METHODS 24, 309–321 (2001

)
doi:10.1006/meth.2001.1201, available online at http://www.idealibrary.com on

Use of Fractal Theory in Neuroscience:
Methods, Advantages, and Potential Problems
Eduardo Ferna´ndez* and Herbert F. Jelinek†,1
*Instituto de Bioingenierı´a, Universidad Miguel Herna´ndez, Elche, Spain; and
†School of Community Health, Charles Sturt University, Albury, Australia

Fractal analysis has already found widespread application
in the field of neuroscience and is being used in many other
areas. Applications are many and include ion channel kinetics
of biological membranes and classification of neurons according to their branching characteristics. In this article we
review some practical methods that are now available to allow
the determination of the complexity and scaling relationships
of anatomical and physiological patterns. The problems of describing fractal dimensions are discussed and the concept of
fractal dimensionality is introduced. Several related methodological considerations, such as preparation of the image and
estimation of the fractal dimensions from the data points,
as well as the advantages and problems of fractal geometric
analysis, are discussed. 䉷 2001 Academic Press

One of the basic tenets of neurobiology holds that
the function of a nerve cell is largely dependent on
its structure. To understand how a neuron integrates
its myriad synaptic inputs to generate an appropriate response, a thorough understanding of the
cell’s morphology and geometry is required. Thus
many quantitative parameters have been used to
characterize the morphology of nerve cells. The simple models that have been used so far for shapes,
such as spheres, ellipsoids, and polyhedra, and their
corresponding two-dimensional profiles, are useful
for many purposes, including estimates of volume or
size distribution, but certainly fall short in dealing
1
To whom all correspondence and reprint requests should be
addressed at School of Community Health, Charles Sturt University, Albury, 2640, N.S.W., Australia. Fax: ⫹61 2 516772. E-mail:
hjelinek@csu.edu.au.

1046-2023/01 $35.00
Copyright 䉷 2001 by Academic Press
All rights of reproduction in any form reserved.

with neuronal complexity. One method for describing
the irregular shape of feature profiles has been to
“unroll” that shape by plotting distance from the
centroid as a function of angle, and then to perform
a Fourier analysis on the resulting curve (1). Besides
being computationally demanding, this approach has
difficulty in dealing with shapes so irregular that
the radius line may intersect the outline more than
once. Other approaches such as the form factor are
also not useful since it can be the same for different
shapes and is not able for instance to measure quantitatively the complexity or degree of dendritic
branching (1). Sholl analysis has been widely used
to analyze branching characteristics of neurons. This
method determines the number of intersections of
dendrites with a set of concentric circles (2). Sholl
analysis is not an ideal method to measure neuronal
complexity as it has several problems with determining the number of processes at each level of measurement. Furthermore it is insensitive to the border
ruggedness of neurons (3, 4).
A new approach to this problem requires us to
think about the concepts of fractal geometry. In the
short period since Mandelbrot defined the term fractal (5), the field of fractal geometry has enjoyed an
enormous surge of popularity. The key observation
is that structures growing according to stochastic
processes are not really as disordered as they seem at
first glance. A nontrivial, scale invariant symmetry
over several orders of magnitude has been found to be
a typical principle of order for such growth processes,
which can be quantified by the fractal dimension.
Although this parameter can be used to quantify the
309

D is called fractal because it usually is not an integer. E. which increases in value with increasing structural complexity and describes the “fractured” nature of objects in nature (10). Helge von Koch in 1904. Figure 1 shows an approximation of an ideal/ theoretical fractal with a fractal dimension of 1. and some of its current applications in neuroscience. We also review some of the methodologies available for calculating the fractal dimension. an ideal plane has a dimension of 2. The fractal dimension for this purpose is therefore not intended to indicate whether the image is a fractal object. FRACTAL DIMENSIONS An important parameter in fractal analysis of biological structures is the fractional or fractal dimension (D). the advantages and problems of fractal geometry. The form of this object is complex since any change in magnification/scale will show more detail to the resolution limit as the magnification is increased. Raising equilateral triangles from the middle third of each of the line segments in the object produces the image in (C). Thus fractals are always described by power functions since homogeneous power laws lack natural scales. Then the middle third is raised to produce an equilateral triangle (B). such as the Koch curve. At higher stages of construction (D. Thus natural patterns display statistical self-similarity only between an upper and lower bound. time. THEORETICAL CONSIDERATIONS OF FRACTAL GEOMETRY AND NATURALLY OCCURRING FRACTALS An object is said to be fractal if certain criteria such as the object being self-similar or scale invariant are met. Construction of the Koch curve with a D of 1. 17). in contrast with mathematical fractals. that is. Many patterns in biology display a limited self-similarity or approximate self-similarity. In this paper we emphasize that fractal analysis is a useful tool for improving image description and for categorizing images representing morphologically complex objects based on the value of the fractal dimension. they do not have a characteristic unit of length. or mass (16). 1. and so on) the fine detail of the complex curve would be lost due to the resolution limits of the printing process. like the dendritic field area or the size of the soma. an ideal line has a dimension of 1. This addition of detail results in an ideal fractal object having an infinite boundary length (16. FIG. it is equal to the standard Euclidean dimension for which an ideal point has a dimension of 0.26 that was described by the Swedish mathematician. Limitations are also imposed by recording and imaging techniques. Further it should be kept in mind that all natural objects are. . Computer-generated fractals. and does not necessarily imply any biological process nor mechanism involved in their development. and a perfectly solid volume has a dimension of 3.310 ´ NDEZ AND JELINEK FERNA complexity of the borders of a neuron (6–12) and to measure how completely the branches of a neuron fill its dendritic field (13. unavoidably finite and limited in scale by their own nature. The final value of the amount of detail or irregularity at different scales associated with a natural object can then be determined by the use of fractal analysis. When D takes an integer value. It is called dimension because it provides a measure of how completely an object fills space.26. 14) it should be noted that the fractal dimension is only a descriptive parameter. They are generally held to be statistically self-similar. are sometimes termed prefractals since they are limited resolution images and therefore do not realize the detail implicit in the complete mathematical formulation (15). The sequential construction of this fractal begins with a straight line (A). Mandelbrot has shown that the boundary length of a fractal object can be mathematically expressed as a power law.

For the Koch curve shown in Fig.USE OF FRACTAL THEORY IN NEUROSCIENCE Since. Various other aspects of fractal analysis and D are discussed formally by other authors (15. say 1. with their associated box counting dimensions. Hausdorff (32) suggested that the volume or measure of the sphere should be eD where e equals the resolution of measurement. 20–31). Examples of different cat ganglion cells.26. It is not easy to give a precise definition of a fractal (15). The D-dimensional Hausdorff measure of an image is finite only when D (the dimension value) equals the dimension of the image. This definition of dimension was extended and put into a more systematic framework by Besicovitch (33). 16. It is calculated by covering an object with countable spheres whose radii are not greater than the image but decrease to zero. or other experimental data obtained from presentations of natural objects. The beta cell (on the right) has a more profuse branching pattern than the gamma cell (on the left) or the alpha cell (in the center). for instance. The fractal dimensions are sufficiently different to suggest that they represent distinct ganglion cell types.2. Table 1 lists some of the most important fractal dimensions with their synonyms and context. Measuring any self-similar set with spheres of integer dimension. 18. Since many of these fractal dimensions are used mainly in pure mathematics or applied physics. 3). their D values fall between 1 and 2.45 (Fig. nerve cells seen in two dimensions are not straight lines. A straight line is drawn from the cell silhouette to its value on the D axis. Calculating the Hausdorff dimension is generally FIG. a very good estimate of D can be achieved by different fractal analysis methods (Fig. the volume goes either to zero or infinity. and there are in the literature many different types of fractal dimensions so that even research mathematicians are not agreed on their names or equivalence (18. All methods rely on the relationship between a measuring device and the object’s spatial distribution. METHODS FOR DETERMINING FRACTAL DIMENSIONS Although the mathematically rigorous determination of D is impossible for a fractal point set obtained 311 from digitized photographs. neurons with low D values. 19). drawn from throughout the retina. For example. this would be when D ⫽ log 4/log 3 ⫽ 1. 2. and they do not completely cover the two-dimensional area. would have relatively few dendritic branches and cover the two-dimensional area less completely than neurons with higher D values like 1. Hausdorff Dimension The original intention of Hausdorff was to define a parameter that was independent of the resolution of measurement and was applicable to all shapes (16). we consider only those that are potentially useful in neuroscience. 2). 1. . drawings.

Basel. Lange (1998) in Fractals in Biology and Medicine (Nonnenmacher. The “capacity dimension” has become the fundamental definition of fractal dimension in the minds of many. (A) Calliper method. but it cannot be strictly applied to natural objects due to its finite range of fractal structure Easier to evaluate than DH. 1990 (6) Jelinek and Fernandez.). See text for more details. mosaic amalgamation dimension. A.. divider. To determine D.. box dimension. (C) Dilation method. Birhauser. divider dimension. 3. 1983 (5) Hausdorff. Note loss of border detail shown in (A) and (B) (D) Mass method example after application of six groups of concentric disks. with various diameters and centered on the border of the Koch island. and Weibel. The capacity dimension is related to the box counting and mass– radius methods that are its applied.(B) Box counting method. E. D(2) in multifractal analysis Context Generic term for fractal dimension Widely used in pure mathematics. two-dimensional embodiment and described below. compass dimension. a ruler of decreasing size r is used to measure the boundary or coastline of an image. 35). was introduced by Kolmogorov (34. 1989 (10) Mandelbrot.. usually is greater than or equal to the Hausdorff dimension Often used in calculating the fractal dimension of outlines Used for calculating the fractal dimensions of many biological structures in 2D and 3D Used in the context of clusters and networks. That is. F.. Some methods used for determination of fractal dimensions of a Koch triadic island with a D ⫽ 1. 1995 (7) Caserta et al. 1998 (59) very difficult and a more practical parameter of D. The length of the coastline then equals the size of the ruler times the number of steps r has taken to trace the coast. . and G. the capacity dimension. After dilation with a disk kernel diameter of 16 pixels. Losa.´ NDEZ AND JELINEK FERNA 312 TABLE 1 Some of the Most Widely Used Fractal Dimensions with their Synonyms and Contexts Dimension Symbol Synonyms Fractal Hausdorff D DH Minkowski– Bouligand DM Minkowsky sausage dimension.. An algorithm based on the Hausdorff dimension is the calliper dimension (also known as the compass. 1983 (5) Tatsumi et al. T. Figures 3A and 4 show examples of this method. Jr. Calliper Method FIG. 1983 (5) Takayasu.. This method is based on counting the number of steps that give a polygonal representation of an arbitrary object using different calliper spans.. The difference from the Hausdorff– Besicovitch dimension is that the set is now covered with spheres of identical radius (16). Smith. D(0) in multifractal analysis Mass DMR Hausdorff–Besicovitch dimension Mass fractal dimension.. All centers lie within the radius of gyration (large circle). or yardstick dimension). with permission. dilation dimension Calliper DC Box counting DB Richardson dimension. 1989 (45) Smith et al. 1989 (10) Takayasu. Eds. 1919 (32) Besicovitch. One finds that the boundary length is a function of the span of the calliper employed in the measurement. perimeter dimension Capacity dimension. 1991 (29) Richardson. 1935 (33) Mandelbrot. mass radius dimension.. G. 1990 (16) Smith et al. R.. 1983 (5) Smith et al. D. Reprinted. the length does not converge to a stable value but keeps increasing as the calliper span decreases. G. 1961 Mandelbrot. can also be applied to surfaces and biological structures Reference Mandelbrot. Kolmogorov dimension.. 1983 (5) Mandelbrot. 1992 (48) Caserta et al. from T. 1990 (16) Peitgen et al.50. 1989 (10) Schroeder.

(B) Graph of resulting log–log plot. Calliper method for ascertaing the boundary length of an image. . (A) Measuring the length of the coastline of the Australian continent. 4.USE OF FRACTAL THEORY IN NEUROSCIENCE 313 FIG.

The logarithm of N(r) versus the logarithm of r gives a line whose gradient corresponds to D. The fractal dimension is then estimated from the slope of the log–log plot of length against diameter. 3C. Note that the terms box counting method and grid intercept method refer generally to two different methods but are used interchangeably in the literature (16. 7).g. and the slope (on the usual log–log plot) gives the dimension. The NIH Image program and its many macros can be fetched in a number of ways. Euclidean curve. 6B. as reported by the above articles. where r ⫽ resolution of image (number of pixels per unit length). and K Minkowski–Bouligand Dimension The Minkowski–Bouligand dimension is different from the Hausdorff dimension (18). (10) and others (11.nih. The following methods can be used for noncontiguous structures as well as for 2D and 3D images. A circle is swept continuously along the line and the area that is covered. A common form of this algorithm. For a fractal curve the length will continue to increase as the radius of the circles decreases. Box Counting Method To estimate D. replaces each pixel of the border by a circle whose diameter ranges from 3 to 61 pixels (Fig. 36. The important difference between this and the calliper method is that the circle is moved so that its center lies on every point of the line..txt. n ⫽ the number of pixels intersecting a portion of the image.” is determined. (10). as devised by Flook (50). for use with NIH Image image processing software. has been implemented by Smith et al. The box counting method applies to any structure in the plane and can be adapted for structures in three-dimensional space (7.gov/pub/nih-image/user-macros/ box count macro. the Euclidean space containing the image is divided into a grid of boxes of size r.nih. This is equivalent to the “grid” method described by Smith et al.html. with the initial box size being the size of the image. 51–56). the points will fall on a straight line between an upper and lower bound with negative slope.gov/ . S. For a smooth. is a constant. The sequence of box sizes for grids is usually reduced by a factor of 1/2 from one grid to the next. called the Minkowski “sausage. 6A). the result will be the length of the curve. Measurement of N(r) at larger scaling factor (lower resolutions) is usually done by zooming down the image using the memory frame with four adjacent pixels making one pixel (Figs. The calliper method has previously been used to characterize neurons (10. The length of the border for each respective diameter is determined by the area of the outline divided by the diameter. A Macintosh program for calculating D using this method can be found at the following URL: http://plantecohost. This filters out structures smaller than the current diameter of the circle. The method is illustrated in Fig.harvard. This is done by application of a convolution procedure which is part of the image analysis program (dilation macro from NIH).314 ´ NDEZ AND JELINEK FERNA If the length of the boundary (coastline) versus the calliper length is plotted on a log–log scale. These are detailed at http://rsb. 47–49). 46). or gradient is related to the fractal dimension by D ⫽ 1 ⫺ S (5). D is then calculated by fitting a linear regression to the following equation: log(r) ⫽ ⫺D* log(n) ⫹ K. Pixel Dilation Method The pixel dilation method is based on the Minkowski–Bouligand dimension. (5. 44). The slope.edu/gmbWWW/ APPL. The pixel dilation method. The grid intercept method relies on progressively coarsening the image representation (by pixels having different scaling factors) and counting the number of pixels intersecting a portion of the image (Fig. 37–42). This value is then plotted as a function of the circle diameter. In practice. the image is digitized by pixels having a given scaling factor r (Fig. A macro for this method can be obtained from the National Institutes of Health (NIH) at ftp://codon. 6). N(r). One major drawback of the calliper method. 29). r is then made progressively smaller and the corresponding number of nonempty boxes. noncontiguous structures or closed loops within a structure). Figure 5 shows an example of this calculation for a retinal ganglion cell. The number N(r) of pixels constituting the image is counted and at the same time the scaling factor r of pixels is recorded. 43. is counted (Fig. A method similar to the box counting technique is the grid intercept method (45. 36). is that images composed of more than one simple perimeter cannot be processed accurately (e. Many research reports using this scheme to analyze neuron structures are found in the literature (10. 12.info. 10. 3B). 6C).

7. FIG. (B) Scaling factor ⫽ 2 ␮m. Box counting method applied to a retinal ganglion cell (cat beta cell) demonstrates the halving of the box sizes of the grids that overlay the image. Each image frame has pixels with a different scaling factor.USE OF FRACTAL THEORY IN NEUROSCIENCE 315 FIG. (B) Minkowski “sausage” created by application of the dilation method to (A). . FIG. Digitized images of a turtle ganglion cell. The perimeter is calculated as the area of this figure divided by the diameter of the dilating disk. 5. 6. (A) Binary image of a turtle ganglion cell. (A) Scaling factor ⫽ 1 ␮m. (C) Scaling factor ⫽ 4 ␮m.

that increases with the increase in the radius r (see Fig. 59). adsorption sites on a surface. The choice of format is related to the spacefilling attributes of the image and the attributes of the image one deems to be important. the cell body interior and that of the dendrites do fill a plane completely and hence have a D of 2. named Fractop. However.au/ fractop/ and discussed by Jones and Jelinek in this issue. the entire border falls outside. the filled interior is solid. steps of a random walk. One is related to how image presentation may influence the possible scaling relationship of the image and the associated estimated D. border roughness (Fig.edu. on the other hand. with a D of 2 (60). there are two further considerations. all appropriate fractal analysis methods approach the same limit. A multiplatform version for computing the mass fractal dimension is available from http://life. had significantly lower D values (58) since they represented only the dendritic branching and do not reflect the other characteristic of complexity. Therefore when calculating the D using complete binary images of neurons there may be a space-filling effect that can lead to a higher D or a D of 2. For strictly self-similar mathematical fractals such as the Koch curve. Usually the quantity of interest is the area of the image.txt) and other user contributed macros are found at ftp://codon. of the radius of gyration. However. This can happen with a known fractal such as a Koch snowflake. When analyzing neurons. 3D). Note that it is necessary to sample all local origins to sample as many data points belonging to the image as possible. . The histological techniques used may also lead to incomplete staining of the peripheral parts of the cell. For instance. binary images with cell body and axon removed. or border-only images of cat retinal ganglion cells as long as the dendrites are thin with respect to the cell body.nih. With neurons specifically. skeletonized or border-only images. has the added advantage of providing a choice for the number of centers and the fraction of the radius of gyration required.gov/pub/ nih-image/user-macros/. There are various versions for various Macintosh computers. This premise stems from the fact that the computer screen has a limited resolution and may not be able to represent branching patterns below the size of one pixel. In addition. the Hausdorff dimension (29. This particular macro (fractal dilation. 57). Mandelbrot (5) stated that an object that fills a plane completely has a dimension value of 2. If one takes a fraction of the radius of gyration. The radius of gyration is introduced as a method of avoiding the outer edges of the figure based on the premise that the peripheral parts of the image that represent natural objects such as neurons is incomplete. can be used and every point within this limit is then chosen as a local origin and the cluster mass (number of pixels occupied) within a distance r of this local origin calculated. the cell body and/or the axon may also be removed from binary or outline representations of neurons.csu.316 ´ NDEZ AND JELINEK FERNA nih-image/download. depending on the relationship between the internal area and the contour. monomers in a polymer chain. Skeletonized images. To implement this method for the analysis of 2D images. this finding is dependent on the type of cell one analyzes and does not hold for glia cells (60). etc. primary particles of a colloidal aggregate. The double logarithmic plot of M(r) against r gives a quantitative value for D. This version. This is due to the area of the cell body and dendrites being much smaller than the extent of the border (58. The sites may be pixels obtained from box counting.html. All possible choices of local origin are averaged and the average cluster mass M(r) is obtained. say 0. Mass–Radius Method The mass–radius dimension is defined by the relationship between the sites of an image found within a sphere or circle of a certain radius covering the image. when most of the mass is concentrated in a convex outer border the method totally fails because the radius of gyration falls tightly within the border itself.6. Having decided which analysis method to use. a circle or sphere of radius r is laid over the image. Previous results (58) have demonstrated no significant difference between the estimated D of binary images. To lessen computation time a fraction. M. and the other is related to estimating D from the data points. The method first computes the center of gravity and then the radius of gyration. 8). IMAGE PREPARATION AND DETERMINATION OF D Digitized images can be presented as binary. one could claim that it is only the border that is fractal.

the number of data points being related to the number of measuring steps.USE OF FRACTAL THEORY IN NEUROSCIENCE GRAPHIC DETERMINATION OF THE FRACTAL DIMENSION How the actual D value is obtained from the log– log data points can lead to differences in the magnitude of D. was not scale-invariant under this transformation and method (58). . 7) using the mass–radius method. 61). some investigators have obtained linear plots using skeletonized images of neurons. Differences in the linearity of the log–log data points was observed between binary. (6. Montague and Friedlander (14). The actual data points generally do not lie on a straight line for more than one to two decades. 8. and skeletonized images (58). D is related to the slope of the line. The figures on the bottom are the associated graphs of their fractal dimensions. This dependency on the analysis method to produce linear log–log plots with skeletonized images may explain the conclusions of Panico and Sterling (61).2) led at times to a sigmoid log–log data point distribution. Box counting analysis of the same turtle ganglion cell. indicating the 317 same image. also obtained linear log–log plots with skeletonized images. as described above. However. using a different implementation of the box counting method (greater number of box sizes) and different image handling (rotation of image and using multiple centers). outline-only. using binary (A). outlined (B). These authors used two variants of the box counting method and the mass–radius method with skeletonized images and concluded that FIG. for instance. This limited self-similarity or scale invariance is characteristic of biological material and is a focus of some controversy (51. Analysis of skeletonized images using the original NIH Image box counting method (Version 1. and skeletonized (C) images. Caserta et al. obtained linear log–log plots (⬎ 2 generations) with skeletonized images of retinal neurons. In such a plot. when skeletonized.

If the linear fit is accepted then the image is fractal. 9) has also been reported (12). and indeed it should not be expected to. The linear region can also be calculated by determining the local slopes. Their conclusion was that the region of true linearity of the local slopes was less than one generation and therefore the images analyzed were not self-similar and could not be fractal. Clearly then. Alternatively. which will have an F distribution (49).25).318 ´ NDEZ AND JELINEK FERNA cat retinal ganglion cells are not fractal due to their limited linearity. and the improvement in the fit may not be that great. This method considers the D of the cell drawing to be the one with the longest linear range (1. The region in which the local slopes are constant is then taken as the linear region (7). from a statistical point of view such a method would not be justified. be able to fit the data better. Method for the graphic determination of fractal dimensions. The test is performed using a critical value of F ( p ⫽ 0. described by Caserta et al. Thus fractal analysis . of course. When this method produces multiple values of D. Because of the limited scale invariance of neurons different authors have used different methods to determine D from log–log values. has several flaws. other methods included only points that fell on the straight part of the line and excluded data points obtained from the peripheral parts of the image (41). The simplest method of obtaining D is to fit a regression line to all data points and determine the slope of this line. Deciding on the range of linearity and especially if it is significance has been addressed by Russ (49). The D with the smallest linear range (1. 52. The higher-degree polynomial will always. personal communication). however. as the difference in log N(r) divided by log (r) for every n successive points. The test is based on the ratio of reduced ␹2 values.07 (filled circles). use of the value with the longest linear range is suggested. This technique allows the detection of changes in D at different scales of measurement and compensates for the finite size effects induced by the limited resolution of the images. (7) for the mass–radius method.41 (open circles) and 1. open circles). The use of a hierarchical cluster analysis to compute particular subsets of the log–log values that achieve the best linear fittings (Fig. One of the main ones is that the sensitivity changes as the window over which the local slopes is obtained is decreased (62). the questions of whether an image is fractal and whether an image belongs to a certain group based on the D value are different and need to be disentangled. but it uses up one more degree of freedom in the process. Therefore the linearity region increases as the window is increased and makes this is a very subjective method. 53. The right-hand side shows a plot of the box counting measurements. The left-hand side shows the digitized image of a retinal bipolar cell. ADVANTAGES AND POTENTIAL PROBLEMS OF FRACTAL DIMENSIONS FIG. The range of linearity is not important if the D obtained in this way is used in differentiating between different cell types (Caserta. filled circles) could be attributed to finite size effects at very low scales. it is a statistically significant parameter for identifying and differentiating neuronal cell classes. In a recent study (12) we posed the following question: Can the estimate of D resolve differences in neuronal branching when simpler metrical analysis alone cannot? Our results indicated that although D alone does not completely specify a cell’s morphology. Several publications have used this method to determine the slope of the data points (10.41.07. 9. Panico and Sterling (61) also used the local slope method to determine the D of their images. is to calculate the n-point local slopes. 37. as biological objects display statistical self-similarity only between a short range of dimensions. This method. 61). One method for this. however. who suggested that comparing the fit of the data points to a straight line and to a higher-degree polynomial can clarify whether a straight-line fit is an appropriate model of the data. A hierarchical cluster analysis yielded two regression lines with two different D values: 1.

Unlike mathematically generated fractals. and does not necessarily imply any underlying mechanism of form generation. 12). Thus many quantitative parameters have been used to characterize the morphology of nerve cells. it remains that in many situations a single number. Finally. 42. 319 Furthermore whether a higher fractal dimension would correlate with a more complex physiological response is still an unresolved issue (9. For example. to more complicated global. however. 39. biological data that have a linear fit of more than two orders of magnitude are extremely rare (66–69). 19). Thus. It has thus become important to establish some criteria for choosing a particular method and how these methods compare in order to standardize the computation of D (59). Thus in almost all circumstances the fractional component of dimension is retained when a fractal object is projected to a lower-order dimension (18. A further advantage of fractal analysis is that shrinkage or expansion of a specimen will not affect D as long as the artifact acts equally in all directions and the measured points still lie on the linear segment of the graph (19). These data reinforce the idea that comparison of measurements of different profiles using the same measurement method may be useful and valid even if the exact numeric value of the dimension is not necessarily very accurate. the connection between empirical values of D and any specific growth mechanism should be avoided and require the answering of further experimental questions. 62. CONCLUSIONS AND FUTURE DEVELOPMENTS Fractal analysis has already found widespread application in the field of neuroscience and is being used in many other areas. 64). an example being the projection of three-dimensional retinal ganglion cells onto a two-dimensional film or drawing (7). 62).USE OF FRACTAL THEORY IN NEUROSCIENCE has an important role in characterizing natural objects. space filling. Furthermore whether scale invariance is observed for a particular image is dependent on image presentation and the analysis program applied to obtain the final D (59. Not even the “coastline of Britain” example in Mandelbrot’s seminal work (5) has a power law behavior spanning more that one or two orders of magnitude (69). could immensely aid in the morphological discernment of different neuron types or neurons that . Notwithstanding the above-mentioned limitations. Our results using different methods to compute the D values show that although different measurement procedures and even the same algorithm performed by different computer programs and/or experimenters may give slightly different numerical values of D. A basic consideration is that most measurements cover only a relatively short range of dimensions. Many neurons display irregular shapes and discontinuous morphogenetic patterns in support and in connection with their functional diversity. Although all analysis methods rely on the relationship between a measuring device and the object’s spatial distribution. These parameters range from simple metrical descriptors. or complexity of neurons. the fractal dimension. Furthermore fractal geometry has some other advantages over its integer-dimensional counterparts. 63). In general. This means that D values of specimens that have been processed in different batches or at different laboratories can usually be compared directly (as long as the same methodology to calculate fractal dimension is used). summarizes concisely and meaningfully the amount of detail. real data cannot be ideally fractal over all scales. 59. not all methods give identical results for the same form. descriptors such as D. To capture all this richness of this complex structure into a theoretical model is one of the major challenges of modern theoretical biology (64). that can be used for an objective assessment of the degree of complexity (a concept heretofore not readily quantifiable) of developing and mature neurons. 12. Thus determining D of a neuron. in addition to the other morphometric criteria typically used. such us dendritic field extent and total dendritic length. be kept in mind that D is only a descriptive parameter. like the dendritic field area or the number of segments of a dendritic tree. This contrasts with integer-dimensional measurement of anisotropic objects which require multiple samples through the thickness of the threedimensional objects (1). It should. and various authors have discussed classification systems of neurons using fractal analysis (7. the results are always consistent. higher D values are obtained by using the mass fractal methods than by using the pixel dilation and box counting procedures. some of the images analyzed using fractal analysis may not demonstrate self-similarity or scale invariance over more than one or two levels of magnification and may not be fractal (61.

. (1958) Dokl. and Smith. B. Hausman. (1990) Fractal Physiology and Chaos in Medicine. 24.. 173–186. B. W. (1989) Proc. W. H.. 136. H. 159–166.. 15. Takeda. C. (1989) Fractal Growth Phenomena. 95–98. Akad. It gives an index of the space-filling capacity of an object but does not give information about the pattern of space filling. J. Wingate. G. Plenum. T. Tauci. F. 87. New York. Eldred. 101–113. Plenum. T.320 ´ NDEZ AND JELINEK FERNA show morphological alterations due to development. 28. Ed. T. 9. 39. Zhang. Plenum. 397–408. 18. A.). (1990) Ann. Behar. Basel. M. and Weibel. Ann. R. Paris. N. H. 33.. In addition we should remember that estimating D is only one of the tools of fractal geometry.. and Khokha.. A 42. R. Smith. E. 22. Daccord. (1990) Fractals. Neurol. Jr. T. Ammermu¨ller. T. A. New York. We thank Peter Bowdren and Cherryl Kolbe for technical assistance. Y. D. (1919) Math.. Sholl.. Cross. D. This means that any single number cannot be characteristic of the mixture (48).. J.. D. T. R. M. A. Caserta. D. E. Cambridge Univ. (1959) Dokl. T. G. R. 25. 16. (1989) Neurosci. World Scientific.. Boca Raton.. 5. B. 3.. Weinheim. Stanford. (1991) Neuroscience 41. 449–474. W.. Montague. SpringerVerlag. (1986) The Beauty of Fractals. E. pp. Freeman. Comp. Ishikawa. and Lange. 1440–1457. Morigiwa. Nauk SSSR 119. B.. Neurosci. G. Acad. Fernandez. Res. (1994) Micron 25. A. E. and Kolb. (1989) A Random Walk through Fractal Dimensions. Stanley. and Spence. Manchester Univ. T. Montague. Eds. D. Mandelbrot. 31. I. Fernandez.. J.. A. 27. Kolmogorov.. A. Eldred.). A. Block. E. USA 86. Basel. F. J. I. pp. J. H. Kaye. Birkauser. Fitzgibbon. Aust. T. M. A. Jr. G. Jr.. 387–406. Singapore. Jr. A criticism that could be leveled at almost all the implementations of measuring D is that it is not always an adequate descriptor of a determined profile. S131–S140. J. and Weibel. Federer. A. Ohtomo. P. Fractal and Dynamics. Fischer.. Sapoval. Dordrecht. T. (1990) Fractals in the Physical Sciences.. 7223–7227. 32. Springer-Verlag. SpringerVerlag. 38. Takayasu. Neurosci. 7. Sci. New York. 26. Vis. CRC Press. Neurosci.. V. . D. Res. (1986) On Growth and Form: Fractal and Non-fractal Patterns in Physics. 2. 115. S. T. M. and Ostrovsky. R.. (1990) Parameter.. 20. Russ. (1985) Chaos. Other measurements such as the lacunarity and the spectra of multifractal dimensions may contribute to these issues (65).. 29. Biomed. R. K. (1994) J... Fernandez. F. N. 35. H. W. Freeman. Natl. (Nonnenmacher. each one with a different value of D. and Eldred. G. P. A. (1994) Fractals in Science. 11.. W. Caserta. Further a structure can be a mixture of different fractals. 10. Ammermu¨ller. and Neale. N. (1969) Geometric Parameter Theory. 1869–1874. and Friedlander. FL. G. D. Eldred. and Fukuda. (1992) Neurosci. H. (1992) Invest. M... and Neale. New York. Bunde. Smith.. Rev. Cambridge.-O. J.F. and Friedlander. Comp. Berlin. E. Some investigators are starting to use multifractals as a more comprehensive methodology which gives information about the distribution of fractal dimensions in a structure. Losa. Eds. P. E. 8. Jr. Hausman. H. 3. H. von Bloh. R. Stanley... 121. Lange. New York. L. Lett.. H. A. Methods 27. J. E. New York. Chaos and Power Laws. G. 19. and Veerwer. H. von Bloh. G. Besicovitch... (1953) J. REFERENCES 1. (1985) The Geometry of Fractal Sets. New York. H. (Capowski. Soc. K. T. 13. 37. Berlin. G. A. W. D. Aditech. H. (1989) J. S. (1990) Phys. 940. Lange. Webb. 79.. H. F. G. A. Neurosci. 323... M.. Jr. (1995) J. Guiloff. 157–179. Rev... and Stanley.... N. New York. Losa. Hausman. (1935) Math. Press.. Y. D. E.. Eng. Kobayashi. Smith. E. Kimmel. Topology and Fractal Geometry. (1989) in Computer Techniques in Neuroanatomy. 19–31. Lett. 133–144. (1988) Fractals. Eldred.. E. Methods 56. R. Cox. F. E. M. Marcel Dekker. E.. (1994) Fractals in Biology and Medicine. 754–755. B. W. 5... C.. and Nittmann. J. Vicsek. 10. B. Suppl. Kolb. E. W. W. (1991) J.. R. Singapore. (1991) Fractals. Verlag Chemie. F. Ophthalmol.. 173–180. (1990) Phys. 14. 30. K. Neurol.. 34. Edgar. Press. D. Jelinek. D. P. 23. West. 135–149.. Behar. and Havlin. N. 33. 12. disease. 17.. 198–202. Sci. 861–864. J. H. Manchester. B.. S. Nauk SSSR 124. and Neale.). (1994) in Fractals in Biology and Medicine. World Scientific. Ann. J. Block. A. 64. I. Birkauser. T. G. Springer-Verlag. 21. Feder. West. 41. (1992) Neurosci. J. and Stanley.. Nonnenmacher. H. G.. and Richter. 296–329. 13. E.. (1983) The Fractal Geometry of Nature. Smith. Sheriff. van Pelt. Caserta. 210–220. J.. or experimental treatments. Suppl. (1986) Practical Stereology.. 347. 18. P.. W. and Matsuoka. (1994) in Fractal Geometry in Biological Systems: An Analytical Approach (Iannacconne. Schroeder. F. Smith for allowing us to use Fig. T.. Anat. Uylings. Schwarzer. S. G. 40.. R. 36. Nijhoff.. E. Falconer. D. ACKNOWLEDGMENTS We thank Dr. (1992) Proc. Bulderev. and Schellnhuber. Kolmogorov. Peitgen.. 6.. J. W. Akad. and Thompson. S. (1992) J. Sheriff. This work was supported by DGICYT Research Grant PB94/ 1509 to E. 3. 4. Hausdorff. H.

Porter. Embryol.. H. Springer-Verlag. Bowers. R. G. (1994) Brain Res. and Smith. A. and Weibel. (1994) J. Berlin. 48. Jelinek. Neurosci. 47. Brauer. K.. Jr.. 45. Birkhauser. Jelinek... K. 361. (1998) J. Losa. 295–298. 369–371. Kolb. Comp. Fernandez. Ghosh. Panico. D. Neurol. and Malcai. Neurol. (1994) Ann. Lange.. Ju¨rgens. 634. 402–406. D. Flook. 9–18. A. Springer-Verlag. 61. Methods 69. E. G. Amthor. F. Res. H. 60. New York.. Birkauser.. and Vahle-Hinz. . 783–784. G. Plenum. and Marks. Neurosci.. and Lange.. B. Kniffki. G.. Murray. L.. G.. 43. Eds. (Nonnenmacher. F. D.. Smith. 785–786. Bot. 52. 34... 21.. D. Senitz. 49. and Spence. H.USE OF FRACTAL THEORY IN NEUROSCIENCE 42. F.. 65. O. E. Methods 81.. G. (1996) J. 73. 57. Neurol. M... Pawlak. Losa.-O. Comp. Russ. M. F. W.. and Fernandez... Jelinek. Smith.. Jr. Jelinek. 54–66.. K. (1992) Fractals for the Classroom. 54. Eds. Bernston. Y. T. 171–178. Bot. Neurosci. and Eiston. 479–490. (1998) Science 279. 62. 343. 281–284. 50.. 68. Schouten. J. C. Brauer. T.. 784 69. G. G. Neurosci. D. E. O. A. (1994) Fractal Surfaces. G. Thesis. 64. in press.. 602. Jr. F. E.. Yamauchi. 130. 14. Tatsumi. 112–116. H. T. and Avnir. F.). T. O. T.. K.. Linberg. Landini. H. A. S.). Jr. (1995) J. Smith. Pfeifer. M.. 64. Hanke.. and Reichenbach.. Lidar. D. and Fisher. (1988) Soc. Neale. 46. (1996) The Use of Fractal Analysis in Cat Retinal Ganglion Cell Classification.. Jr. T. University of Sydney. (1994) in Fractals in Biology and Medicine (Nonnenmacher. S.. 59. P... 331. I. Lange.. Neurol. Lidar. D. H. Kniffki. 547–550. Ahnelt. E. G. R. P. (1993) CABIOS 9. C. Basel. (1997) Fractals. (1991) Neurosci. Jr. T. Reichenbach. (1988) Science 279. (1993) J.-O. 5. B. K. 370–386. and Saupe. (1989) Ann.. J. and Kono. B.. 56. T. (1988) The Science of Fractal Images.. Avnir. W.. Lett. K. Smith. (1978) Powder Technol.. 44. G.. G. (1993) J. Malcai. A. S. and Saupe..-D. D. (1998) Science 279. and Rippin. R. A. 183. (1991) Anat. R. F. 181–190. (1998) in Fractals in Biology and Medicine. Pawlak. A. (1997) Fractals. New York. C. (1993) Fractals 1. 321 55. J. G. 605–612. T. G. In Press. Basel. 51. Bilham. (1998) Science 279. A. N. O. (1995) J. J. D. Bilham. A. 361. M. 66.. P. and Behar. D.. H.-D. T. G. J. and Vahle-Hinz. 63. and Smith. Peitgen. G. Siegel. 123–136. 53. 673–684. J. 499–503. and Weibel.. Comp. Part One. 58. and Sterling. Mandelbrot. H. Peitgen. and Smith.. D. 39–40 67. Comp...

Sign up to vote on this title
UsefulNot useful