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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

. Mandelbrot has shown that the boundary length of a fractal object can be mathematically expressed as a power law. 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. Further it should be kept in mind that all natural objects are. they do not have a characteristic unit of length. such as the Koch curve. 1. When D takes an integer value. that is. Figure 1 shows an approximation of an ideal/ theoretical fractal with a fractal dimension of 1. FIG. Raising equilateral triangles from the middle third of each of the line segments in the object produces the image in (C). Limitations are also imposed by recording and imaging techniques. and does not necessarily imply any biological process nor mechanism involved in their development. Then the middle third is raised to produce an equilateral triangle (B). or mass (16). 17). the advantages and problems of fractal geometry. 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 natural patterns display statistical self-similarity only between an upper and lower bound. This addition of detail results in an ideal fractal object having an infinite boundary length (16. an ideal plane has a dimension of 2. and so on) the fine detail of the complex curve would be lost due to the resolution limits of the printing process. and some of its current applications in neuroscience. At higher stages of construction (D. like the dendritic field area or the size of the soma. Helge von Koch in 1904. unavoidably finite and limited in scale by their own nature. Computer-generated fractals. We also review some of the methodologies available for calculating the fractal dimension. 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. 14) it should be noted that the fractal dimension is only a descriptive parameter. They are generally held to be statistically self-similar. in contrast with mathematical fractals. Many patterns in biology display a limited self-similarity or approximate self-similarity. Thus fractals are always described by power functions since homogeneous power laws lack natural scales. D is called fractal because it usually is not an integer. it is equal to the standard Euclidean dimension for which an ideal point has a dimension of 0. The fractal dimension for this purpose is therefore not intended to indicate whether the image is a fractal object.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. are sometimes termed prefractals since they are limited resolution images and therefore do not realize the detail implicit in the complete mathematical formulation (15). E. which increases in value with increasing structural complexity and describes the “fractured” nature of objects in nature (10). It is called dimension because it provides a measure of how completely an object fills space. The sequential construction of this fractal begins with a straight line (A). an ideal line has a dimension of 1. and a perfectly solid volume has a dimension of 3. 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. time. Construction of the Koch curve with a D of 1.26 that was described by the Swedish mathematician. FRACTAL DIMENSIONS An important parameter in fractal analysis of biological structures is the fractional or fractal dimension (D).

For example. we consider only those that are potentially useful in neuroscience.26. It is not easy to give a precise definition of a fractal (15). Since many of these fractal dimensions are used mainly in pure mathematics or applied physics. the volume goes either to zero or infinity. . and they do not completely cover the two-dimensional area. this would be when D ⫽ log 4/log 3 ⫽ 1. A straight line is drawn from the cell silhouette to its value on the D axis. 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. or other experimental data obtained from presentations of natural objects. Hausdorff (32) suggested that the volume or measure of the sphere should be eD where e equals the resolution of measurement. This definition of dimension was extended and put into a more systematic framework by Besicovitch (33). for instance. 20–31). Table 1 lists some of the most important fractal dimensions with their synonyms and context.USE OF FRACTAL THEORY IN NEUROSCIENCE Since. Examples of different cat ganglion cells.2. Calculating the Hausdorff dimension is generally FIG. drawings. For the Koch curve shown in Fig. 19).45 (Fig. would have relatively few dendritic branches and cover the two-dimensional area less completely than neurons with higher D values like 1. Measuring any self-similar set with spheres of integer dimension. neurons with low D values. say 1. Various other aspects of fractal analysis and D are discussed formally by other authors (15. METHODS FOR DETERMINING FRACTAL DIMENSIONS Although the mathematically rigorous determination of D is impossible for a fractal point set obtained 311 from digitized photographs. nerve cells seen in two dimensions are not straight lines. their D values fall between 1 and 2. a very good estimate of D can be achieved by different fractal analysis methods (Fig. 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). 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. with their associated box counting dimensions. 1. 2. drawn from throughout the retina. 16. 2). 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. 3). 18. All methods rely on the relationship between a measuring device and the object’s spatial distribution.

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

4. (B) Graph of resulting log–log plot. (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.

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

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

the entire border falls outside. The method first computes the center of gravity and then the radius of gyration. One is related to how image presentation may influence the possible scaling relationship of the image and the associated estimated D.nih. For strictly self-similar mathematical fractals such as the Koch curve. 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. Note that it is necessary to sample all local origins to sample as many data points belonging to the image as possible. the cell body interior and that of the dendrites do fill a plane completely and hence have a D of 2.6. on the other hand. 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 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. A multiplatform version for computing the mass fractal dimension is available from http://life. IMAGE PREPARATION AND DETERMINATION OF D Digitized images can be presented as binary. and the other is related to estimating D from the data points. with a D of 2 (60). Mandelbrot (5) stated that an object that fills a plane completely has a dimension value of 2.txt) and other user contributed macros are found at ftp://codon. However. the filled interior is solid. that increases with the increase in the radius r (see Fig. For instance. 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 circle or sphere of radius r is laid over the image. This is due to the area of the cell body and dendrites being much smaller than the extent of the border (58. M.html. There are various versions for various Macintosh computers. all appropriate fractal analysis methods approach the same limit. 8). border roughness (Fig.edu. To implement this method for the analysis of 2D images. When analyzing neurons. To lessen computation time a fraction. primary particles of a colloidal aggregate. the cell body and/or the axon may also be removed from binary or outline representations of neurons. adsorption sites on a surface. The choice of format is related to the spacefilling attributes of the image and the attributes of the image one deems to be important. With neurons specifically. . The sites may be pixels obtained from box counting. This particular macro (fractal dilation. However. had significantly lower D values (58) since they represented only the dendritic branching and do not reflect the other characteristic of complexity. 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. skeletonized or border-only images. named Fractop.gov/pub/ nih-image/user-macros/. 59). If one takes a fraction of the radius of gyration. steps of a random walk.316 ´ NDEZ AND JELINEK FERNA nih-image/download. Previous results (58) have demonstrated no significant difference between the estimated D of binary images. The histological techniques used may also lead to incomplete staining of the peripheral parts of the cell. of the radius of gyration. say 0. 3D). In addition. one could claim that it is only the border that is fractal. 57).au/ fractop/ and discussed by Jones and Jelinek in this issue. monomers in a polymer chain. This version. binary images with cell body and axon removed. depending on the relationship between the internal area and the contour. there are two further considerations. Usually the quantity of interest is the area of the image. This can happen with a known fractal such as a Koch snowflake. 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. Skeletonized images. The double logarithmic plot of M(r) against r gives a quantitative value for D.csu. etc. the Hausdorff dimension (29. has the added advantage of providing a choice for the number of centers and the fraction of the radius of gyration required. All possible choices of local origin are averaged and the average cluster mass M(r) is obtained. Having decided which analysis method to use. or border-only images of cat retinal ganglion cells as long as the dendrites are thin with respect to the cell body. this finding is dependent on the type of cell one analyzes and does not hold for glia cells (60).

This limited self-similarity or scale invariance is characteristic of biological material and is a focus of some controversy (51. In such a plot. outline-only. The figures on the bottom are the associated graphs of their fractal dimensions. was not scale-invariant under this transformation and method (58).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. The actual data points generally do not lie on a straight line for more than one to two decades. . and skeletonized (C) images. D is related to the slope of the line. some investigators have obtained linear plots using skeletonized images of neurons. 7) using the mass–radius method. also obtained linear log–log plots with skeletonized images. as described above. Analysis of skeletonized images using the original NIH Image box counting method (Version 1.2) led at times to a sigmoid log–log data point distribution. 8. Box counting analysis of the same turtle ganglion cell. obtained linear log–log plots (⬎ 2 generations) with skeletonized images of retinal neurons. outlined (B). This dependency on the analysis method to produce linear log–log plots with skeletonized images may explain the conclusions of Panico and Sterling (61). Differences in the linearity of the log–log data points was observed between binary. These authors used two variants of the box counting method and the mass–radius method with skeletonized images and concluded that FIG. using binary (A). and skeletonized images (58). Caserta et al. Montague and Friedlander (14). (6. when skeletonized. However. for instance. indicating the 317 same image. using a different implementation of the box counting method (greater number of box sizes) and different image handling (rotation of image and using multiple centers). the number of data points being related to the number of measuring steps. 61).

The left-hand side shows the digitized image of a retinal bipolar cell. Because of the limited scale invariance of neurons different authors have used different methods to determine D from log–log values. personal communication). If the linear fit is accepted then the image is fractal.41 (open circles) and 1. Alternatively. 53. The test is based on the ratio of reduced 2 values. 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.318 ´ NDEZ AND JELINEK FERNA cat retinal ganglion cells are not fractal due to their limited linearity. 61). (7) for the mass–radius method. of course. The D with the smallest linear range (1. ADVANTAGES AND POTENTIAL PROBLEMS OF FRACTAL DIMENSIONS FIG.07 (filled circles). When this method produces multiple values of D. The use of a hierarchical cluster analysis to compute particular subsets of the log–log values that achieve the best linear fittings (Fig. The region in which the local slopes are constant is then taken as the linear region (7). 9) has also been reported (12). use of the value with the longest linear range is suggested. as the difference in log N(r) divided by log (r) for every n successive points.25). Panico and Sterling (61) also used the local slope method to determine the D of their images. 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. A hierarchical cluster analysis yielded two regression lines with two different D values: 1. described by Caserta et al. 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). 9. and indeed it should not be expected to. Therefore the linearity region increases as the window is increased and makes this is a very subjective method. One of the main ones is that the sensitivity changes as the window over which the local slopes is obtained is decreased (62). be able to fit the data better. but it uses up one more degree of freedom in the process. it is a statistically significant parameter for identifying and differentiating neuronal cell classes. One method for this. has several flaws. Thus fractal analysis . is to calculate the n-point local slopes. however. from a statistical point of view such a method would not be justified. In a recent study (12) we posed the following question: Can the estimate of D resolve differences in neuronal branching when simpler metrical analysis alone cannot? Our results indicated that although D alone does not completely specify a cell’s morphology. the 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. Clearly then. The higher-degree polynomial will always. filled circles) could be attributed to finite size effects at very low scales. Deciding on the range of linearity and especially if it is significance has been addressed by Russ (49). The test is performed using a critical value of F ( p ⫽ 0.41. 37. Several publications have used this method to determine the slope of the data points (10. This method considers the D of the cell drawing to be the one with the longest linear range (1. The right-hand side shows a plot of the box counting measurements. This method. Method for the graphic determination of fractal dimensions. however. The simplest method of obtaining D is to fit a regression line to all data points and determine the slope of this line. The linear region can also be calculated by determining the local slopes. and the improvement in the fit may not be that great. which will have an F distribution (49). 52. open circles).07. as biological objects display statistical self-similarity only between a short range of dimensions. 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. The range of linearity is not important if the D obtained in this way is used in differentiating between different cell types (Caserta.

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). it remains that in many situations a single number. 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). A basic consideration is that most measurements cover only a relatively short range of dimensions. some of the images analyzed using fractal analysis may not demonstrate self-similarity or scale invariance over more than one or two levels of magnification and may not be fractal (61. To capture all this richness of this complex structure into a theoretical model is one of the major challenges of modern theoretical biology (64). Thus in almost all circumstances the fractional component of dimension is retained when a fractal object is projected to a lower-order dimension (18. 319 Furthermore whether a higher fractal dimension would correlate with a more complex physiological response is still an unresolved issue (9. 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). however. such us dendritic field extent and total dendritic length.USE OF FRACTAL THEORY IN NEUROSCIENCE has an important role in characterizing natural objects. 19). 12). 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). 42. descriptors such as D. space filling. like the dendritic field area or the number of segments of a dendritic tree. Thus. and various authors have discussed classification systems of neurons using fractal analysis (7. real data cannot be ideally fractal over all scales. 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. could immensely aid in the morphological discernment of different neuron types or neurons that . It should. Unlike mathematically generated fractals. In general. Thus determining D of a neuron. 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. 39. Thus many quantitative parameters have been used to characterize the morphology of nerve cells. the results are always consistent. Notwithstanding the above-mentioned limitations. Although all analysis methods rely on the relationship between a measuring device and the object’s spatial distribution. biological data that have a linear fit of more than two orders of magnitude are extremely rare (66–69). 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. CONCLUSIONS AND FUTURE DEVELOPMENTS Fractal analysis has already found widespread application in the field of neuroscience and is being used in many other areas. not all methods give identical results for the same form. Furthermore fractal geometry has some other advantages over its integer-dimensional counterparts. 62. in addition to the other morphometric criteria typically used. and does not necessarily imply any underlying mechanism of form generation. 63). 59. to more complicated global. These parameters range from simple metrical descriptors. For example. 64). an example being the projection of three-dimensional retinal ganglion cells onto a two-dimensional film or drawing (7). be kept in mind that D is only a descriptive parameter. or complexity of neurons. the fractal dimension. 62). 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). summarizes concisely and meaningfully the amount of detail. 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. Finally. 12. the connection between empirical values of D and any specific growth mechanism should be avoided and require the answering of further experimental questions.

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