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

This addition of detail results in an ideal fractal object having an infinite boundary length (16. 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.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. Figure 1 shows an approximation of an ideal/ theoretical fractal with a fractal dimension of 1. FIG. Construction of the Koch curve with a D of 1. Thus natural patterns display statistical self-similarity only between an upper and lower bound. that is. Computer-generated fractals. It is called dimension because it provides a measure of how completely an object fills space. 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. it is equal to the standard Euclidean dimension for which an ideal point has a dimension of 0. FRACTAL DIMENSIONS An important parameter in fractal analysis of biological structures is the fractional or fractal dimension (D). We also review some of the methodologies available for calculating the fractal dimension. Limitations are also imposed by recording and imaging techniques. they do not have a characteristic unit of length. such as the Koch curve. and so on) the fine detail of the complex curve would be lost due to the resolution limits of the printing process. and a perfectly solid volume has a dimension of 3. Thus fractals are always described by power functions since homogeneous power laws lack natural scales. the advantages and problems of fractal geometry. E. The fractal dimension for this purpose is therefore not intended to indicate whether the image is a fractal object. Raising equilateral triangles from the middle third of each of the line segments in the object produces the image in (C). 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. and does not necessarily imply any biological process nor mechanism involved in their development. 17). When D takes an integer value. Then the middle third is raised to produce an equilateral triangle (B). time. Mandelbrot has shown that the boundary length of a fractal object can be mathematically expressed as a power law.26 that was described by the Swedish mathematician. Further it should be kept in mind that all natural objects are. Helge von Koch in 1904. Many patterns in biology display a limited self-similarity or approximate self-similarity. unavoidably finite and limited in scale by their own nature. in contrast with mathematical fractals. and some of its current applications in neuroscience. are sometimes termed prefractals since they are limited resolution images and therefore do not realize the detail implicit in the complete mathematical formulation (15).26. . D is called fractal because it usually is not an integer. At higher stages of construction (D. The sequential construction of this fractal begins with a straight line (A). an ideal line has a dimension of 1. 14) it should be noted that the fractal dimension is only a descriptive parameter. They are generally held to be statistically self-similar. or mass (16). which increases in value with increasing structural complexity and describes the “fractured” nature of objects in nature (10). 1. like the dendritic field area or the size of the soma. 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. an ideal plane has a dimension of 2.

For example. Hausdorff (32) suggested that the volume or measure of the sphere should be eD where e equals the resolution of measurement. drawn from throughout the retina. 2. 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). 3). It is calculated by covering an object with countable spheres whose radii are not greater than the image but decrease to zero. nerve cells seen in two dimensions are not straight lines. 19). It is not easy to give a precise definition of a fractal (15). with their associated box counting dimensions. the volume goes either to zero or infinity. 1. 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. Examples of different cat ganglion cells. a very good estimate of D can be achieved by different fractal analysis methods (Fig. neurons with low D values.45 (Fig. Since many of these fractal dimensions are used mainly in pure mathematics or applied physics. A straight line is drawn from the cell silhouette to its value on the D axis. Calculating the Hausdorff dimension is generally FIG. for instance.2. Table 1 lists some of the most important fractal dimensions with their synonyms and context. 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). All methods rely on the relationship between a measuring device and the object’s spatial distribution. Various other aspects of fractal analysis and D are discussed formally by other authors (15. would have relatively few dendritic branches and cover the two-dimensional area less completely than neurons with higher D values like 1. or other experimental data obtained from presentations of natural objects.USE OF FRACTAL THEORY IN NEUROSCIENCE Since. say 1. This definition of dimension was extended and put into a more systematic framework by Besicovitch (33). 2). 20–31). this would be when D ⫽ log 4/log 3 ⫽ 1.26. . The fractal dimensions are sufficiently different to suggest that they represent distinct ganglion cell types. 16. their D values fall between 1 and 2. METHODS FOR DETERMINING FRACTAL DIMENSIONS Although the mathematically rigorous determination of D is impossible for a fractal point set obtained 311 from digitized photographs. and they do not completely cover the two-dimensional area. The D-dimensional Hausdorff measure of an image is finite only when D (the dimension value) equals the dimension of the image. 18. Measuring any self-similar set with spheres of integer dimension. drawings. For the Koch curve shown in Fig. we consider only those that are potentially useful in neuroscience.

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

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

The following methods can be used for noncontiguous structures as well as for 2D and 3D images. 6B.html. as reported by the above articles. A common form of this algorithm. The method is illustrated in Fig. noncontiguous structures or closed loops within a structure). 3C. 7). The number N(r) of pixels constituting the image is counted and at the same time the scaling factor r of pixels is recorded. A Macintosh program for calculating D using this method can be found at the following URL: http://plantecohost. S. called the Minkowski “sausage. The length of the border for each respective diameter is determined by the area of the outline divided by the diameter. (10). The box counting method applies to any structure in the plane and can be adapted for structures in three-dimensional space (7. the result will be the length of the curve. For a fractal curve the length will continue to increase as the radius of the circles decreases. The sequence of box sizes for grids is usually reduced by a factor of 1/2 from one grid to the next. The NIH Image program and its many macros can be fetched in a number of ways. the points will fall on a straight line between an upper and lower bound with negative slope. (5. A circle is swept continuously along the line and the area that is covered. 47–49). In practice. The fractal dimension is then estimated from the slope of the log–log plot of length against diameter. 36.314 ´ 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).” is determined. This is done by application of a convolution procedure which is part of the image analysis program (dilation macro from NIH). the image is digitized by pixels having a given scaling factor r (Fig.harvard. with the initial box size being the size of the image. 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. 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.g. is counted (Fig. The pixel dilation method.nih. D is then calculated by fitting a linear regression to the following equation: log(r) ⫽ ⫺D* log(n) ⫹ K. where r ⫽ resolution of image (number of pixels per unit length)..info. 43. These are detailed at http://rsb. 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. Many research reports using this scheme to analyze neuron structures are found in the literature (10. This value is then plotted as a function of the circle diameter. Pixel Dilation Method The pixel dilation method is based on the Minkowski–Bouligand dimension. is that images composed of more than one simple perimeter cannot be processed accurately (e. (10) and others (11. 36).gov/pub/nih-image/user-macros/ box count macro. A method similar to the box counting technique is the grid intercept method (45. 46). 51–56). or gradient is related to the fractal dimension by D ⫽ 1 ⫺ S (5).gov/ . for use with NIH Image image processing software. 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. Euclidean curve. r is then made progressively smaller and the corresponding number of nonempty boxes. 6). Figure 5 shows an example of this calculation for a retinal ganglion cell. This filters out structures smaller than the current diameter of the circle. 37–42). The calliper method has previously been used to characterize neurons (10. and K Minkowski–Bouligand Dimension The Minkowski–Bouligand dimension is different from the Hausdorff dimension (18). 3B). and the slope (on the usual log–log plot) gives the dimension. Box Counting Method To estimate D.nih. The logarithm of N(r) versus the logarithm of r gives a line whose gradient corresponds to D. One major drawback of the calliper method. 12. has been implemented by Smith et al. 6A). 44). N(r). This is equivalent to the “grid” method described by Smith et al. 10. n ⫽ the number of pixels intersecting a portion of the image.txt. 29).edu/gmbWWW/ APPL. is a constant. The slope. replaces each pixel of the border by a circle whose diameter ranges from 3 to 61 pixels (Fig. For a smooth. 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. 6C).

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

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

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

07 (filled circles). however. of course. use of the value with the longest linear range is suggested. which will have an F distribution (49). The linear region can also be calculated by determining the local slopes.41 (open circles) and 1. be able to fit the data better. One method for this. The simplest method of obtaining D is to fit a regression line to all data points and determine the slope of this line. however. and the improvement in the fit may not be that great. The range of linearity is not important if the D obtained in this way is used in differentiating between different cell types (Caserta.07. The D with the smallest linear range (1. (7) for the mass–radius method.318 ´ NDEZ AND JELINEK FERNA cat retinal ganglion cells are not fractal due to their limited linearity. The use of a hierarchical cluster analysis to compute particular subsets of the log–log values that achieve the best linear fittings (Fig. 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. Thus fractal analysis . When this method produces multiple values of D. as the difference in log N(r) divided by log (r) for every n successive points. The higher-degree polynomial will always. This method. 9) has also been reported (12). personal communication). The left-hand side shows the digitized image of a retinal bipolar cell. The test is performed using a critical value of F ( p ⫽ 0. The region in which the local slopes are constant is then taken as the linear region (7).41. it is a statistically significant parameter for identifying and differentiating neuronal cell classes. 53.25). but it uses up one more degree of freedom in the process. 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. Deciding on the range of linearity and especially if it is significance has been addressed by Russ (49). 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). Panico and Sterling (61) also used the local slope method to determine the D of their images. 61). from a statistical point of view such a method would not be justified. is to calculate the n-point local slopes. ADVANTAGES AND POTENTIAL PROBLEMS OF FRACTAL DIMENSIONS FIG. 52. The right-hand side shows a plot of the box counting measurements. and indeed it should not be expected to. Because of the limited scale invariance of neurons different authors have used different methods to determine D from log–log values. A hierarchical cluster analysis yielded two regression lines with two different D values: 1. described by Caserta et al. 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. The test is based on the ratio of reduced ␹2 values. filled circles) could be attributed to finite size effects at very low scales. 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. 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. Several publications have used this method to determine the slope of the data points (10. Alternatively. One of the main ones is that the sensitivity changes as the window over which the local slopes is obtained is decreased (62). has several flaws. 37. Method for the graphic determination of fractal dimensions. Clearly then. 9. Therefore the linearity region increases as the window is increased and makes this is a very subjective method. If the linear fit is accepted then the image is fractal. as biological objects display statistical self-similarity only between a short range of dimensions. This method considers the D of the cell drawing to be the one with the longest linear range (1. open circles).

higher D values are obtained by using the mass fractal methods than by using the pixel dilation and box counting procedures. To capture all this richness of this complex structure into a theoretical model is one of the major challenges of modern theoretical biology (64). summarizes concisely and meaningfully the amount of detail. 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). 64). 319 Furthermore whether a higher fractal dimension would correlate with a more complex physiological response is still an unresolved issue (9. 62). Although all analysis methods rely on the relationship between a measuring device and the object’s spatial distribution. Many neurons display irregular shapes and discontinuous morphogenetic patterns in support and in connection with their functional diversity. This contrasts with integer-dimensional measurement of anisotropic objects which require multiple samples through the thickness of the threedimensional objects (1). 19). 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). space filling. the connection between empirical values of D and any specific growth mechanism should be avoided and require the answering of further experimental questions. such us dendritic field extent and total dendritic length. to more complicated global. Furthermore fractal geometry has some other advantages over its integer-dimensional counterparts. CONCLUSIONS AND FUTURE DEVELOPMENTS Fractal analysis has already found widespread application in the field of neuroscience and is being used in many other areas. These parameters range from simple metrical descriptors. 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). however. 39. 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). 12. not all methods give identical results for the same form. and various authors have discussed classification systems of neurons using fractal analysis (7. Finally. be kept in mind that D is only a descriptive parameter. Notwithstanding the above-mentioned limitations. Thus. 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. 12). 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. Thus many quantitative parameters have been used to characterize the morphology of nerve cells. Thus in almost all circumstances the fractional component of dimension is retained when a fractal object is projected to a lower-order dimension (18. the fractal dimension. and does not necessarily imply any underlying mechanism of form generation. the results are always consistent. it remains that in many situations a single number. an example being the projection of three-dimensional retinal ganglion cells onto a two-dimensional film or drawing (7). 63). 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. 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. In general. Unlike mathematically generated fractals. Thus determining D of a neuron. A basic consideration is that most measurements cover only a relatively short range of dimensions. like the dendritic field area or the number of segments of a dendritic tree. that can be used for an objective assessment of the degree of complexity (a concept heretofore not readily quantifiable) of developing and mature neurons. could immensely aid in the morphological discernment of different neuron types or neurons that . 42. biological data that have a linear fit of more than two orders of magnitude are extremely rare (66–69). For example. 62. It should. or complexity of neurons. 59. in addition to the other morphometric criteria typically used.USE OF FRACTAL THEORY IN NEUROSCIENCE has an important role in characterizing natural objects. real data cannot be ideally fractal over all scales. descriptors such as D.

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