Plant and Soil

, Volume 185, Issue 1, pp 75–97 | Cite as

The ecological significance of plasticity in root weight ratio in response to nitrogen: Opinion

  • H. L. Reynolds
  • C. D'Antonio
Opinion Articles


We analyzed data on root weight ratio from a range of experimental studies documenting plant allocation changes in response to altered nitrogen availability. Our goal was to determine the degree to which plasticity in allocation between roots and shoots exists and to search for patterns in such plasticity among species. Our survey included 77 studies representing 206 cases and 129 species. As expected, we found that root weight ratio decreased with increased nitrogen availability in the majority of cases examined, and this response was most consistent when plants were grown individually or in intraspecific competition (versus interspecific competition). Surprisingly, however, we found no evidence to support existing hypotheses that fast-growing species adapted to high soil fertilities exhibit the highest levels of morphological plasticity, or that plasticity is positively associated with competitive ability. Rather, we found that average amounts of plasticity in root weight ratio in response to nitrogen availability were similar among species grouped by maximum relative growth rate and habitat fertility. Similar results were obtained for species categorized by life form, life history or root weight ratio itself, and plasticity in root weight ratio also had no consistent relationship with competitive ability. Numerous difficulties are associated with the attempt to search for pattern using independent studies, however our results lead to the conclusion that strong patterns in plasticity of root weight ratio in response to nitrogen availability among species do not exist. We discuss two reasons for this: (1) the costs of plasticity relative to its benefits are lower than previously predicted and (2) plasticity in traits other than root weight ratio is more important to plant foraging ability.

Key words

allocation competitive ability nitrogen availability plasticity root weight ratio 


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  1. Aerts R, Boot R G A and van der Aart P J M 1991 The relation between above- and belowground biomass allocation patterns and competitive ability. Oecologia 87, 551–559.Google Scholar
  2. Aerts R, de Caluwe H and Konings H 1992 Seasonal allocation of biomass and nitrogen in fourCarex species from mesotrophic and eutrophic fens as affected by nitrogen supply. J. Ecol. 80, 653–664.Google Scholar
  3. Ågren G I and Ingestad T 1987 Root:shoot ratio as a balance between nitrogen productivity and photosynthesis. Plant Cell Environ. 10, 579–586.Google Scholar
  4. Atkinson C J 1985 Nitrogen acquisition in four co-existing species from an upland acidic grassland. Physiol. Plant. 63, 375–387.Google Scholar
  5. Bernston G M, Farnsworth E J and Bazzaz F A 1995 Allocation, within and between organs, and the dynamics of root length changes in two birch species. Oecologia 101, 439–447.Google Scholar
  6. Bhat K K S, Nye P H and Brereton A J 1979 The possibility of predicting solute uptake and plant growth response from independently measured soil and plant characteristics. VI. The growth and uptake of rape in solutions of constant nitrate concentration. Plant and Soil 53, 137–167.Google Scholar
  7. Bilbao B and Medina E 1990 Nitrogen-use efficiency for growth in a cultivated African grass and a native South American pasture grass. J. Biogeogr. 17, 421–425.Google Scholar
  8. Birk E M and Vitousek P M 1986 Nitrogen availability and nitrogen use efficiency in loblolly pine stands. Ecology 67, 69–79.Google Scholar
  9. Bloom A J, Chapin F S III and Mooney H A 1985 Resource limitation in plants-an economic analogy. Annu. Rev. Ecol. Syst. 16, 363–392.Google Scholar
  10. Boot R G A 1990 The significance of size and morphology of root systems for nutrient acquisition and competition.In Causes and Consequences of Variation in Growth Rate and Productivity of Higher Plants. Eds. H Lambers, M L Cambridge, H Konings and T L Pons. pp 299–311. SPB Academic Publishing, The Hague, the Netherlands.Google Scholar
  11. Boot R G A and Mensink M 1990 Size and morphology of root systems of perennial grasses from contrasting habitats as affected by nitrogen supply. Plant and Soil 129, 291–299.Google Scholar
  12. Boot R G A and den Dubbelden K C 1990 Effects of nitrogen supply on growth, allocation and gas exchange characteristics of two perennial grasses from inland dunes. Oecologia 85, 115–121.Google Scholar
  13. Boot R G A, Schildwacht P M and Lambers H 1992 Partitioning of nitrogen and biomass at a range of N-addition rates and their consequences for growth and gas exchange in two perennial grasses from inland dunes. Physiol. Plant. 86, 152–160.Google Scholar
  14. Bradshaw A D 1965 Evolutionary significance of phenotypic plasticity in plants. Adv. Genet. 13, 115–155.Google Scholar
  15. Brouwer R 1966 Root growth of grasses and cereals.In The Growth of Cereals and Grasses. Eds. F L Milthorpe and J D Ivins. pp 153–166. Butterworth and Co. Ltd., London, UK.Google Scholar
  16. Burke M K, Raynal D J and Mitchell M J 1992 Soil nitrogen availability influences seasonal carbon allocation patterns in sugar maple (Acer saccharum). Can. J. For. Res. 22, 447–456.Google Scholar
  17. Bulisani E A and Warner R L 1980 Seed protein and nitrogen effects upon seedling vigor in wheat. Agron. J. 72, 657–662.Google Scholar
  18. Caldwell M, Richards J, Johnson D, Nowak R and Dzurec R 1981 Coping with herbivory: photosynthetic capacity and resource allocation in two semi-aridAgropyron bunchgrasses. Oecologia 50, 14–24.Google Scholar
  19. Caloin M, Khodre A El and Atry M 1980 Effect of nitrate concentration on the root:shoot ratio inDactylis glomerata L. and on the kinetics of growth in the vegetative phase. Ann. Bot. 46, 165–173.Google Scholar
  20. Campbell B D, Grime J P and Mackey J M L 1991 A trade-off between scale and precision in resource foraging. Oecologia 87, 532–538.Google Scholar
  21. Chambers J C, MacMahon J A and Brown R W 1987 Response of an early seral dominant alpine grass and a late seral dominant alpine forb to N and P availability. Rec. Rev. Res. 6, 219–234.Google Scholar
  22. Chapin F S III 1980 The mineral nutrition of wild plants. Annu. Rev. Ecol. Syst. 11, 233–260.Google Scholar
  23. Chapin F S III 1988 Ecological aspects of plant mineral nutrition. Adv. Min. Nutr. 3, 161–190.Google Scholar
  24. Cheplick G P 1995 Genotype variation and plasticity of clonal growth in relation to nutrient availability inAmphibromus scabrivalvis. J. Ecol. 83, 459–468.Google Scholar
  25. Christie E K and Moorby J 1975 Physiological responses of semiarid grasses I. The influence of phosphorus supply on growth and phosphorus absorption. Aust. J. Agric. Res. 26, 423–436.Google Scholar
  26. Coughenour M, McNaughton S and Wallace L L 1985 Responses of an African graminoid (Themeda trandra Forsk) to frequent defoliation. nitrogen, and water: a limit to adaptation to herbivory. Oecologia 68, 105–110.Google Scholar
  27. Crick J C and Grime J P 1987 Morphological plasticity and mineral nutrient capture in two herbaceous species of contrasted ecology. New Phytol. 107, 403–414.Google Scholar
  28. Crist J W and Stout G J 1929 Relation between top and root size in herbaceous plants. Plant Physiol. 4, 63–85.Google Scholar
  29. Cromer R N, Wheeler A M and Barr N J 1984 Mineral nutrition and growth ofEucalyptus seedlings. N. Z. J. For. Sci. 14, 229–239.Google Scholar
  30. Cure J D, Israel D W and Rufty T W Jr 1988 Nitrogen stress effects on growth and seed yield of nonnodulated soybean exposed to elevated carbon dioxide. Crop Sci. 28, 671–677.Google Scholar
  31. Dakheel A J, Radosevich S R and Barbour M G 1993 Effect of nitrogen and phosphorus on growth and interference betweenBromus tectorum andTaeniatherum asperum. Weed Res. 33, 415–422.Google Scholar
  32. Davidson R L 1969 Effects of soil nutrients and moisture on root/shoot ratios inLolium perenne L. andTrifolium repens L. Ann. Bot. 33, 571–577.Google Scholar
  33. Elberse W Th and Berendse F 1993 A comparative study of the growth and morphology of eight grass species from habitats with different nutrient availabilities. Func. Ecol. 7, 232–229.Google Scholar
  34. Ericsson T 1981 Effects of varied nitrogen stress on growth and nutrition in threeSalix clones. Physiol. Plant. 51, 423–429.Google Scholar
  35. Fetene M, Möller I and Beck E 1993 The effect of nitrogen supply toUrtica dioica L. plants on the distribution of assimilate between shoot and roots. Bot. Acta 106, 228–234.Google Scholar
  36. Fichtner K and Schulze E-D 1992 The effect of nitrogen nutrition on growth and biomass partitioning of annual plants originating from habitats of different nitrogen availability. Oecologia 92, 236–241.Google Scholar
  37. Figiel C R, Collins B and Wein G 1995 Variation in survival and biomass of two wetland grasses at different nutrient and water levels over a six week period. Bull. Torrey Bot. Club 122, 24–29.Google Scholar
  38. Firbank L G, Watkinson A R, Norton L R and Ashenden T W 1995 Plant populations and global environmental change: the effects of different temperature, carbon dioxide and nutrient regimes on density dependence in populations ofVulpia ciliata. Func. Ecol. 9, 432–441.Google Scholar
  39. Freijsen A H J and Otten H 1993 Utilization of the ambient concentration as a criterion for steady-state after exponential growth: Some culture experiments with optimum or suboptimum N nutrition. Plant and Soil 151, 265–271.Google Scholar
  40. Garnier E, Koch G W, Roy J and Mooney H A 1989 Responses of wild plants to nitrate availability: Relationships between growth rate and nitrate uptake parameters, a case study with twoBromus species, and a survey. Oecologia 79, 542–550.Google Scholar
  41. Gebauer R L E, Reynolds J F and Tenhunen J D 1995 Growth and allocation of the arctic sedgesEriophorum angustifolium andE. vaginatum: effects of variable soil oxygen and nutrient availability. Oecologia 104, 330–339.Google Scholar
  42. Gleeson S K 1993 Optimization of tissue nitrogen and root-shoot allocation. Ann. Bot. 71, 23–31.Google Scholar
  43. Georgiadis N J, Ruess R W, McNaughton S J and Western D 1989 Ecological conditions that determine when grazing stimulates grass production. Oecologia 81, 316–322.Google Scholar
  44. Griffin K L, Winner W E and Strain B R 1995 Growth and dry matter partitioning in loblolly and ponderosa pine seedlings in response to carbon and nitrogen availability. New Phytol. 129, 547–556.Google Scholar
  45. Grime J P 1977 Evidence for the existence of three primary strategies in plants and its relevance to ecological and evolutionary theory. Am. Nat. 111, 1169–1194.Google Scholar
  46. Grime J P 1979 Plant strategies and vegetation processes. John Wiley and Sons, Chichester, UK. 222 p.Google Scholar
  47. Grime J P and Hunt R 1975 Relative growth-rate: its range and adaptive significance in a local flora. J. Ecol. 63, 393–422.Google Scholar
  48. Grime J P, Crick J C and Rincon J E 1986 The ecological significance of plasticity.In Plasticity in Plants. Eds. D H Jennings and A J Trewavas. pp 5–30. The Company of Biologists, Cambridge, UK.Google Scholar
  49. Grime J P, Campbell B D, Mackey J M L and Crick J C 1991 Root plasticity, nitrogen capture and competitive ability.In Plant Root Growth: an Ecological Perspective. Ed. D Atkinson. pp 381–397. Blackwell Scientific Publications, Oxford, UK.Google Scholar
  50. Gross K L, Peters A and Pregitzer K S 1993 Fine root growth and demographic responses to nutrient patches in four old-field plant species. Oecologia 95, 61–64.Google Scholar
  51. Gulmon S L and Chu C C 1981 The effects of light and nitrogen on photosynthesis, leaf characteristics, and dry matter allocation in the chaparral shrub,Diplacus aurantiacus. Oecologia 49, 207–212.Google Scholar
  52. Gurevitch J, Wilson P, Stone J L, Teese P and Stoutenburgh R J 1990 Competition among old-field perennials at different levels of soil fertility and available space. J. Ecol. 78, 727–744.Google Scholar
  53. Hansson M L 1994 Response ofAnthriscus silvestris (L.) Hoffm. to defoliation and different nitrogen supply levels. Swed. J. Agric. Res. 24, 21–29.Google Scholar
  54. Hansson M L and Goransson A 1993 Growth and biomass partitioning ofAnthriscus sylvestris (L.) Hoffm. andFestucu ovina (L.) at different relative addition rates of nitrogen. Plant and Soil 155/156, 187–190.Google Scholar
  55. Hawthorn W R and Cavers P B 1982 Dry weight and resource allocation patterns among individuals in populations ofPlantago major andP. rugelii. Can. J. Bot. 60, 2424–2439.Google Scholar
  56. Hendricks J J, Nadelhoffer K J and Aber J D 1993 Assessing the role of fine roots in carbon and nutrient cycling. Trends Ecol. Evol. 8, 174–178.Google Scholar
  57. Hilbert D W 1990 Optimization of plant root:shoot ratios and internal nitrogen concentration. Ann. Bot. 66, 91–99.Google Scholar
  58. Hilbert D W and Reynolds J 1991 A model allocating growth among leaf proteins, shoot structure and root biomass to produce balanced activity. Ann. Bot. 68, 417–425.Google Scholar
  59. Holopainen J K. Rikala R, Kainulainen P and Oksanen J 1995 Resource partitioning to growth, storage and defense in nitrogen-fertilized Scots pine and susceptibility of the seedlings to the tarnished plant bugLygus rugulipennis. New Phytol. 131, 521–532.Google Scholar
  60. Hunt R 1975 Further observations on root-shoot equilibria in perennial ryegrass (Lolium perenne L.). Ann. Bot. 39, 745–755.Google Scholar
  61. Hunt R and Nicholls A O 1986 Stress and the coarse control of growth and root-shoot partitioning in herbaceous plants. Oikos 47, 149–158.Google Scholar
  62. Hutchings M J and de Kroon H 1994 Foraging in plants: the role of morphological plasticity in resource acquisition. Adv. Ecol. Res. 25, 159–238.Google Scholar
  63. Ingestad T and Ågren G I 1991 The influence of plant nutrition on biomass allocation. Ecol. Applic. 1, 168–174.Google Scholar
  64. Ingestad T and Lund A-B 1979 Nitrogen stress in birch seedlings I. Growth technique and growth. Physiol. Plant. 45, 137–148.Google Scholar
  65. Jackson R B, Manwaring J H and Caldwell M M 1990 Rapid physiological adjustment of roots to localized soil enrichment. Nature 344, 58–60.Google Scholar
  66. Jackson R B and Reynolds H L 1996 Nitrate and ammonium uptake for single-and mixed-species communities grown at elevated CO2. Oecologia 105, 74–80.Google Scholar
  67. Jameson D 1963 Responses of individual plants to harvesting. Bot. Rev. 29, 532–593.Google Scholar
  68. Jarvis S G 1987 The effects of low, regulated supplies of nitrate and ammonium nitrogen on the growth and composition of perennial ryegrass. Plant and Soil 100, 99–112.Google Scholar
  69. Johnson I R and Thornley J H M 1987 A model of shoot:root partitioning with optimal growth. Ann. Bot. 60, 133–142.Google Scholar
  70. Karlsson P S and Nordell K O 1987 Growth ofBetula pubescens andPinus sylvestris seedlings in a subarctic environment. Func. Ecol. 1, 37–44.Google Scholar
  71. Karrou M and Maranville J W 1994 Response of wheat cultivars to different soil nitrogen and moisture regimes: I. Dry matter partitioning and root growth. J. Plant Nutr. 17, 729–744.Google Scholar
  72. Körner C and Renhardt U 1987 Dry matter partitioning and root length/leaf area ratios in herbacous perennial plants with diverse altitudinal distribution. Oecologia 74, 411–418.Google Scholar
  73. Kuiper D, Schuit J and Kuiper P J C 1988 Effects of internal and external cytokinin concentrations on root growth and shoot to root ratio ofPlantago major ssp.pleiosperma at different nutrient conditions. Plant and Soil 111, 231–236.Google Scholar
  74. Lambers H and Dijkstra P 1987 A physiological analysis of genotypic variation in relative growth rate: can growth rate confer ecological advantage?In Disturbance in Grasslands. Eds. J van Andel, J P Bakker and R W Snaydon. pp 237–252. Junk Publishers, Dordrecht, the Netherlands.Google Scholar
  75. Lajtha K and Klein M 1988 The effect of varying nitrogen and phosphorus availability on nutrient use byLarrea tridentata, a desert evergreen shrub. Oecologia 75, 348–353.Google Scholar
  76. Larigauderie A, Hilbert D W and Oechel W C 1988 Effect of CO2 enrichment and nitrogen availability on resource acquisition and resource allocation in a grass,Bromus mollis. Oecologia 77, 544–549.Google Scholar
  77. Ledig F T, Bormann F H and Wenger K F 1970 The distribution of dry matter growth between shoot and roots in loblolly pine. Bot. Gaz. 131, 349–359.Google Scholar
  78. Lieffers V J and Titus S J 1989 The effects of stem density and nutrient status on size inequality and resource allocation in lodgepole pine and white spruce seedlings. Can. J. Bot. 67, 2900–2903.Google Scholar
  79. Macduff H, Jarvis S C, Larsson C M and Oscarson P 1993 Plant growth in relation to the supply and uptake of NO3: A comparison between relative addition rate and external concentration as driving variables. J. Exp. Bot. 44, 1475–1484.Google Scholar
  80. McDonald A J S, Lohammar T and Ericsson A 1986 Growth responses to step-decreases in nutrient availability in small birch (Betula pendula Roth). Plant Cell Environ. 9, 427–432.Google Scholar
  81. McGraw J B and Chapin F S III 1989 Competitive ability and adaptation to fertile and infertile soils in twoEriophorum species. Ecology 70, 736–749.Google Scholar
  82. McNaughton S J, Wallace L and Coughenour M 1983 Plant adaptation in an ecosystem context: effects of defoliation, nitrogen, and water on growth of an African C4 sedge. Ecology 64, 307–318.Google Scholar
  83. McNaughton S J and Chapin F S III 1985 Effects of phosphorus nutrition and defoliation on C4 graminoids from the Serengeti plains. Ecology 66, 1617–1629.Google Scholar
  84. Mihaliak C A and Lincoln D E 1989 Plant biomass partitioning and chemical defense: response to defoliation and nitrate limitation. Oecologia 80, 122–126.Google Scholar
  85. Millard P and Neilsen G H 1989 The influence of nitrogen supply on the uptake and remobilization of stored N for the seasonal growth of apple trees. Ann. Bot. 63, 301–309.Google Scholar
  86. Mortimer S 1992 Root length/leaf area ratios of chalk grassland perennials and their importance for competitive interactions. J. Veg. Sci. 3, 665–672.Google Scholar
  87. Muller B and Garnier E 1990 Components of relative growth rate and sensitivity to nitrogen availability in annual and perennial species ofBromus. Oecologia 84, 513–518.Google Scholar
  88. Munson A D and Timmer V R 1990 Site-specific growth and nutrition of plantedPicea mariana in the Ontario Clay Belt. III. Biomass and nutrient allocation. Can. J. For. Res. 20, 1165–1171.Google Scholar
  89. Newman E I 1973 Competition and diversity in herbaceous vegetation. Nature 244, 310.Google Scholar
  90. Olff H, van Andel J and Bakker J P 1990 Biomass and shoot/root allocation of five species from a grassland succession series at different combinations of light and nutrient supply. Func. Ecol. 4, 193–200.Google Scholar
  91. Peuke A D, Hartung W and Jeschke W D 1994 The uptake and flow of C, N and ions between roots and shoots inRicinus communis L. II. Grown with low or high nitrate supply. J. Exp. Bot. 45, 733–740.Google Scholar
  92. Polley H W and Detling J K 1988 Herbivory tolerance ofAgropyron smithii populations with different grazing histories. Oecologia 77, 261–267.Google Scholar
  93. Poorter H 1990 Interspecific variation in relative growth rate: on ecological causes and physiological consequences.In Causes and Consequences of Variation in Growth Rate and Productivity of Higher Plants. Eds. H Lambers, M L Cambridge, H Konings and T L Pons. pp 45–68. SPB Academic Publishing, The Hague, the Netherlands.Google Scholar
  94. Poorter H, van de Vijver C A D M, Boot R G A and Lambers H 1995 Growth and carbon economy of a fast-growing and a slow-growing grass species as dependent on nitrate supply. Plant and Soil 171, 217–227.Google Scholar
  95. Poorter H and Remkes C 1990 Leaf area ratio and net assimilation rate of 24 wild species differing in relative growth rate. Oecologia 83, 553–559.Google Scholar
  96. Redente E F, Friedlander J E and McLendon T 1992 Response of early and late semiarid seral species to nitrogen and phosphorus gradients. Plant and Soil 140, 127–135.Google Scholar
  97. Reynolds J F and Chen J 1996 Modelling whole-plant allocation in relation to carbon and nitrogen supply: Coordination versus optimization: Opinion. Plant and Soil 185, 65.Google Scholar
  98. Reynolds J F and Thornley J H M 1982 A shoot:root partitioning model. Ann. Bot. 49, 585–597.Google Scholar
  99. Robinson D 1994 Tansley review no. 73, The responses of plants to non-uniform supplies of nutrients. New Phytol. 127, 635–674.Google Scholar
  100. Robinson D and Rorison I H 1988 Plasticity in grass species in relation to nitrogen supply. Func. Ecol. 2, 249–257.Google Scholar
  101. Roder W, Mason S C, Clegg M D, Doran J W and Kniep K R 1988 Plant and microbial responses to sorghum-soybean cropping systems and fertility management. Soil Sci. Soc. Am. J. 52, 1337–1342.Google Scholar
  102. Rozijn N A M G, Ernst W H O, Van Andel J and Nelissen H J M 1990 Growth responses to different levels of soil moisture and soil fertility in four winter annual species during their life-cycle. Flora 184, 303–312.Google Scholar
  103. Ruess R W 1988 The interaction of defoliation and nutrient uptake inSporobolus kentrophyllus, a short-grass species from the Serengeti Plains. Oecologia 77, 550–556.Google Scholar
  104. Rufty T W Jr, Rapier C D Jr and Huber S C 1984 Alterations in internal partitioning of carbon in soybean plants in response to nitrogen stress. Can. J. Bot. 62, 501–508.Google Scholar
  105. Ryle G J A, Arnott R A and Powell C E 1981 Distribution of dry weight between root and shoot in white clover dependent on N2 fixation or utilizing abundant nitrate nitrogen. Plant and Soil 60, 29–39.Google Scholar
  106. Ryser P and Lambers H 1995 Root and leaf attributes accounting for the performance of fast- and slow-growing grasses at different nutrient supply. Plant and Soil 170, 251–265.Google Scholar
  107. Sattelmacher B, Klotz F and Marschner H 1990 Influence of the nitrogen level on root growth and morphology of two potato varieties differing in nitrogen acquisition. Plant and Soil 123, 131–137.Google Scholar
  108. Schlichting C D 1986 The evolution of phenotypic plasticity in plants. Annu. Rev. Ecol. Syst. 17, 667–693.Google Scholar
  109. Schlichting C D and Levin D A 1984 Phenotypic plasticity of annual phlox: tests of some hypotheses. Am. J. Bot, 71, 252–260.Google Scholar
  110. Schlichting C D and Levin D A 1990 Phenotypic plasticity inPhlox. III. Variation among natural populations ofP. drummondii. J. Evol. Biol. 3, 411–428.Google Scholar
  111. Sievanen R, Hari P, Orava P J and Pelkonen P 1988 A model for the effect of photosynthate allocation and soil nitrogen on plant growth. Ecol. Model. 41, 55–65.Google Scholar
  112. Smolders E and Merckx R 1992 Growth and shoot:root partitioning of spinach plants as affected by nitrogen supply. Plant Cell Environ. 15, 795–807.Google Scholar
  113. Srutek M 1995 Growth responses ofUrtica dioica to nutrient supply. Can. J. Bot. 73, 843–851.Google Scholar
  114. Steinger T and Müller-Schärer H 1992 Physiological and growth responses ofCentaurea maculosa (Asteraceae) to root herbivory under varying levels of interspecific plant competition and soil nitrogen availability. Oecologia 91, 141–149.Google Scholar
  115. Stribley D P, Read D J and Hunt R 1975 The biology of mycorrhiza in the Ericaceae. New Phytol. 75, 119–130.Google Scholar
  116. SYSTAT 1992 Statistics. Version 5.2.1. SYSTAT, Inc., Evanston, IL, USA.Google Scholar
  117. Tilman D and Wedin D 1991 Plant traits and resource reduction for five grasses growing on a nitrogen gradient. Ecology 72, 685–700.Google Scholar
  118. Troughton A 1956 Studies on the growth of young grass plants with special reference to the relationship between the shoot and root systems. J. Br. Grassl. Soc. 6, 56–65.Google Scholar
  119. Van de Vijver C A D M, Boot R G A, Poorter H and Lambers H 1993 Phenotypic plasticity in response to nitrate supply of an inherently fast-growing species from a fertile habitat and an inherently slow-growing species from an infertile habitat. Oecologia 96, 548–554.Google Scholar
  120. Van der Werf A, van Nuenen M, Visser A J and Lambers H 1993 Contribution of physiological and morphological plant traits to a species' competitive ability at high and low nitrogen supply: A hypothesis for inherently fast- and slow-growing monocotyle-donous species. Oecologia 94, 434–440.Google Scholar
  121. Van der Werf A, Visser A J, Schieving F and Lambers H 1993 Evidence for optimal partitioning of biomass and nitrogen at a range of nitrogen availabilities for a fast- and slow-growing species. Func. Ecol. 7, 63–74.Google Scholar
  122. Van der Werf A and Nagel O W 1996 Carbon allocation to shoots and roots in relation to nitrogen supply is mediated by cytokinins and sucrose: Opinion. Plant and Soil 185, 21.Google Scholar
  123. Vose P B 1962 Nutritional responses and shoot/root ratio as factors in the composition and yield of genotypes of perennial ryegrass,Lolium perenne L. Ann. Bot. N.S. 26, 425–437.Google Scholar
  124. Wallace L L, McNaughton S J and Coughenour M B 1982 The effects of clipping and fertilization on nitrogen nutrition and allocation by mycorrhizal and nonmycorrhizalPanicum coloratum L., a C4 grass. Oecologia 54, 68–71.Google Scholar
  125. Wilson J B 1988 A review of evidence on the control of shoot:root ratio, in relation to models. Ann. Bot. 61, 433–449.Google Scholar
  126. Wilson S D 1991 Plasticity, morphology and distribution in twelve lakeshore plants. Oikos 62, 292–298.Google Scholar
  127. Witowski E T F 1991 Growth and competition between seedlings ofProtea repens (L.) L. and the alien invasive,Acacia saligna (Labill.) Wendl. in relation to nutrient availability. Func. Ecol. 5, 101–110.Google Scholar
  128. Yeager T H and Wright R D 1981 Influence of nitrogen and phosphorus on shoot:root ratio ofIlex crenta Thunb. ‘Helleri’. HortScience 16, 564–565.Google Scholar
  129. Zangerl A R and Bazzaz F A 1983 Responses of an early and a late successional species ofPolygonum to variations in resource availability. Oecologia 56, 397–404.Google Scholar

Copyright information

© Kluwer Academic Publishers 1996

Authors and Affiliations

  • H. L. Reynolds
    • 1
    • 2
  • C. D'Antonio
    • 1
    • 2
  1. 1.W.K. Kellogg Biological StationHickory CornersUSA
  2. 2.Department of Integrative BiologyUniversity of CaliforniaBerkeleyUSA

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