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
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:
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
they do not have a characteristic
unit of length.
and does not necessarily imply any biological process
nor mechanism involved in their development. 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. that is. and so on) the fine
detail of the complex curve would be lost due to the resolution
limits of the printing process.
Helge von Koch in 1904. At higher stages of construction (D. an ideal
line has a dimension of 1. Thus natural patterns display statistical self-similarity only between
an upper and lower bound. They are generally held to be
statistically self-similar. and some of its current applications in
addition of detail results in an ideal fractal object
having an infinite boundary length (16. unavoidably finite and limited in scale by their own nature. Construction of the Koch curve with a D of 1.
it is equal to the standard Euclidean dimension for
which an ideal point has a dimension of 0. or mass (16). an ideal plane has a dimension of 2. Many patterns in
biology display a limited self-similarity or approximate self-similarity. Figure 1 shows an approximation of an ideal/
theoretical fractal with a fractal dimension of 1. Then the middle third is raised to produce an equilateral
triangle (B). time. Thus fractals are always described by power
functions since homogeneous power laws lack natural scales. Computer-generated fractals. Mandelbrot has shown that the boundary length of a fractal
object can be mathematically expressed as a power
sequential construction of this fractal begins with a straight line
(A). 17). E.
. are sometimes termed
prefractals since they are limited resolution images
and therefore do not realize the detail implicit in the
complete mathematical formulation (15). Further it should be kept
in mind that all natural objects are. 14) it should be noted that
the fractal dimension is only a descriptive parameter.
like the dendritic field area or the size of the soma. D is called fractal
because it usually is not an integer. and a perfectly solid volume has a dimension of 3. The fractal dimension for this purpose is
therefore not intended to indicate whether the image
is a fractal object. the advantages and problems of fractal geometry.
An important parameter in fractal analysis of biological structures is the fractional or fractal dimension (D). Raising equilateral triangles from the middle third
of each of the line segments in the object produces the image in
(C). It is called dimension because it provides a measure of how completely
an object fills space. 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.26. Limitations are also imposed by recording and imaging techniques.
THEORETICAL CONSIDERATIONS OF
FRACTAL GEOMETRY AND NATURALLY
An object is said to be fractal if certain criteria such
as the object being self-similar or scale invariant are
met. When D takes an integer value.
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. which increases in value with increasing
structural complexity and describes the “fractured”
nature of objects in nature (10).310
´ NDEZ AND JELINEK
complexity of the borders of a neuron (6–12) and to
measure how completely the branches of a neuron
fill its dendritic field (13. We also review some of the methodologies available for calculating the fractal dimension. in contrast with
mathematical fractals. such as the Koch curve.26
that was described by the Swedish mathematician.
and there are in the literature many different
types of fractal dimensions so that even research
mathematicians are not agreed on their names or
equivalence (18. 20–31).45 (Fig. 1. we consider only those that are potentially
useful in neuroscience. Hausdorff (32) suggested that the volume or measure of the sphere
should be eD where e equals the resolution of measurement. neurons with
low D values. drawings. for instance. their D values fall between 1 and 2. nerve cells seen in two dimensions are not straight lines. the volume
goes either to zero or infinity.26.
METHODS FOR DETERMINING FRACTAL
Although the mathematically rigorous determination of D is impossible for a fractal point set obtained
from digitized photographs. 2. 18. The
fractal dimensions are sufficiently different to suggest that they represent distinct ganglion cell types. 16. or other experimental data obtained from presentations of natural objects. with their associated box counting dimensions. Measuring any self-similar set with spheres of integer dimension.2. a very good estimate of D can be achieved
by different fractal analysis methods (Fig. The D-dimensional Hausdorff measure of
an image is finite only when D (the dimension value)
equals the dimension of the image. would have relatively few dendritic branches and cover the two-dimensional area
less completely than neurons with higher D values
like 1. 19). For example.
Calculating the Hausdorff dimension is generally
FIG. For the Koch
curve shown in Fig. All
methods rely on the relationship between a measuring device and the object’s spatial distribution. and they do not completely cover the two-dimensional area. Various other aspects of fractal analysis and D are discussed formally by other
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).USE OF FRACTAL THEORY IN NEUROSCIENCE
. drawn from throughout the retina.
It is not easy to give a precise definition of a fractal
(15). say 1. 3). A straight line is drawn from the
cell silhouette to its value on the D axis. It is calculated by covering an object with countable spheres whose radii are not greater than the
image but decrease to zero. This definition of dimension was extended and put into a more systematic framework
by Besicovitch (33). Table 1 lists some of the most
important fractal dimensions with their synonyms
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). Since many of these fractal dimensions
are used mainly in pure mathematics or applied
physics. 2). Examples of different cat ganglion cells. this would be when D ⫽ log
4/log 3 ⫽ 1.
dimension. from T. A. dilation
Richardson dimension. Losa. The difference from the Hausdorff–
Besicovitch dimension is that the set is now covered
with spheres of identical radius (16). perimeter
Capacity dimension. 1961
Mandelbrot. (C) Dilation method. F.
and G. 1983 (5)
Smith et al. was introduced by Kolmogorov (34. The length of the
coastline then equals the size of the ruler times the
number of steps r has taken to trace the coast. The “capacity
dimension” has become the fundamental definition of
fractal dimension in the minds of many. (A) Calliper method. 1992 (48)
Caserta et al..
but it cannot be strictly applied
to natural objects due to its finite
range of fractal structure
Easier to evaluate than DH. R. 1990 (16)
Smith et al. 1990 (16)
Peitgen et al. 1983 (5)
Takayasu. and Weibel. with various
diameters and centered on the border of the Koch island. can also be applied
to surfaces and biological
Mandelbrot. Jr. 1935 (33)
Mandelbrot. Birhauser. Figures 3A and 4 show examples of this method.
D(0) in multifractal analysis
Mass fractal dimension.
FIG. 1919 (32)
Besicovitch. D(2) in
Generic term for fractal dimension
Widely used in pure mathematics. To determine D. One
finds that the boundary length is a function of the
span of the calliper employed in the measurement. 1983 (5)
Tatsumi et al. 1989 (45)
Smith et al. with permission. or yardstick dimension). The capacity
dimension is related to the box counting and mass–
radius methods that are its applied. See text for
more details. T. Lange (1998) in Fractals in Biology and Medicine (Nonnenmacher.
This method is based on counting the number of steps that give
a polygonal representation of an arbitrary object using different
very difficult and a more practical parameter of D... Reprinted.. Note loss of
border detail shown in (A) and (B) (D) Mass method example
after application of six groups of concentric disks. two-dimensional
embodiment and described below. All
centers lie within the radius of gyration (large circle).
..). 35). 1989 (10)
An algorithm based on the Hausdorff dimension
is the calliper dimension (also known as the compass. 1991 (29)
Richardson. 1989 (10)
dimension..(B) Box counting method. D. 1983 (5)
usually is greater than or
equal to the Hausdorff
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. Some methods used for determination of fractal dimensions of a Koch triadic island with a D ⫽ 1. Eds. a
ruler of decreasing size r is used to measure the
boundary or coastline of an image.
the capacity dimension. 3.. Basel. 1989 (10)
Takayasu. 1983 (5)
Mandelbrot. 1990 (6)
Jelinek and Fernandez. the length does not converge to a stable value
but keeps increasing as the calliper span decreases. mass
radius dimension.´ NDEZ AND JELINEK
Some of the Most Widely Used Fractal Dimensions with their Synonyms and Contexts
divider. 1995 (7)
Caserta et al.. G. mosaic
amalgamation dimension. E.
That is. After
dilation with a disk kernel diameter of 16 pixels.. G. compass
USE OF FRACTAL THEORY IN NEUROSCIENCE
. Calliper method for ascertaing the boundary length of an image. (B) Graph of resulting log–log plot. (A) Measuring the length of the coastline of the Australian
For a fractal curve the length
will continue to increase as the radius of the circles decreases. 43.
6B. 3C.harvard. N(r). The NIH Image program
and its many macros can be fetched in a number of
Many research reports using this scheme to analyze neuron structures are found in the literature
(10. The pixel dilation method. S. The logarithm of N(r) versus the logarithm of r
gives a line whose gradient corresponds to D. 44).
and the slope (on the usual log–log plot) gives the
dimension. 6). as reported by
the above articles. and K
The Minkowski–Bouligand dimension is different
from the Hausdorff dimension (18). The fractal dimension is
then estimated from the slope of the log–log plot of
length against diameter. 36). A common form of
47–49). the Euclidean space containing the
image is divided into a grid of boxes of size r. D is then calculated by fitting a
linear regression to the following equation: log(r)
⫽ ⫺D* log(n) ⫹ K. as devised by Flook (50). noncontiguous structures
or closed loops within a structure). 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. (10) and others (11. called
the Minkowski “sausage.
is a constant. 7). The calliper method has previously been
used to characterize neurons (10. 36.nih.
3B). 46). where r ⫽ resolution of image
(number of pixels per unit length). A Macintosh program for calculating D
using this method can be found at the following
URL: http://plantecohost. This value
is then plotted as a function of the circle diameter. or
gradient is related to the fractal dimension by D ⫽
1 ⫺ S (5). This is done by application of a convolution
procedure which is part of the image analysis program (dilation macro from NIH). has been
implemented by Smith et al.nih. the result will be the
length of the curve. Euclidean curve. replaces each pixel of the border
by a circle whose diameter ranges from 3 to 61 pixels
(Fig. is that images composed of more than one simple perimeter cannot be
processed accurately (e. for use with NIH Image image processing software.txt. 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.
This is equivalent to the “grid” method described by
Smith et al.
51–56).html. (5. 37–42).. In practice.edu/gmbWWW/
APPL. the image is digitized by
pixels having a given scaling factor r (Fig. 12. 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.gov/pub/nih-image/user-macros/
box count macro. A circle is swept continuously
along the line and the area that is covered.
Box Counting Method
To estimate D.
The following methods can be used for noncontiguous structures as well as for 2D and 3D images. The method is
illustrated in Fig.
A method similar to the box counting technique
is the grid intercept method (45. Figure 5
shows an example of this calculation for a retinal
ganglion cell.g. 6A).gov/
. The slope. with
the initial box size being the size of the image. 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 macro for this method can be obtained from the National Institutes of Health (NIH)
at ftp://codon. For
a smooth. The
sequence of box sizes for grids is usually reduced by
a factor of 1/2 from one grid to the next. This filters out
structures smaller than the current diameter of the
circle. The important difference between this
and the calliper method is that the circle is moved
so that its center lies on every point of the line. the points
will fall on a straight line between an upper and
lower bound with negative slope.
Pixel Dilation Method
The pixel dilation method is based on the Minkowski–Bouligand dimension. 6C). The box counting method applies to
any structure in the plane and can be adapted for
structures in three-dimensional space (7. 10. 29). These are detailed at http://rsb.” is determined. The length of the border for each respective
diameter is determined by the area of the outline
divided by the diameter.314
´ NDEZ AND JELINEK
If the length of the boundary (coastline) versus the
calliper length is plotted on a log–log scale. r is
then made progressively smaller and the corresponding number of nonempty boxes.info. is counted (Fig. (10). One major
drawback of the calliper method.
(C) Scaling factor ⫽ 4 m.
FIG.USE OF FRACTAL THEORY IN NEUROSCIENCE
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. 5.
The perimeter is calculated as the area of this figure divided by
the diameter of the dilating disk. (A) Scaling factor
⫽ 1 m. 6. Digitized images of a turtle ganglion cell. (B) Scaling factor ⫽ 2 m. Each image
frame has pixels with a different scaling factor.
. (A) Binary image of a turtle ganglion cell. (B) Minkowski
“sausage” created by application of the dilation method to (A). 7.
monomers in a
polymer chain. In
addition. binary images with
cell body and axon removed. Previous results (58)
have demonstrated no significant difference between
the estimated D of binary images. the cell body interior and that of the
dendrites do fill a plane completely and hence have
a D of 2. Note that it is necessary to sample all local
origins to sample as many data points belonging to
the image as possible.edu. skeletonized or border-only images.html. However. The sites may be pixels obtained from box
counting. All possible choices
of local origin are averaged and the average cluster
mass M(r) is obtained. The histological techniques used may also lead to incomplete staining of
the peripheral parts of the cell. adsorption sites on a surface. the Hausdorff
dimension (29. When analyzing neurons. Usually the
quantity of interest is the area of the image.
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
Mandelbrot (5) stated that an object that fills a plane
completely has a dimension value of 2. The method first computes the center of
gravity and then the radius of gyration. etc. To lessen
computation time a fraction. 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. For instance.
IMAGE PREPARATION AND DETERMINATION
Digitized images can be presented as binary. the filled interior is solid.gov/pub/
nih-image/user-macros/.6. Skeletonized images.
For strictly self-similar mathematical fractals
such as the Koch curve.txt) and other user contributed macros are found at ftp://codon.au/
fractop/ and discussed by Jones and Jelinek in this
roughness (Fig. had
significantly lower D values (58) since they represented only the dendritic branching and do not reflect
the other characteristic of complexity.csu. 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.nih. 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. The choice of format is related to the spacefilling attributes of the image and the attributes of
the image one deems to be important. There are various versions for various Macintosh computers. 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. 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. with a D of
2 (60). A
multiplatform version for computing the mass fractal
dimension is available from http://life. of the radius
of gyration. However. If one takes
a fraction of the radius of gyration. the entire border
falls outside. 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. depending on the relationship between the
internal area and the contour. on the other hand. has the added
advantage of providing a choice for the number of
centers and the fraction of the radius of gyration
required. one could claim that it is only the border
that is fractal. say 0. Having decided which analysis
method to use. This particular macro (fractal dilation.316
´ NDEZ AND JELINEK
nih-image/download. With neurons
One is related to how image presentation may influence the possible scaling relationship of the image
and the associated estimated D.
To implement this method for the analysis of 2D
images. This can happen with a known fractal
such as a Koch snowflake. or border-only images of
cat retinal ganglion cells as long as the dendrites are
thin with respect to the cell body. M. primary
particles of a colloidal aggregate. the cell body and/or the axon may also be removed from binary or outline representations of
. This is due to
the area of the cell body and dendrites being much
smaller than the extent of the border (58. there are two further considerations. The double logarithmic plot
of M(r) against r gives a quantitative value for D. and the other is
related to estimating D from the data points. 8). that
increases with the increase in the radius r (see
Fig. 3D). steps of a random walk. named Fractop. 59). all appropriate fractal analysis methods approach the same limit. This version.
and skeletonized images (58). obtained linear log–log plots (⬎ 2 generations)
with skeletonized images of retinal neurons. 8. for instance. and skeletonized (C) images. Analysis of skeletonized images using the original NIH Image box
counting method (Version 1. 61). These authors used two variants
of the box counting method and the mass–radius
method with skeletonized images and concluded that
FIG. also obtained linear log–log plots with
skeletonized images. (6. 7) using the mass–radius method. Montague and Friedlander (14). was not scale-invariant under this transformation and method (58). indicating the
same image. when skeletonized.
. 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). some investigators have obtained linear
plots using skeletonized images of neurons.2) led at times to a sigmoid log–log data point distribution. outlined (B). The actual data
points generally do not lie on a straight line for more
than one to two decades. This limited self-similarity
or scale invariance is characteristic of biological material and is a focus of some controversy (51. using binary (A). D is related to the slope of
Differences in the linearity of the log–log data
points was observed between binary.USE OF FRACTAL THEORY IN NEUROSCIENCE
GRAPHIC DETERMINATION OF THE
How the actual D value is obtained from the log–
log data points can lead to differences in the magnitude of D. outline-only.
However. The figures
on the bottom are the associated graphs of their fractal dimensions. Caserta
et al. the number of data points being related
to the number of measuring steps. This dependency on the analysis method to produce linear log–log plots with skeletonized images may explain the conclusions of Panico
and Sterling (61). Box counting analysis of the same turtle ganglion cell. In such a plot. as described above.
Deciding on the range of
linearity and especially if it is significance has been
addressed by Russ (49). (7) for
the mass–radius method. 61). The left-hand side shows the digitized image of a retinal
bipolar cell. One of the main ones is that the sensitivity changes as the window over which the local
slopes is obtained is decreased (62). 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.
The higher-degree polynomial will always.
be able to fit the data better.
from a statistical point of view such a method would
not be justified. as the difference in log N(r) divided by
log (r) for every n successive points. This method. Method for the graphic determination of fractal dimensions. 9.25). The
D with the smallest linear range (1. 37. The right-hand side shows a plot of the box counting
measurements. 53. 9)
has also been reported (12).41 (open circles) and 1. however. The use of a hierarchical cluster
analysis to compute particular subsets of the log–log
values that achieve the best linear fittings (Fig.
use of the value with the longest linear range is suggested. 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. of course. 52.07
(filled circles). it is a statistically significant parameter for identifying and differentiating neuronal cell classes. open circles).318
´ NDEZ AND JELINEK
cat retinal ganglion cells are not fractal due to their
limited linearity. The simplest method
of obtaining D is to fit a regression line to all data
points and determine the slope of this line.41. The test is
based on the ratio of reduced 2 values. The region in
which the local slopes are constant is then taken as
the linear region (7). Several publications have used this
method to determine the slope of the data points
(10. described by Caserta et al. as biological objects display statistical self-similarity only between a short range of dimensions. filled circles) could be
attributed to finite size effects at very low scales. The test is performed
using a critical value of F ( p ⫽ 0.
Because of the limited scale invariance of neurons
different authors have used different methods to determine D from log–log values.
Clearly then. which will
have an F distribution (49).
and indeed it should not be expected to. Thus fractal analysis
. personal communication). and the improvement in the fit may not be that great. 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).
ADVANTAGES AND POTENTIAL PROBLEMS
OF FRACTAL DIMENSIONS
FIG. The range of linearity is not important if the D obtained in this way is used in differentiating between different cell types (Caserta. The linear region can also be
calculated by determining the local slopes. 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. Alternatively. but it uses up one more
degree of freedom in the process. If the linear
fit is accepted then the image is fractal. One
method for this.07. This method considers the D of the cell drawing to
be the one with the longest linear range (1. A hierarchical cluster analysis yielded two regression lines with two different D values: 1. has
When this method produces multiple values of D. however. is to calculate the n-point
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.
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. Therefore the
linearity region increases as the window is increased
and makes this is a very subjective method.
values are obtained by using the mass fractal methods than by using the pixel dilation and box counting
Although all analysis methods rely on the relationship between a measuring device and the object’s
spatial distribution. the
fractal dimension. that can be used for an objective
assessment of the degree of complexity (a concept
heretofore not readily quantifiable) of developing and
mature neurons. the connection
between empirical values of D and any specific
growth mechanism should be avoided and require
the answering of further experimental questions. descriptors such as D. 64). 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.
It should. This contrasts with integer-dimensional
measurement of anisotropic objects which require
multiple samples through the thickness of the threedimensional objects (1). 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). the results
are always consistent. however.
A basic consideration is that most measurements
cover only a relatively short range of dimensions. or complexity of neurons. 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. such us dendritic field extent and total
dendritic length. an
example being the projection of three-dimensional
retinal ganglion cells onto a two-dimensional film or
drawing (7). to more complicated global.USE OF FRACTAL THEORY IN NEUROSCIENCE
has an important role in characterizing natural objects. 12. biological data that have a linear fit of more
than two orders of magnitude are extremely rare
(66–69). real data
cannot be ideally fractal over all scales.
These parameters range from simple metrical descriptors.
A further advantage of fractal analysis is that
shrinkage or expansion of a specimen will not affect
D as long as the artifact acts equally in all directions
and the measured points still lie on the linear segment of the graph (19). it remains that in many situations a single number. 59. 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. For example. 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). Thus. space filling. Many neurons display
irregular shapes and discontinuous morphogenetic
patterns in support and in connection with their
functional diversity. 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). Notwithstanding the above-mentioned limitations. 42.
Furthermore whether a higher fractal dimension
would correlate with a more complex physiological
response is still an unresolved issue (9. summarizes concisely and meaningfully the amount of detail. 12).
Furthermore fractal geometry has some other advantages over its integer-dimensional counterparts. 62).
CONCLUSIONS AND FUTURE
Fractal analysis has already found widespread application in the field of neuroscience and is being
used in many other areas. and
does not necessarily imply any underlying mechanism of form generation. In general. Finally. like the dendritic field area
or the number of segments of a dendritic tree. Thus many quantitative parameters have been
used to characterize the morphology of nerve cells. and various authors have discussed classification systems of neurons using fractal analysis (7. 63). be kept in mind that D is only
a descriptive parameter. 19).
Unlike mathematically generated fractals. 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. not all methods give identical
results for the same form.
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 capture all this richness of
this complex structure into a theoretical model is one
of the major challenges of modern theoretical biology
(64). 62. in
addition to the other morphometric criteria typically
used. could immensely aid in the morphological discernment of different neuron types or neurons that
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