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

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

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

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

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

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

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

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

This method. (7) for the mass–radius method. The test is based on the ratio of reduced 2 values. 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). from a statistical point of view such a method would not be justified. and indeed it should not be expected to. The linear region can also be calculated by determining the local slopes. be able to fit the data better. The region in which the local slopes are constant is then taken as the linear region (7). 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. A hierarchical cluster analysis yielded two regression lines with two different D values: 1.41 (open circles) and 1. The simplest method of obtaining D is to fit a regression line to all data points and determine the slope of this line. Thus fractal analysis . The D with the smallest linear range (1.07. use of the value with the longest linear range is suggested.41. The higher-degree polynomial will always. 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. Deciding on the range of linearity and especially if it is significance has been addressed by Russ (49). which will have an F distribution (49). The left-hand side shows the digitized image of a retinal bipolar cell. ADVANTAGES AND POTENTIAL PROBLEMS OF FRACTAL DIMENSIONS FIG.318 ´ NDEZ AND JELINEK FERNA cat retinal ganglion cells are not fractal due to their limited linearity. 52. described by Caserta et al. One of the main ones is that the sensitivity changes as the window over which the local slopes is obtained is decreased (62). If the linear fit is accepted then the image is fractal. however. however. is to calculate the n-point local slopes. 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.25). but it uses up one more degree of freedom in the process. Alternatively. The use of a hierarchical cluster analysis to compute particular subsets of the log–log values that achieve the best linear fittings (Fig. Several publications have used this method to determine the slope of the data points (10. Because of the limited scale invariance of neurons different authors have used different methods to determine D from log–log values. personal communication). as the difference in log N(r) divided by log (r) for every n successive points. 53. it is a statistically significant parameter for identifying and differentiating neuronal cell classes. open circles). The test is performed using a critical value of F ( p ⫽ 0. 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. has several flaws. When this method produces multiple values of D. This method considers the D of the cell drawing to be the one with the longest linear range (1. Clearly then. Therefore the linearity region increases as the window is increased and makes this is a very subjective method. 37. 9) has also been reported (12).07 (filled circles). 9. filled circles) could be attributed to finite size effects at very low scales. 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. as biological objects display statistical self-similarity only between a short range of dimensions. Panico and Sterling (61) also used the local slope method to determine the D of their images. One method for this. Method for the graphic determination of fractal dimensions. The range of linearity is not important if the D obtained in this way is used in differentiating between different cell types (Caserta. The right-hand side shows a plot of the box counting measurements. and the improvement in the fit may not be that great. of course. 61).

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

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