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# METHODS 24, 309–321 (2001

)

doi:10.1006/meth.2001.1201, available online at http://www.idealibrary.com on

**Use of Fractal Theory in Neuroscience:
**

Methods, Advantages, and Potential Problems

Eduardo Ferna´ndez* and Herbert F. Jelinek†,1

*Instituto de Bioingenierı´a, Universidad Miguel Herna´ndez, Elche, Spain; and

†School of Community Health, Charles Sturt University, Albury, Australia

**Fractal analysis has already found widespread application
**

in the field of neuroscience and is being used in many other

areas. Applications are many and include ion channel kinetics

of biological membranes and classification of neurons according to their branching characteristics. In this article we

review some practical methods that are now available to allow

the determination of the complexity and scaling relationships

of anatomical and physiological patterns. The problems of describing fractal dimensions are discussed and the concept of

fractal dimensionality is introduced. Several related methodological considerations, such as preparation of the image and

estimation of the fractal dimensions from the data points,

as well as the advantages and problems of fractal geometric

analysis, are discussed. 䉷 2001 Academic Press

**One of the basic tenets of neurobiology holds that
**

the function of a nerve cell is largely dependent on

its structure. To understand how a neuron integrates

its myriad synaptic inputs to generate an appropriate response, a thorough understanding of the

cell’s morphology and geometry is required. Thus

many quantitative parameters have been used to

characterize the morphology of nerve cells. The simple models that have been used so far for shapes,

such as spheres, ellipsoids, and polyhedra, and their

corresponding two-dimensional profiles, are useful

for many purposes, including estimates of volume or

size distribution, but certainly fall short in dealing

1

To whom all correspondence and reprint requests should be

addressed at School of Community Health, Charles Sturt University, Albury, 2640, N.S.W., Australia. Fax: ⫹61 2 516772. E-mail:

hjelinek@csu.edu.au.

**1046-2023/01 $35.00
**

Copyright 䉷 2001 by Academic Press

All rights of reproduction in any form reserved.

**with neuronal complexity. One method for describing
**

the irregular shape of feature profiles has been to

“unroll” that shape by plotting distance from the

centroid as a function of angle, and then to perform

a Fourier analysis on the resulting curve (1). Besides

being computationally demanding, this approach has

difficulty in dealing with shapes so irregular that

the radius line may intersect the outline more than

once. Other approaches such as the form factor are

also not useful since it can be the same for different

shapes and is not able for instance to measure quantitatively the complexity or degree of dendritic

branching (1). Sholl analysis has been widely used

to analyze branching characteristics of neurons. This

method determines the number of intersections of

dendrites with a set of concentric circles (2). Sholl

analysis is not an ideal method to measure neuronal

complexity as it has several problems with determining the number of processes at each level of measurement. Furthermore it is insensitive to the border

ruggedness of neurons (3, 4).

A new approach to this problem requires us to

think about the concepts of fractal geometry. In the

short period since Mandelbrot defined the term fractal (5), the field of fractal geometry has enjoyed an

enormous surge of popularity. The key observation

is that structures growing according to stochastic

processes are not really as disordered as they seem at

first glance. A nontrivial, scale invariant symmetry

over several orders of magnitude has been found to be

a typical principle of order for such growth processes,

which can be quantified by the fractal dimension.

Although this parameter can be used to quantify the

309

an ideal line has a dimension of 1. unavoidably finite and limited in scale by their own nature. Helge von Koch in 1904. 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. Further it should be kept in mind that all natural objects are. It is called dimension because it provides a measure of how completely an object fills space. FIG. We also review some of the methodologies available for calculating the fractal dimension. and does not necessarily imply any biological process nor mechanism involved in their development. they do not have a characteristic unit of length. This addition of detail results in an ideal fractal object having an infinite boundary length (16. . THEORETICAL CONSIDERATIONS OF FRACTAL GEOMETRY AND NATURALLY OCCURRING FRACTALS An object is said to be fractal if certain criteria such as the object being self-similar or scale invariant are met.26. E. Thus fractals are always described by power functions since homogeneous power laws lack natural scales. Thus natural patterns display statistical self-similarity only between an upper and lower bound. 1. time. 14) it should be noted that the fractal dimension is only a descriptive parameter. FRACTAL DIMENSIONS An important parameter in fractal analysis of biological structures is the fractional or fractal dimension (D). Limitations are also imposed by recording and imaging techniques. 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 0.26 that was described by the Swedish mathematician. in contrast with mathematical fractals. and a perfectly solid volume has a dimension of 3. 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. like the dendritic field area or the size of the soma. are sometimes termed prefractals since they are limited resolution images and therefore do not realize the detail implicit in the complete mathematical formulation (15). Mandelbrot has shown that the boundary length of a fractal object can be mathematically expressed as a power law. 17). such as the Koch curve. and some of its current applications in neuroscience. and so on) the fine detail of the complex curve would be lost due to the resolution limits of the printing process. that is. 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. Computer-generated fractals. Construction of the Koch curve with a D of 1.310 ´ 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. or mass (16). When D takes an integer value. 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). the advantages and problems of fractal geometry. which increases in value with increasing structural complexity and describes the “fractured” nature of objects in nature (10). They are generally held to be statistically self-similar. The sequential construction of this fractal begins with a straight line (A). an ideal plane has a dimension of 2. Then the middle third is raised to produce an equilateral triangle (B). Many patterns in biology display a limited self-similarity or approximate self-similarity. At higher stages of construction (D.

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

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

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

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

7. (B) Scaling factor ⫽ 2 m. 5. Each image frame has pixels with a different scaling factor. (A) Scaling factor ⫽ 1 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. Digitized images of a turtle ganglion cell. (A) Binary image of a turtle ganglion cell. 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. FIG. FIG. . (C) Scaling factor ⫽ 4 m. 6.

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

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

Because of the limited scale invariance of neurons different authors have used different methods to determine D from log–log values. The use of a hierarchical cluster analysis to compute particular subsets of the log–log values that achieve the best linear fittings (Fig. which will have an F distribution (49).41.07 (filled circles). as the difference in log N(r) divided by log (r) for every n successive points. The D with the smallest linear range (1. 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). 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. This method considers the D of the cell drawing to be the one with the longest linear range (1. The linear region can also be calculated by determining the local slopes. it is a statistically significant parameter for identifying and differentiating neuronal cell classes. Clearly then. The test is based on the ratio of reduced 2 values. Deciding on the range of linearity and especially if it is significance has been addressed by Russ (49). described by Caserta et al. Method for the graphic determination of fractal dimensions. One method for this. 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. Alternatively. be able to fit the data better. 52. Thus fractal analysis . open circles). as biological objects display statistical self-similarity only between a short range of dimensions. The left-hand side shows the digitized image of a retinal bipolar cell.318 ´ NDEZ AND JELINEK FERNA cat retinal ganglion cells are not fractal due to their limited linearity. 61). however. ADVANTAGES AND POTENTIAL PROBLEMS OF FRACTAL DIMENSIONS FIG. This method. Therefore the linearity region increases as the window is increased and makes this is a very subjective method. and indeed it should not be expected to. 9. of course. Panico and Sterling (61) also used the local slope method to determine the D of their images. The simplest method of obtaining D is to fit a regression line to all data points and determine the slope of this line. but it uses up one more degree of freedom in the process. The right-hand side shows a plot of the box counting measurements. has several flaws. is to calculate the n-point local slopes. The higher-degree polynomial will always. The range of linearity is not important if the D obtained in this way is used in differentiating between different cell types (Caserta. 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.41 (open circles) and 1. One of the main ones is that the sensitivity changes as the window over which the local slopes is obtained is decreased (62).25). 53. 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. Several publications have used this method to determine the slope of the data points (10. 9) has also been reported (12). If the linear fit is accepted then the image is fractal. The region in which the local slopes are constant is then taken as the linear region (7). and the improvement in the fit may not be that great. use of the value with the longest linear range is suggested. The test is performed using a critical value of F ( p ⫽ 0. 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. filled circles) could be attributed to finite size effects at very low scales. 37. from a statistical point of view such a method would not be justified. personal communication). When this method produces multiple values of D. A hierarchical cluster analysis yielded two regression lines with two different D values: 1.07. (7) for the mass–radius method. however.

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

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