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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

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

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

Since many of these fractal dimensions are used mainly in pure mathematics or applied physics.2. This definition of dimension was extended and put into a more systematic framework by Besicovitch (33). would have relatively few dendritic branches and cover the two-dimensional area less completely than neurons with higher D values like 1. 1. neurons with low D values. drawings. For the Koch curve shown in Fig. a very good estimate of D can be achieved by different fractal analysis methods (Fig. A straight line is drawn from the cell silhouette to its value on the D axis. Measuring any self-similar set with spheres of integer dimension. 16. with their associated box counting dimensions. For example. 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. 20–31). nerve cells seen in two dimensions are not straight lines.45 (Fig. Examples of different cat ganglion cells. Table 1 lists some of the most important fractal dimensions with their synonyms and context. 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). . It is calculated by covering an object with countable spheres whose radii are not greater than the image but decrease to zero. and they do not completely cover the two-dimensional area. say 1. this would be when D ⫽ log 4/log 3 ⫽ 1.USE OF FRACTAL THEORY IN NEUROSCIENCE Since. 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. 19). we consider only those that are potentially useful in neuroscience. or other experimental data obtained from presentations of natural objects. drawn from throughout the retina.26. 2). The fractal dimensions are sufficiently different to suggest that they represent distinct ganglion cell types. All methods rely on the relationship between a measuring device and the object’s spatial distribution. Hausdorff (32) suggested that the volume or measure of the sphere should be eD where e equals the resolution of measurement. Various other aspects of fractal analysis and D are discussed formally by other authors (15. 18. It is not easy to give a precise definition of a fractal (15). 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). Calculating the Hausdorff dimension is generally FIG. The D-dimensional Hausdorff measure of an image is finite only when D (the dimension value) equals the dimension of the image. 3). for instance. their D values fall between 1 and 2. the volume goes either to zero or infinity.

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

Calliper method for ascertaing the boundary length of an image.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.

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

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

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

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

Deciding on the range of linearity and especially if it is significance has been addressed by Russ (49).07 (filled circles). Because of the limited scale invariance of neurons different authors have used different methods to determine D from log–log values. 53. 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 .07. is to calculate the n-point local slopes. The left-hand side shows the digitized image of a retinal bipolar cell. One of the main ones is that the sensitivity changes as the window over which the local slopes is obtained is decreased (62). 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. and the improvement in the fit may not be that great. If the linear fit is accepted then the image is fractal. The test is performed using a critical value of F ( p ⫽ 0. 61). The linear region can also be calculated by determining the local slopes. however. 9. The region in which the local slopes are constant is then taken as the linear region (7). The D with the smallest linear range (1. described by Caserta et al. 52. 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. however. The right-hand side shows a plot of the box counting measurements. Therefore the linearity region increases as the window is increased and makes this is a very subjective method. ADVANTAGES AND POTENTIAL PROBLEMS OF FRACTAL DIMENSIONS FIG. 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. Panico and Sterling (61) also used the local slope method to determine the D of their images. The higher-degree polynomial will always. 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. When this method produces multiple values of D. but it uses up one more degree of freedom in the process. Method for the graphic determination of fractal dimensions. it is a statistically significant parameter for identifying and differentiating neuronal cell classes. of course. from a statistical point of view such a method would not be justified. A hierarchical cluster analysis yielded two regression lines with two different D values: 1. Clearly then. and indeed it should not be expected to.25). One method for this. filled circles) could be attributed to finite size effects at very low scales. The test is based on the ratio of reduced 2 values. personal communication). as the difference in log N(r) divided by log (r) for every n successive points. This method. 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). be able to fit the data better.318 ´ NDEZ AND JELINEK FERNA cat retinal ganglion cells are not fractal due to their limited linearity. This method considers the D of the cell drawing to be the one with the longest linear range (1. Alternatively.41 (open circles) and 1. The range of linearity is not important if the D obtained in this way is used in differentiating between different cell types (Caserta. use of the value with the longest linear range is suggested. has several flaws. which will have an F distribution (49). open circles). 9) has also been reported (12). 37. The use of a hierarchical cluster analysis to compute particular subsets of the log–log values that achieve the best linear fittings (Fig. (7) for the mass–radius method. The simplest method of obtaining D is to fit a regression line to all data points and determine the slope of this line.41. as biological objects display statistical self-similarity only between a short range of dimensions.

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

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