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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
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:
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
E. 17). an ideal line has a dimension of 1. are sometimes termed prefractals since they are limited resolution images and therefore do not realize the detail implicit in the complete mathematical formulation (15). Construction of the Koch curve with a D of 1. 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. This addition of detail results in an ideal fractal object having an infinite boundary length (16. they do not have a characteristic unit of length. Computer-generated fractals. 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. 1. Helge von Koch in 1904. . Thus natural patterns display statistical self-similarity only between an upper and lower bound. it is equal to the standard Euclidean dimension for which an ideal point has a dimension of 0. The sequential construction of this fractal begins with a straight line (A). Many patterns in biology display a limited self-similarity or approximate self-similarity. such as the Koch curve. an ideal plane has a dimension of 2. unavoidably finite and limited in scale by their own nature. 14) it should be noted that the fractal dimension is only a descriptive parameter. D is called fractal because it usually is not an integer.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. time. and some of its current applications in neuroscience. like the dendritic field area or the size of the soma. 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. Thus fractals are always described by power functions since homogeneous power laws lack natural scales. and does not necessarily imply any biological process nor mechanism involved in their development.26. Further it should be kept in mind that all natural objects are. and a perfectly solid volume has a dimension of 3. or mass (16). Limitations are also imposed by recording and imaging techniques. When D takes an integer value. and so on) the fine detail of the complex curve would be lost due to the resolution limits of the printing process. The fractal dimension for this purpose is therefore not intended to indicate whether the image is a fractal object. 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. in contrast with mathematical fractals. Raising equilateral triangles from the middle third of each of the line segments in the object produces the image in (C). Then the middle third is raised to produce an equilateral triangle (B). Figure 1 shows an approximation of an ideal/ theoretical fractal with a fractal dimension of 1. 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. FIG. It is called dimension because it provides a measure of how completely an object fills space. They are generally held to be statistically self-similar. that is.26 that was described by the Swedish mathematician. which increases in value with increasing structural complexity and describes the “fractured” nature of objects in nature (10). Mandelbrot has shown that the boundary length of a fractal object can be mathematically expressed as a power law. At higher stages of construction (D. We also review some of the methodologies available for calculating the fractal dimension.
1. 19). Calculating the Hausdorff dimension is generally FIG. neurons with low D values. nerve cells seen in two dimensions are not straight lines. It is calculated by covering an object with countable spheres whose radii are not greater than the image but decrease to zero. the volume goes either to zero or infinity. for instance. this would be when D ⫽ log 4/log 3 ⫽ 1.USE OF FRACTAL THEORY IN NEUROSCIENCE Since.45 (Fig. This definition of dimension was extended and put into a more systematic framework by Besicovitch (33). 20–31). or other experimental data obtained from presentations of natural objects. . Various other aspects of fractal analysis and D are discussed formally by other authors (15. 2. Hausdorff (32) suggested that the volume or measure of the sphere should be eD where e equals the resolution of measurement. a very good estimate of D can be achieved by different fractal analysis methods (Fig. For the Koch curve shown in Fig. drawn from throughout the retina. Table 1 lists some of the most important fractal dimensions with their synonyms and context. with their associated box counting dimensions. 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). their D values fall between 1 and 2. All methods rely on the relationship between a measuring device and the object’s spatial distribution. 16. Since many of these fractal dimensions are used mainly in pure mathematics or applied physics.26.2. drawings. say 1. would have relatively few dendritic branches and cover the two-dimensional area less completely than neurons with higher D values like 1. Measuring any self-similar set with spheres of integer dimension. A straight line is drawn from the cell silhouette to its value on the D axis. 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). 18. and they do not completely cover the two-dimensional area. 2). 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. 3). The D-dimensional Hausdorff measure of an image is finite only when D (the dimension value) equals the dimension of the image. For example. It is not easy to give a precise definition of a fractal (15). METHODS FOR DETERMINING FRACTAL DIMENSIONS Although the mathematically rigorous determination of D is impossible for a fractal point set obtained 311 from digitized photographs. we consider only those that are potentially useful in neuroscience. Examples of different cat ganglion cells. The fractal dimensions are sufficiently different to suggest that they represent distinct ganglion cell types.
This method is based on counting the number of steps that give a polygonal representation of an arbitrary object using different calliper spans. G. with permission. mass radius dimension. 1998 (59) very difficult and a more practical parameter of D. The difference from the Hausdorff– Besicovitch dimension is that the set is now covered with spheres of identical radius (16).´ 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. E. the length does not converge to a stable value but keeps increasing as the calliper span decreases. 1991 (29) Richardson. but it cannot be strictly applied to natural objects due to its finite range of fractal structure Easier to evaluate than DH. All centers lie within the radius of gyration (large circle). Smith. a ruler of decreasing size r is used to measure the boundary or coastline of an image.. with various diameters and centered on the border of the Koch island. R. divider dimension. 1989 (10) Takayasu. Birhauser.. Losa. and Weibel. Calliper Method FIG. That is. divider. . F. 1989 (10) Mandelbrot. 1961 Mandelbrot.. D(0) in multifractal analysis Mass DMR Hausdorff–Besicovitch dimension Mass fractal dimension. (C) Dilation method. The capacity dimension is related to the box counting and mass– radius methods that are its applied. compass dimension. 35). Note loss of border detail shown in (A) and (B) (D) Mass method example after application of six groups of concentric disks. D. An algorithm based on the Hausdorff dimension is the calliper dimension (also known as the compass. two-dimensional embodiment and described below. 1983 (5) Smith et al.. 1992 (48) Caserta et al. Reprinted. Figures 3A and 4 show examples of this method.. (A) Calliper method. 1983 (5) Tatsumi et al. or yardstick dimension). G.. and G. dilation dimension Calliper DC Box counting DB Richardson dimension. To determine D. 1919 (32) Besicovitch.. Jr. 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. See text for more details. perimeter dimension Capacity dimension.50.. Some methods used for determination of fractal dimensions of a Koch triadic island with a D ⫽ 1. One finds that the boundary length is a function of the span of the calliper employed in the measurement. the capacity dimension. 1990 (16) Peitgen et al. 1995 (7) Caserta et al. Eds. The “capacity dimension” has become the fundamental definition of fractal dimension in the minds of many..).. 1990 (16) Smith et al. was introduced by Kolmogorov (34. A. 3. 1983 (5) Hausdorff. Basel. 1983 (5) Takayasu.(B) Box counting method. After dilation with a disk kernel diameter of 16 pixels. 1989 (45) Smith et al. from T. D(2) in multifractal analysis Context Generic term for fractal dimension Widely used in pure mathematics. T. Kolmogorov dimension. 1935 (33) Mandelbrot. Lange (1998) in Fractals in Biology and Medicine (Nonnenmacher. The length of the coastline then equals the size of the ruler times the number of steps r has taken to trace the coast. 1990 (6) Jelinek and Fernandez. 1989 (10) Schroeder. 1983 (5) Mandelbrot. can also be applied to surfaces and biological structures Reference Mandelbrot. box dimension.. mosaic amalgamation dimension.
.USE OF FRACTAL THEORY IN NEUROSCIENCE 313 FIG. 4. (A) Measuring the length of the coastline of the Australian continent. (B) Graph of resulting log–log plot. Calliper method for ascertaing the boundary length of an image.
3B).nih. 6C). The logarithm of N(r) versus the logarithm of r gives a line whose gradient corresponds to D. the points will fall on a straight line between an upper and lower bound with negative slope. 7). 47–49). the result will be the length of the curve. (10). the image is digitized by pixels having a given scaling factor r (Fig. In practice. Box Counting Method To estimate D. 51–56). The calliper method has previously been used to characterize neurons (10. For a smooth. The box counting method applies to any structure in the plane and can be adapted for structures in three-dimensional space (7. 36). Many research reports using this scheme to analyze neuron structures are found in the literature (10. A circle is swept continuously along the line and the area that is covered.g.nih.” is determined. A method similar to the box counting technique is the grid intercept method (45. The slope. is counted (Fig. 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. 6). 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. The pixel dilation method. has been implemented by Smith et al. and K Minkowski–Bouligand Dimension The Minkowski–Bouligand dimension is different from the Hausdorff dimension (18). A macro for this method can be obtained from the National Institutes of Health (NIH) at ftp://codon. the Euclidean space containing the image is divided into a grid of boxes of size r. for use with NIH Image image processing software. 6B.edu/gmbWWW/ APPL. r is then made progressively smaller and the corresponding number of nonempty boxes. 12. The number N(r) of pixels constituting the image is counted and at the same time the scaling factor r of pixels is recorded. 6A).txt. n ⫽ the number of pixels intersecting a portion of the image.html. This filters out structures smaller than the current diameter of the circle.314 ´ NDEZ AND JELINEK FERNA If the length of the boundary (coastline) versus the calliper length is plotted on a log–log scale. as devised by Flook (50). Figure 5 shows an example of this calculation for a retinal ganglion cell.info. A common form of this algorithm.gov/ .harvard. The sequence of box sizes for grids is usually reduced by a factor of 1/2 from one grid to the next. 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. S. A Macintosh program for calculating D using this method can be found at the following URL: http://plantecohost. where r ⫽ resolution of image (number of pixels per unit length). and the slope (on the usual log–log plot) gives the dimension. (5. replaces each pixel of the border by a circle whose diameter ranges from 3 to 61 pixels (Fig. 44). 46). The fractal dimension is then estimated from the slope of the log–log plot of length against diameter. The NIH Image program and its many macros can be fetched in a number of ways. as reported by the above articles. Euclidean 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. This is equivalent to the “grid” method described by Smith et al. 37–42).gov/pub/nih-image/user-macros/ box count macro. 36. 43. or gradient is related to the fractal dimension by D ⫽ 1 ⫺ S (5). with the initial box size being the size of the image. is a constant. 10. The method is illustrated in Fig. Pixel Dilation Method The pixel dilation method is based on the Minkowski–Bouligand dimension. The length of the border for each respective diameter is determined by the area of the outline divided by the diameter. These are detailed at http://rsb. D is then calculated by fitting a linear regression to the following equation: log(r) ⫽ ⫺D* log(n) ⫹ K.. For a fractal curve the length will continue to increase as the radius of the circles decreases. is that images composed of more than one simple perimeter cannot be processed accurately (e. (10) and others (11. called the Minkowski “sausage. 3C. 29). The following methods can be used for noncontiguous structures as well as for 2D and 3D images. This value is then plotted as a function of the circle diameter. noncontiguous structures or closed loops within a structure). This is done by application of a convolution procedure which is part of the image analysis program (dilation macro from NIH). N(r). One major drawback of the calliper method.
FIG. 7. (B) Minkowski “sausage” created by application of the dilation method to (A). Each image frame has pixels with a different scaling factor. (A) Scaling factor ⫽ 1 m. (A) Binary image of a turtle ganglion cell. FIG. (B) Scaling factor ⫽ 2 m.USE OF FRACTAL THEORY IN NEUROSCIENCE 315 FIG. 5. (C) Scaling factor ⫽ 4 m. 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. 6. 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.
59). Usually the quantity of interest is the area of the image. This particular macro (fractal dilation. with a D of 2 (60). or border-only images of cat retinal ganglion cells as long as the dendrites are thin with respect to the cell body. that increases with the increase in the radius r (see Fig.edu. Having decided which analysis method to use. In addition. had significantly lower D values (58) since they represented only the dendritic branching and do not reflect the other characteristic of complexity. This can happen with a known fractal such as a Koch snowflake. Note that it is necessary to sample all local origins to sample as many data points belonging to the image as possible. the filled interior is solid. the cell body and/or the axon may also be removed from binary or outline representations of neurons. A multiplatform version for computing the mass fractal dimension is available from http://life. If one takes a fraction of the radius of gyration. on the other hand. 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. One is related to how image presentation may influence the possible scaling relationship of the image and the associated estimated D. of the radius of gyration.gov/pub/ nih-image/user-macros/. one could claim that it is only the border that is fractal.6. 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. However. steps of a random walk. All possible choices of local origin are averaged and the average cluster mass M(r) is obtained.csu.html. When analyzing neurons. 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 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. monomers in a polymer chain. The method first computes the center of gravity and then the radius of gyration. say 0.316 ´ NDEZ AND JELINEK FERNA nih-image/download. With neurons specifically. and the other is related to estimating D from the data points. This is due to the area of the cell body and dendrites being much smaller than the extent of the border (58. The choice of format is related to the spacefilling attributes of the image and the attributes of the image one deems to be important. depending on the relationship between the internal area and the contour. has the added advantage of providing a choice for the number of centers and the fraction of the radius of gyration required.au/ fractop/ and discussed by Jones and Jelinek in this issue. named Fractop. Mandelbrot (5) stated that an object that fills a plane completely has a dimension value of 2. adsorption sites on a surface. binary images with cell body and axon removed.txt) and other user contributed macros are found at ftp://codon. However. 8). primary particles of a colloidal aggregate. This version. skeletonized or border-only images. IMAGE PREPARATION AND DETERMINATION OF D Digitized images can be presented as binary. the entire border falls outside. The sites may be pixels obtained from box counting. For instance. a circle or sphere of radius r is laid over the image. the cell body interior and that of the dendrites do fill a plane completely and hence have a D of 2. 57). . Skeletonized images. To lessen computation time a fraction. The histological techniques used may also lead to incomplete staining of the peripheral parts of the cell. To implement this method for the analysis of 2D images. 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. border roughness (Fig. M. The double logarithmic plot of M(r) against r gives a quantitative value for D. all appropriate fractal analysis methods approach the same limit. this finding is dependent on the type of cell one analyzes and does not hold for glia cells (60).nih. etc. 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. there are two further considerations. There are various versions for various Macintosh computers. For strictly self-similar mathematical fractals such as the Koch curve. 3D). the Hausdorff dimension (29. Previous results (58) have demonstrated no significant difference between the estimated D of binary images.
when skeletonized. This dependency on the analysis method to produce linear log–log plots with skeletonized images may explain the conclusions of Panico and Sterling (61). obtained linear log–log plots (⬎ 2 generations) with skeletonized images of retinal neurons. and skeletonized (C) images. 7) using the mass–radius method. Caserta et al. and skeletonized images (58). as described above. These authors used two variants of the box counting method and the mass–radius method with skeletonized images and concluded that FIG. However. 61). The figures on the bottom are the associated graphs of their fractal dimensions. Analysis of skeletonized images using the original NIH Image box counting method (Version 1. for instance.2) led at times to a sigmoid log–log data point distribution. This limited self-similarity or scale invariance is characteristic of biological material and is a focus of some controversy (51. indicating the 317 same image. 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). was not scale-invariant under this transformation and method (58). also obtained linear log–log plots with skeletonized images. The actual data points generally do not lie on a straight line for more than one to two decades. Box counting analysis of the same turtle ganglion cell. Differences in the linearity of the log–log data points was observed between binary. Montague and Friedlander (14). . outlined (B). D is related to the slope of the line. (6. In such a plot. outline-only. 8. using binary (A). the number of data points being related to the number of measuring steps. some investigators have obtained linear plots using skeletonized images of neurons.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.
9) has also been reported (12). from a statistical point of view such a method would not be justified. and indeed it should not be expected to. The D with the smallest linear range (1. If the linear fit is accepted then the image is fractal. however. 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 higher-degree polynomial will always. Therefore the linearity region increases as the window is increased and makes this is a very subjective method. 9. Panico and Sterling (61) also used the local slope method to determine the D of their images. 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.25). Method for the graphic determination of fractal dimensions.07.07 (filled circles). Thus fractal analysis .318 ´ NDEZ AND JELINEK FERNA cat retinal ganglion cells are not fractal due to their limited linearity. has several flaws. (7) for the mass–radius method. it is a statistically significant parameter for identifying and differentiating neuronal cell classes. This method considers the D of the cell drawing to be the one with the longest linear range (1. as biological objects display statistical self-similarity only between a short range of dimensions. is to calculate the n-point local slopes. The right-hand side shows a plot of the box counting measurements. When this method produces multiple values of D. open circles). One method for this. 61). The left-hand side shows the digitized image of a retinal bipolar cell. and the improvement in the fit may not be that great. 53. be able to fit the data better. use of the value with the longest linear range is suggested. The test is performed using a critical value of F ( p ⫽ 0. A hierarchical cluster analysis yielded two regression lines with two different D values: 1. 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. One of the main ones is that the sensitivity changes as the window over which the local slopes is obtained is decreased (62). The use of a hierarchical cluster analysis to compute particular subsets of the log–log values that achieve the best linear fittings (Fig. however. 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. The test is based on the ratio of reduced 2 values. 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.41 (open circles) and 1. described by Caserta et al. personal communication). which will have an F distribution (49). The region in which the local slopes are constant is then taken as the linear region (7). Several publications have used this method to determine the slope of the data points (10.41. of course. The range of linearity is not important if the D obtained in this way is used in differentiating between different cell types (Caserta. but it uses up one more degree of freedom in the process. filled circles) could be attributed to finite size effects at very low scales. ADVANTAGES AND POTENTIAL PROBLEMS OF FRACTAL DIMENSIONS FIG. as the difference in log N(r) divided by log (r) for every n successive points. 52. 37. Alternatively. This method. Deciding on the range of linearity and especially if it is significance has been addressed by Russ (49). Because of the limited scale invariance of neurons different authors have used different methods to determine D from log–log values. The linear region can also be calculated by determining the local slopes. Clearly then. The simplest method of obtaining D is to fit a regression line to all data points and determine the slope of this line. 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.
be kept in mind that D is only a descriptive parameter. Thus in almost all circumstances the fractional component of dimension is retained when a fractal object is projected to a lower-order dimension (18.USE OF FRACTAL THEORY IN NEUROSCIENCE has an important role in characterizing natural objects. Thus many quantitative parameters have been used to characterize the morphology of nerve cells. or complexity of neurons. summarizes concisely and meaningfully the amount of detail. 12). and does not necessarily imply any underlying mechanism of form generation. could immensely aid in the morphological discernment of different neuron types or neurons that . 63). 39. 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. biological data that have a linear fit of more than two orders of magnitude are extremely rare (66–69). like the dendritic field area or the number of segments of a dendritic tree. it remains that in many situations a single number. Thus. 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). 19). 12. such us dendritic field extent and total dendritic length. 62). 62. 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). 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. To capture all this richness of this complex structure into a theoretical model is one of the major challenges of modern theoretical biology (64). For example. 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). and various authors have discussed classification systems of neurons using fractal analysis (7. Although all analysis methods rely on the relationship between a measuring device and the object’s spatial distribution. that can be used for an objective assessment of the degree of complexity (a concept heretofore not readily quantifiable) of developing and mature neurons. the fractal dimension. Furthermore fractal geometry has some other advantages over its integer-dimensional counterparts. Many neurons display irregular shapes and discontinuous morphogenetic patterns in support and in connection with their functional diversity. Finally. space filling. A basic consideration is that most measurements cover only a relatively short range of dimensions. 59. 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. to more complicated global. 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 all methods give identical results for the same form. Notwithstanding the above-mentioned limitations. an example being the projection of three-dimensional retinal ganglion cells onto a two-dimensional film or drawing (7). These parameters range from simple metrical descriptors. 42. 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). This contrasts with integer-dimensional measurement of anisotropic objects which require multiple samples through the thickness of the threedimensional objects (1). descriptors such as D. real data cannot be ideally fractal over all scales. the results are always consistent. 319 Furthermore whether a higher fractal dimension would correlate with a more complex physiological response is still an unresolved issue (9. in addition to the other morphometric criteria typically used. Unlike mathematically generated fractals. 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). however. Thus determining D of a neuron. higher D values are obtained by using the mass fractal methods than by using the pixel dilation and box counting procedures. In general. the connection between empirical values of D and any specific growth mechanism should be avoided and require the answering of further experimental questions. It should.
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