Root competition: beyond resource depletion

Summary 1 Root competition is defined as a reduction in the availability of a soil resource to roots that is caused by other roots. Resource availability to competitors can be affected through resource depletion (scramble competition) and by mechanisms that inhibit access of other roots to resources (contest competition, such as allelopathy). 2 It has been proposed that soil heterogeneity can cause size-asymmetric root competition. Support for this hypothesis is limited and contradictory, possibly because resource uptake is affected more by the amount and spatial distribution of resource-acquiring organs, relative to the spatial distribution of resources, than by root system size per se . 3 Root competition intensity between individual plants generally decreases as resource availability (but not necessarily habitat productivity) increases, but the importance of root competition relative to other factors that structure communities may increase with resource availability. 4 Soil organisms play important, and often species-specific, roles in root interactions. 5 The findings that some roots can detect other roots, or inert objects, before they are contacted and can distinguish between self and non-self roots create experimental challenges for those attempting to untangle the effects of self/non-self root recognition, self-inhibition and root segregation or proliferation in response to competition. Recent studies suggesting that root competition may represent a ‘tragedy-of-the-commons’ may have failed to account for this complexity. 6 Theories about potential effects of root competition on plant diversity (and vice versa) appear to be ahead of the experimental evidence, with only one study documenting different effects of root competition on plant diversity under different levels of resource availability. 7 Roots can interact with their biotic and abiotic environments using a large variety of often species-specific mechanisms, far beyond the traditional view that plants interact mainly through resource depletion. Research on root interactions between exotic invasives and native species holds great promise for a better understanding of the way in which root competition may affect community structure and plant diversity, and may create new insights into coevolution of plants, their competitors and the soil community.

[1]  P. Keddy,et al.  Root and shoot competition intensity along a soil depth gradient , 1995 .

[2]  Robert B. Jackson,et al.  PLANT COMPETITION UNDERGROUND , 1997 .

[3]  Joel s. Brown,et al.  Intra-plant versus Inter-plant Root Competition in Beans: avoidance, resource matching or tragedy of the commons , 2002, Plant Ecology.

[4]  Thomas J. Givnish,et al.  On the economy of plant form and function. , 1988 .

[5]  D. Goldberg,et al.  Root competition can cause a decline in diversity with increased productivity , 2003 .

[6]  J. Lepš,et al.  Influence of soil heterogeneity and competition on growth features of three meadow species , 2004 .

[7]  R. K. Dixon,et al.  Effect of Container Size and Shape on the Growth of Northern Red Oak Seedlings , 1987, HortScience.

[8]  T. Schoener Presence and Absence of Habitat Shift in Some Widespread Lizard Species , 1975 .

[9]  B. Blair,et al.  Effect of soil nutrient heterogeneity on the symmetry of belowground competition , 2001, Plant Ecology.

[10]  Robert B. Jackson,et al.  DEFINING A PLANT'S BELOWGROUND ZONE OF INFLUENCE , 2003 .

[11]  C. Donald,et al.  The interaction of competition for light and for nutrients , 1958 .

[12]  J. Maron,et al.  A biogeographical approach to plant invasions: the importance of studying exotics in their introduced and native range , 2005 .

[13]  J. Cahill Interactions between root and shoot competition vary among species , 2002 .

[14]  E. Haugland,et al.  Improving grasslands: the influence of soil moisture and nitrogen fertilization on the establishment of seedlings , 1999 .

[15]  J. Harper Population Biology of Plants , 1979 .

[16]  J. Connell Apparent versus "real" competition in plants. , 1990 .

[17]  T. Rajaniemi Explaining productivity‐diversity relationships in plants , 2003 .

[18]  S. Wilson,et al.  Competition Intensity along a Productivity Gradient in a Low‐Diversity Grassland , 1998, The American Naturalist.

[19]  C. Welden,et al.  The Intensity of Competition Versus its Importance: An Overlooked Distinction and Some Implications , 1986, The Quarterly Review of Biology.

[20]  R. Callaway,et al.  Novel Weapons: Invasive Success and the Evolution of Increased Competitive Ability , 2004 .

[21]  T. Weir,et al.  Retracted: Dual role for an allelochemical: (±)‐catechin from Centaurea maculosa root exudates regulates conspecific seedling establishment , 2005 .

[22]  R. B. Jackson,et al.  Rooting depths, lateral root spreads and below‐ground/above‐ground allometries of plants in water‐limited ecosystems , 2002 .

[23]  F. Pugnaire,et al.  Changes in plant interactions along a gradient of environmental stress , 2001 .

[24]  E. Morris,et al.  How does fertility of the substrate affect intraspecific competition? Evidence and synthesis from self-thinning , 2003, Ecological Research.

[25]  Jacob Weiner,et al.  Are invasive plant species better competitors than native plant species? - evidence from pair-wise experiments , 2004 .

[26]  D. Tilman,et al.  PLANT COMPETITION AND RESOURCE AVAILABILITY IN RESPONSE TO DISTURBANCE AND FERTILIZATION , 1993 .

[27]  E. Salisbury Competition between Plants. , 1930, Nature.

[28]  W. Bouten,et al.  Towards understanding tree root profiles: simulating hydrologically optimal strategies for root distribution , 2001 .

[29]  D. Hooper THE ROLE OF COMPLEMENTARITY AND COMPETITION IN ECOSYSTEM RESPONSES TO VARIATION IN PLANT DIVERSITY , 1998 .

[30]  R. Callaway,et al.  The detection of neighbors by plants. , 2002 .

[31]  Joel s. Brown,et al.  Tragedy of the commons as a result of root competition , 2001 .

[32]  D. Coomes,et al.  RESPONSES OF JUVENILE TREES TO ABOVE‐ AND BELOWGROUND COMPETITION IN NUTRIENT‐STARVED AMAZONIAN RAIN FOREST , 1998 .

[33]  M. Austin,et al.  BEHAVIOUR OF EXPERIMENTAL PLANT COMMUNITIES ALONG A NUTRIENT GRADIENT , 1980 .

[34]  D. Goldberg,et al.  Patterns and Consequences of Interspecific Competition in Natural Communities: A Review of Field Experiments with Plants , 1992, The American Naturalist.

[35]  M. Litav,et al.  Root Competition Between Two Strains of Spinacia Oleracea. I. Pattern and Processes , 1974 .

[36]  Peter W. Barlow,et al.  Root apices as plant command centres: the unique 'brain-like' status of the root apex transition zone , 2004 .

[37]  J. Silvertown,et al.  Hydrologically defined niches reveal a basis for species richness in plant communities , 1999, Nature.

[38]  Joel s. Brown,et al.  Root proliferation and seed yield in response to spatial heterogeneity of below-ground competition. , 2005, The New phytologist.

[39]  M. Zobel,et al.  Can arbuscular mycorrhiza change the effect of root competition between conspecific plants of different ages , 1998 .

[40]  Jessica Gurevitch,et al.  EMPIRICAL APPROACHES TO QUANTIFYING INTERACTION INTENSITY: COMPETITION AND FACILITATION ALONG PRODUCTIVITY GRADIENTS , 1999 .

[41]  Jacob Weiner,et al.  Mechanisms determining the degree of size asymmetry in competition among plants , 1998, Oecologia.

[42]  K. Wiegand,et al.  Resource pulses, species interactions, and diversity maintenance in arid and semi-arid environments , 2004, Oecologia.

[43]  M. V. Iersel Root Restriction Effects on Growth and Development of Salvia (Salvia splendens) , 1997 .

[44]  Franz Rebele,et al.  Competition and coexistence of rhizomatous perennial plants along a nutrient gradient , 2000, Plant Ecology.

[45]  P. Alpert,et al.  Root cooperation in a clonal plant: connected strawberries segregate roots , 2002, Oecologia.

[46]  Jonathan M. Levine,et al.  Elton revisited: a review of evidence linking diversity and invasibility , 1999 .

[47]  L. Mommer,et al.  Root competition: towards a mechanistic understanding , 2003 .

[48]  H. Olff,et al.  Plant Competition in Relation to Neighbor Biomass: An Intercontinental Study with POA Pratensis , 1994, Ecology.

[49]  J. Cahill Neighbourhood‐scale diversity, composition and root crowding do not alter competition during drought in a native grassland , 2003 .

[50]  H. Kroon,et al.  Soil nutrient heterogeneity alters competition between two perennial grass species , 2001 .

[51]  H. J. Schenk,et al.  Spatial Root Segregation: Are Plants Territorial? , 1999 .

[52]  R. Sands,et al.  Competition for water and nutrients in forests , 1993 .

[53]  Ariel Novoplansky,et al.  Physiologically mediated self/non-self discrimination in roots. , 2004, Proceedings of the National Academy of Sciences of the United States of America.

[54]  J. Duval,et al.  The effect of container size , 1998 .

[55]  J. Fridley Diversity effects on production in different light and fertility environments: an experiment with communities of annual plants , 2003 .

[56]  José L. Hierro,et al.  Allelopathy and exotic plant invasion , 2003, Plant and Soil.

[57]  Fusuo Zhang,et al.  Effect of root contact on interspecific competition and N transfer between wheat and fababean using direct and indirect 15N techniques , 2004, Plant and Soil.

[58]  H. M. Alexander,et al.  Soil pathogenic fungi have the potential to affect the co‐existence of two tallgrass prairie species , 1999 .

[59]  J. Arnone,et al.  Root competition and elevated CO2: effects on seedling growth in Linum usitatissimum populations and Linum–Silene cretica mixtures , 1997 .

[60]  T. Semere,et al.  The effect of pea cultivar and water stress on root and shoot competition between vegetative plants of maize and pea , 2001 .

[61]  R. Callaway,et al.  Effects of Regional Origin and Genotype on Intraspecific Root Communication in the Desert Shrub Ambrosia Dumosa (Asteraceae) , 1996 .

[62]  M. Hutchings,et al.  Does pattern of soil resource heterogeneity determine plant community structure? An experimental investigation , 2005 .

[63]  D. Tilman,et al.  QUADRATIC VARIATION IN OLD-FIELD SPECIES RICHNESS ALONG GRADIENTS OF DISTURBANCE AND NITROGEN , 2002 .

[64]  B. Griffiths,et al.  Plant root proliferation in nitrogen–rich patches confers competitive advantage , 1999, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[65]  Pereira,et al.  Plant diversity and productivity experiments in european grasslands , 1999, Science.

[66]  B. E. Mahall,et al.  Positive and negative plant interactions contribute to a north-south-patterned association between two desert shrub species , 2002, Oecologia.

[67]  Angela Hodge,et al.  The plastic plant: root responses to heterogeneous supplies of nutrients , 2004 .

[68]  M. Austin A silent clash of paradigms : some inconsistencies in community ecology , 1999 .

[69]  M. Loik,et al.  A multi-scale perspective of water pulses in dryland ecosystems: climatology and ecohydrology of the western USA , 2004, Oecologia.

[70]  C. D’Antonio,et al.  Root profiles and competition between the invasive, exotic perennial, Carpobrotus edulis, and two native shrub species in California coastal scrub , 1991 .

[71]  Jonathan P Lynch,et al.  Optimization modeling of plant root architecture for water and phosphorus acquisition. , 2004, Journal of theoretical biology.

[72]  James B. Grace,et al.  Components of resource competition in plant communities. , 1990 .

[73]  M. Litav,et al.  Partial Separation of Roots as a Means of Reducing the Effect of Competition between Two Grass Species , 1971 .

[74]  H. Walter,et al.  Ecology of Southern Africa@@@Grasland, Savanne und Busch der arideren Teile Afrikas in ihrer okologischen Bedingtheit , 1941 .

[75]  K. J. Day,et al.  The effects of spatial pattern of nutrient supply on yield, structure and mortality in plant populations , 2003 .

[76]  James F. Reynolds,et al.  Plant responses to precipitation in desert ecosystems: integrating functional types, pulses, thresholds, and delays , 2004, Oecologia.

[77]  R. Callaway,et al.  Positive interactions among plants , 1995, The Botanical Review.

[78]  B. R. Christie,et al.  Beneficial microbial allelopathies in the root zone: the management of soil quality and plant disease with rhizobacteria , 2003 .

[79]  A review of belowground interactions in agroforestry, focussing on mechanisms and management options , 1999 .

[80]  L. Aarssen,et al.  On the density dependence of replacement-series competition experiments , 1989 .

[81]  B. Wolfe,et al.  Breaking New Ground: Soil Communities and Exotic Plant Invasion , 2005 .

[82]  C. Stern CONCLUDING REMARKS OF THE CHAIRMAN , 1950 .

[83]  Thomas W. Schoener,et al.  Field Experiments on Interspecific Competition , 1983, The American Naturalist.

[84]  Lauren D. Quinn,et al.  Phosphorus uptake, not carbon transfer, explains arbuscular mycorrhizal enhancement of Centaurea maculosa in the presence of native grassland species , 2002 .

[85]  Y. Kitamura,et al.  Legume Growth and Nitrogen Fixation as Affected by Plant Competition for Light and for Soil Nitrogen1 , 1981 .

[86]  J. Bastow Wilson,et al.  Evidence for limiting similarity in a sand dune community , 2004 .

[87]  A. Fitter Darkness visible: reflections on underground ecology , 2005 .

[88]  J. Gurevitch,et al.  Competition among old-field perennials at different levels of soil fertility and available space , 1990 .

[89]  A. Packer,et al.  Plant-soil biota interactions and spatial distribution of black cherry in its native and invasive ranges , 2003 .

[90]  Simon Gilroy,et al.  Allelopathy and Exotic Plant Invasion: From Molecules and Genes to Species Interactions , 2003, Science.

[91]  R. Callaway,et al.  Root herbivores, pathogenic fungi, and competition between Centaurea maculosa and Festuca idahoensis , 2003, Plant Ecology.

[92]  Density-dependent habitat selection in plants , 1998 .

[93]  D. Hertel,et al.  Tree species diversity and soil patchiness in a temperate broad-leaved forest with limited rooting space , 2002 .

[94]  H. Schenk Vertical Vegetation Structure Below Ground: Scaling from Root to Globe , 2005 .

[95]  A. Fitter Influence of mycorrhizal infection on competition for phosphorus and potassium by two grasses , 1977 .

[96]  M. Hutchings,et al.  TOWARD UNDERSTANDING THE CONSEQUENCES OF SOIL HETEROGENEITY FOR PLANT POPULATIONS AND COMMUNITIES , 2003 .

[97]  F. Bazzaz,et al.  IS PHYSICAL SPACE A SOIL RESOURCE , 1991 .

[98]  J. Facelli,et al.  Soil phosphorus heterogeneity and mycorrhizal symbiosis regulate plant intra-specific competition and size distribution , 2002, Oecologia.

[99]  Christopher J. Lortie,et al.  The importance of importance , 2005 .

[100]  F. Chapin,et al.  Consequences of changing biodiversity , 2000, Nature.

[101]  J. Vivanco,et al.  Natural selection for resistance to the allelopathic effects of invasive plants , 2005 .

[102]  E. Aschehoug,et al.  Invasive plants versus their new and old neighbors: a mechanism for exotic invasion. , 2000, Science.

[103]  R. Callaway,et al.  Root Communication Mechanisms and Intracommunity Distributions of Two Mojave Desert Shrubs , 1992 .

[104]  M. Caldwell,et al.  Competing root systems: morphology and models of absorption , 1986 .

[105]  Norma L. Fowler,et al.  The Role of Competition in Plant Communities in Arid and Semiarid Regions , 1986 .

[106]  R. Callaway,et al.  Compensatory growth and competitive ability of an invasive weed are enhanced by soil fungi and native neighbours , 2001 .

[107]  H. Schenk,et al.  Modeling the effects of temperature on growth and persistence of tree species: A critical review of tree population models , 1996 .

[108]  J. Cahill,et al.  Canopy gaps are sites of reduced belowground plant competition in a productive old field , 2004, Plant Ecology.

[109]  Katherine E. Mills,et al.  MAINTENANCE OF DIVERSITY WITHIN PLANT COMMUNITIES: SOIL PATHOGENS AS AGENTS OF NEGATIVE FEEDBACK , 1998 .

[110]  R. Callaway,et al.  SOIL FUNGI AND THE EFFECTS OF AN INVASIVE FORB ON GRASSES: NEIGHBOR IDENTITY MATTERS , 2003 .

[111]  D. Aspinall AN ANALYSIS OF COMPETITION BETWEEN BARLEY AND WHITE PERSICARIA , 1960 .

[112]  H. Schenk Clonal splitting in desert shrubs , 1999, Plant Ecology.

[113]  R. Callaway,et al.  Root communication among desert shrubs. , 1991, Proceedings of the National Academy of Sciences of the United States of America.

[114]  J. Bever Feeback between Plants and Their Soil Communities in an Old Field Community , 1994 .

[115]  B. Casper,et al.  Investigating the relationship between neighbor root biomass and belowground competition: field evidence for symmetric competition belowground , 2000 .

[116]  A. Packer,et al.  Soil pathogens and spatial patterns of seedling mortality in a temperate tree , 2000, Nature.

[117]  Yuan Zhi-you,et al.  Effects of Plant Sizes on the Nitrogen Use Strategy in an Annual Herb, Helianthus annuus (Sunflower) , 2004 .

[118]  H. Hauggaard‐Nielsen,et al.  Facilitative root interactions in intercrops , 2005 .

[119]  P. Keddy,et al.  Above‐ and Belowground Competition Intensity in Two Contrasting Wetland Plant Communities , 1996 .

[120]  K. J. Day,et al.  The effects of spatially heterogeneous nutrient supply on yield, intensity of competition and root placement patterns in Briza media and Festuca ovina , 2003 .

[121]  E. V. von Wettberg,et al.  Larger Triticum aestivum plants do not preempt nutrient-rich patches in a glasshouse experiment , 2003, Plant Ecology.

[122]  J. Vivanco,et al.  How plants communicate using the underground information superhighway. , 2004, Trends in plant science.

[123]  M. Caldwell,et al.  Competition for Phosphorus: Differential Uptake from Dual-Isotope—Labeled Soil Interspaces Between Shrub and Grass , 1985, Science.

[124]  O. Falik,et al.  Root navigation by self inhibition , 2005 .

[125]  D. Hartnett,et al.  MYCORRHIZAE INFLUENCE PLANT COMMUNITY STRUCTURE AND DIVERSITY IN TALLGRASS PRAIRIE , 1999 .

[126]  M. Zobel,et al.  Studying plant competition: from root biomass to general aims , 2002 .

[127]  J. Bunce,et al.  Competitive Exploitation of Soil Water by Five Eastern North American Tree Species , 1977, Botanical Gazette.

[128]  M. Loreau,et al.  Biodiversity and ecosystem functioning : synthesis and perspectives , 2002 .

[129]  Alex C Rodriguez,et al.  Soil biota and exotic plant invasion , 2004, Nature.

[130]  G. Berntson,et al.  CHARACTERIZING THE SIZE DEPENDENCE OF RESOURCE ACQUISITION WITHIN CROWDED PLANT POPULATIONS , 2000 .

[131]  Jacob Weiner,et al.  Size Hierarchies in Experimental Populations of Annual Plants , 1985 .

[132]  T. Rajaniemi Evidence for size asymmetry of belowground competition , 2003 .

[133]  J. Cahill Fertilization effects on interactions between above-and belowground competition in an old field , 1999 .

[134]  K. Kitajima,et al.  MYCORRHIZAE DIFFERENTIALLY ALTER GROWTH, PHYSIOLOGY, AND COMPETITIVE ABILITY OF AN INVASIVE SHRUB , 2003 .

[135]  Uta Berger,et al.  Asymmetric competition as a natural outcome of neighbour interactions among plants: results from the field-of-neighbourhood modelling approach , 2004, Plant Ecology.

[136]  L. Aarssen,et al.  The separation of above- and below-ground competition in plants A review and critique of methodology , 2001, Plant Ecology.

[137]  R. Callaway,et al.  MYCORRHIZAE INDIRECTLY ENHANCE COMPETITIVE EFFECTS OF AN INVASIVE FORB ON A NATIVE BUNCHGRASS , 1999 .

[138]  Jonathan Silvertown,et al.  Plant coexistence and the niche , 2004 .

[139]  O. Leyser,et al.  Root system architecture determines fitness in an Arabidopsis mutant in competition for immobile phosphate ions but not for nitrate ions , 2002, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[140]  D. Lambert COMPETITION BETWEEN PLANTS OF COCKSFOOT (DACTYLIS GLOMERATA) GROWN FOR SEED , 1968 .

[141]  James D. Bever,et al.  GRASSROOTS ECOLOGY: PLANT-MICROBE-SOIL INTERACTIONS AS DRIVERS OF PLANT COMMUNITY STRUCTURE AND DYNAMICS , 2003 .

[142]  D. Coomes,et al.  IMPACTS OF ROOT COMPETITION IN FORESTS AND WOODLANDS: A THEORETICAL FRAMEWORK AND REVIEW OF EXPERIMENTS , 2000 .

[143]  J. Cahill What evidence is necessary in studies which separate root and shoot competition along productivity gradients? , 2002 .

[144]  R. Ford Denison,et al.  Darwinian Agriculture: When Can Humans Find Solutions Beyond The Reach of Natural Selection? , 2003, The Quarterly Review of Biology.

[145]  C. H. Muller Allelopathy as a factor in ecological process , 2004, Vegetatio.

[146]  L. R. Oliver,et al.  Competitive Mechanisms of Common Cocklebur (Xanthium strumarium) and Soybean (Glycine max) During Seedling Growth , 1990, Weed Science.

[147]  S. Hartley,et al.  Soil organic matter distribution and below-ground competition between Calluna vulgaris and Nardus stricta , 2002 .

[148]  Michel Loreau,et al.  Partitioning selection and complementarity in biodiversity experiments , 2001, Nature.

[149]  M. Zobel,et al.  Plant Coexistence in the Interactive Environment: Arbuscular Mycorrhiza Should Not Be out of Mind , 1997 .

[150]  T. Fahey,et al.  Efficiency of Nutrient Acquisition by Fine Roots and Mycorrhizae , 1995 .

[151]  C. Holzapfel,et al.  BIDIRECTIONAL FACILITATION AND INTERFERENCE BETWEEN SHRUBS AND ANNUALS IN THE MOJAVE DESERT , 1999 .

[152]  Jacob Weiner,et al.  How competition for light and nutrients affects size variability in Ipomoea tricolor populations , 1986 .

[153]  Ariel Novoplansky,et al.  Self/non‐self discrimination in roots , 2003 .