85. Wilde, S.A., Corey, R.B., Iyer, J.G., andVoigt, G.K. 1979. Soil and Plant Analysis for Tree Culture. Oxford and IBH Publishing, New Delhi, India.Google Scholar
  86. Willis, R.J. 1985. The historical bases of the concept of allelopathy.J. History Biol. 18:71–102.Google Scholar
  87. Wonnacott, R.J., andWonnacott, T.H. 1985. Introductory Statistics, 4th ed. John Wiley & Sons, New York.Google Scholar
  88. Yen, C.P.,Pham, C.H.,Cox, G.S., andGarrett, H.E. 1978. Soil depth and root development patterns of Missouri black walnut and certain Taiwan hardwoods, pp. 36–42,in Proc. of Symp. of Root Form of Planted Trees, Victoria, B.C., Canada, May 16–19.Google Scholar
  89. Zinke, P.J. 1962. The pattern of influence of individual forest trees on soil properties.Ecology 43:130–133.Google Scholar

Copyright information

© Plenum Publishing Corporation 1990

Authors and Affiliations

  • Bettina de Scisciolo
    • 1
  • Donald J. Leopold
    • 1
  • Daniel C. Walton
    • 1
  1. 1.SUNY College of Environmental Science and ForestrySyracuse

Personalised recommendations