Topological scaling and plant root system architecture: developmental and functional hierarchies

SUMMARY Topology is an important component of the architecture of whole root systems. Unfortunately, most commonly applied indices used for characterizing topology are poorly correlated with one another and thus reflect different aspects of topology. In order to understand better how different methods of characterizing topology vary, this paper presents an exploration of several different methods for assigning order within branched root systems on the basis of (a) developmental (centrifugal) vs. functional (centripetal) ordering sequences and (b) whether orders are assigned to individual links or groups of adjacent links (segments). For each ordering system, patterns of scaling in relation to various aspects of link and segment size are explored using regression analyses. Segment-based ordering systems resulted in better fits for simple scaling relationships with size, but these patterns varied between developmental vs. functional ordering as well as the different size metrics examined. The functional (centripetal), link-based ordering system showed complex, non-linear scaling in relation to numbers of links per order. Using a simple simulation model of root growth, it is demonstrated that this method of characterizing root topology in relation to root size might be a more powerful tool for characterizing root system architecture than in the use of simple, single-index characterizations of topology.

[1]  F. I. Woodward,et al.  The root system architecture and development of Senecio vulgaris in elevated CO_2 and drought , 1992 .

[2]  G Cumming,et al.  Quantitative morphometry of the branching structure of trees. , 1973, Journal of theoretical biology.

[3]  Alastair H. Fitter,et al.  Architectural analysis of plant root systems , 1992 .

[4]  Jeffrey W. Hughes,et al.  Fine Root Dynamics in a Northern Hardwood Forest Ecosystem, Hubbard Brook Experimental Forest, NH , 1994 .

[5]  W. A. Cannon,et al.  A Tentative Classification of Root Systems , 1949 .

[6]  Benoit B. Mandelbrot,et al.  Fractal Geometry of Nature , 1984 .

[7]  P. Burrough Fractal dimensions of landscapes and other environmental data , 1981, Nature.

[8]  A. N. Strahler Hypsometric (area-altitude) analysis of erosional topography. , 1952 .

[9]  R. Chappell,et al.  Fitting bent lines to data, with applications to allometry. , 1989, Journal of theoretical biology.

[10]  S. Caul,et al.  What limits nitrate uptake from soil , 1991 .

[11]  A. Fitter,et al.  Ecological Interactions in Soil. , 1986 .

[12]  Jacob Weiner,et al.  Competition and Allometry in Three Species of Annual Plants , 1992 .

[13]  A. R. Ennos,et al.  Comparative Functional Morphology of the Anchorage Systems of Annual Dicots , 1992 .

[14]  J. Crawford,et al.  A multiple scaled fractal tree , 1990 .

[15]  N. MacDonald,et al.  Trees and networks in biological models , 1983 .

[16]  K. Pregitzer,et al.  Patterns of fine root mortality in two sugar maple forests , 1993, Nature.

[17]  J. Huxley Problems of relative growth , 1932 .

[18]  J. Lawton,et al.  Fractal dimension of vegetation and the distribution of arthropod body lengths , 1985, Nature.

[19]  D. Aspinall,et al.  Quantitative Studies of Root Development II. Growth in the Early Stages of Development , 1967 .

[20]  Alastair H. Fitter,et al.  Morphometric analysis of root systems: application of the technique and influence of soil fertility on root system development in two herbaceous species , 1982 .

[21]  B. Gunnarsson Fractal dimension of plants and body size distribution in spiders , 1992 .

[22]  G. Whitney,et al.  The Bifurcation Ratio as an Indicator of Adaptive Strategy in Woody Plant Species , 1976 .

[23]  L. B. Leopold,et al.  Trees and streams: the efficiency of branching patterns. , 1971, Journal of theoretical biology.

[24]  G Sugihara,et al.  Applications of fractals in ecology. , 1990, Trends in ecology & evolution.

[25]  P. Bottomley,et al.  In situ nuclear magnetic resonance imaging of roots: influence of soil type, ferromagnetic particle content, and soil water , 1987 .

[26]  T. Ebert,et al.  Allometry and model II non-linear regression , 1994 .

[27]  W. Rice ANALYZING TABLES OF STATISTICAL TESTS , 1989, Evolution; international journal of organic evolution.

[28]  Gail W. T. Wilson,et al.  Architectural analysis of plant root systems 1. Architectural correlates of exploitation efficiency , 1991 .

[29]  A. Eshel,et al.  Plant roots : the hidden half , 1991 .

[30]  R. O'Neill,et al.  Landscape patterns in a disturbed environment , 1987 .

[31]  Kurt S. Pregitzer,et al.  THE DEMOGRAPHY OF FINE ROOTS IN A NORTHERN HARDWOOD FOREST , 1992 .

[32]  G. Berntson Modelling root architecture: are there tradeoffs between efficiency and potential of resource acquisition? , 1994 .

[33]  E. David Ford,et al.  Noninvasive studies of conifer roots: nuclear magnetic resonance (NMR) imaging of Douglas-fir seedlings , 1991 .

[34]  S. Gould ALLOMETRY AND SIZE IN ONTOGENY AND PHYLOGENY , 1966, Biological reviews of the Cambridge Philosophical Society.

[35]  E. Weibel Morphometry of the Human Lung , 1965, Springer Berlin Heidelberg.

[36]  Karl J. Niklas,et al.  Botanical Scaling. (Book Reviews: Plant Allometry. The Scaling of Form and Process.) , 1994 .

[37]  D. Aspinall,et al.  Quantitative Studies of Root Development I. The Influence of Nutrient Concentration , 1965 .

[38]  Alastair H. Fitter,et al.  The Topology and Geometry of Plant Root Systems: Influence of Watering Rate on Root System Topology in Trifolium pratense , 1986 .

[39]  D. Clarkson,et al.  The development and function of roots. , 1977 .

[40]  K. Horsfield Are diameter, length and branching ratios meaningful in the lung? , 1980, Journal of theoretical biology.

[41]  Alastair H. Fitter,et al.  Functional significance of root morphology and root system architecture , 1985 .

[42]  D. Park Does Horton's law of branch length apply to open branching systems? , 1985 .

[43]  Alastair H. Fitter,et al.  Architectural analysis of plant root systems 2. Influence of nutrient supply on architecture in contrasting plant species , 1991 .

[44]  D. Waller,et al.  Non-stationarity of tree branching patterns and bifurcation ratios , 1986, Proceedings of the Royal Society of London. Series B. Biological Sciences.

[45]  R. Horton EROSIONAL DEVELOPMENT OF STREAMS AND THEIR DRAINAGE BASINS; HYDROPHYSICAL APPROACH TO QUANTITATIVE MORPHOLOGY , 1945 .

[46]  W. Lyford Rhizography of non-woody roots of trees in the forest floor , 1975 .

[47]  J. S. Smart The analysis of drainage network composition , 1978 .

[48]  Andrei G. Jablokow,et al.  Carbon cost of root systems: an architectural approach , 1994 .

[49]  G. Berntson The Characterization of Topology: A Comparison of Four topological Indices for Rooted Binary Trees , 1995 .