Biomimicry and Fuzzy Modeling:

A Match Made in Heaven

Michael Margaliot∗
November 18, 2007

Abstract
Biomimicry, the design of artificial systems that mimic natural
behavior, is recently attracting considerable interest. Biomimicry re-
quires a reverse engineering process; the behavior of a biological agent
is analyzed in order to mimic this behavior in an artificial system. In
many cases, biologists have already studied the relevant behavior and
provided a detailed verbal description of it. Mimicking the natural be-
havior is then reduced to the following problem: how can we convert
the given verbal description into a well-defined mathematical formula
or algorithm that can be implemented by an artificial system?
Fuzzy modeling (FM), with its ability to handle and manipulate
verbal information, constitutes a natural approach for addressing this
problem. The application of FM in this context may lead to a sys-
tematic approach for biomimcry, namely, given a verbal description
of an animal’s behavior (e.g., the foraging behavior of ants), apply
FM to obtain a mathematical model of this behavior which can be
implemented by artificial systems (e.g., autonomous robots).
The purpose of this position paper is to highlight these issues and
to alert the attention of the computational intelligence community to
this emerging application of FM.

Keywords: Linguistic modeling, vagueness, animal behavior, bio-inspired

systems, mathematical modeling in biology.

∗
Correspondence: Dr. Michael Margaliot, School of Elec. Eng.-Systems, Tel
Aviv University, Tel Aviv 69978, Israel. Tel: +972 3 640 7768. Homepage:
www.eng.tau.ac.il/∼michaelm Email: michaelm@eng.tau.ac.il

1
1 Introduction
Biomimicry1 is a new discipline that studies designs and processes in nature
and then mimics them in order to solve human problems. Studying how
plants absorb and utilize solar energy in order to develop better solar cells is
an example of such engineering inspired by nature.
In order to implement a natural behavior in an artificial system, it is first
necessary to understand the behavior. In many cases, the relevant behav-
ior has already been studied by biologists. Biological theories are generally
descriptive in nature, and are stated using natural language. For example,
Darwin’s original presentation of his evolutionary theory did not include a
single equation [9]. The problem of mimicking the natural behavior is then
reduced to the following problem.

Problem 1 Given a verbal description of some relevant behavior, design an

artificial system that mimics this behavior.
Fuzzy logic theory has been associated with human linguistics ever since
it was first suggested by Zadeh [66, 67]. Decades of successful applications
(see, e.g., [12, 54, 55, 64, 65, 44]) suggest that the real power of fuzzy logic
is in its ability to handle and manipulate verbally-stated information based
on perceptions rather than equations [11, 34, 35, 68]. In particular, fuzzy
modeling (FM) is routinely used to transform the knowledge of a human
expert, stated in natural language, into a fuzzy expert system that imitates
the human experts’ functioning [51, 26].
This suggests that FM may be suitable for addressing Problem 1, and
thus provide a systematic approach for addressing biomimicry. Given a ver-
bal description of the behavior of a biological agent, apply FM to obtain
a mathematical model of this behavior which can be implemented by an
artificial system.
The purpose of this position paper is to summarize some of the lessons
learned from using FM to develop mathematical models of animal behavior,
and to suggest the next natural step, that is, the use of FM in biomimcry.
Special attention is devoted to presenting the potential advantages of FM in
this specific context.
The remainder of this paper is organized as follows. Section 2 briefly
reviews the motivation for imitating processes and designs found in nature.
1
from the Greek bio, life, and mimesis, to imitate.

2
Section 3 describes several approaches for transforming verbal descriptions
into mathematical equations or algorithms. Section 4 describes two recent ap-
plications of FM for transforming verbal descriptions of animal behavior into
well-defined mathematical models. Sections 5 and 6 summarize some lessons
learned from using FM in modeling animal behavior, with an emphasis on
the unique advantages and disadvantages of the FM approach. Section 7 dis-
cusses the suitability of FM in the specific context of biomimcry. The final
section concludes.

2 Biomimicry
Over the course of evolution, living systems have developed efficient and
robust solutions to various problems they encountered in their natural habi-
tats. Plants have developed means for absorbing and utilizing solar energy.
Foraging animals have developed ways of navigating and searching in an un-
known environment. More generally, many natural beings have developed
the capabilities to reason, learn, evolve, adapt, and heal.
Scientists and engineers are interested in implementing such capabilities
in artificial systems. For example, designing efficient schemes for navigat-
ing and searching in an unknown environment is a fundamental problem in
the field of autonomous robots. It is thus natural that considerable research
has recently been devoted to the development of artificial products or ma-
chines that mimic (or are inspired by) various biological phenomena [52, 41].
For example, walking robots offer tremendous potential in replacing humans
where it is unsafe, inaccessible, or too expensive to operate. Currently there
is still a lack of a strong scientific basis for designing such robots. Yet, nature
provides us with an abundance of highly efficient walking agents. In partic-
ular, many insects are capable of traversing very rough terrain with sparse
footholds. This has motivated considerable research on the design of artificial
walking robots by emulating the structure and functioning of insects [46, 4].
Other examples from the field Biomimicry or Bio-inspired systems, in-
clude: evolutionary computation that addresses optimization problems by
imitating some of the ideas in Darwin’s theory of evolution [30, 18]; the design
of artificial structures that mimic shape laws in nature and, in particular, the
structural properties of trees [36]; autonomous robots that mimic the behav-
ior of various animals [3, 8]; the design of computer security systems inspired
by the natural immune system [16, 22]; the design of optimization, search,

3
and clustering algorithms based on the behavior of social insects [6, 10]; and
the design of new materials inspired by nature, such as a self-cleaning glass
based on the drift-repellent surface of the lotus plant [15]. More information
and a suggested reading list can be found at http://www.biomimicry.net/
An important component in biomimicry is a reverse engineering process
of the relevant natural behavior. In some cases, biologists have already stud-
ied the relevant behavior and provided verbal descriptions of the underlying
mechanisms. For example, as noted in [50], it is customary to describe the
behavior of simple animals (e.g., ants) using simple rules of thumb [24]. In
this case, the problem of imitating the relevant behavior is reduced to Prob-
lem 1. In the next section, we review several approaches that are suitable for
addressing this problem.

3 From words to equations

During the 1950s, Jay W. Forrester and his colleagues developed system dy-
namics2 as a method for modeling the dynamic behavior of complex systems.
The basic idea is to represent the causal structure of the system using elemen-
tary structures that include positive, negative, or combined positive and neg-
ative feedback loops. These are depicted graphically, and then transformed
into a set of differential equations. The method was applied successfully
to numerous real-world applications in the social, economic, and industrial
sciences. However, the inherent vagueness of the linguistic description is ig-
nored, and exact terms and phrases (e.g., a temperature of 30◦ Celsius) are
treated in precisely the same way as vague verbal terms (e.g., warm weather).
A more systematic approach to modeling physical systems is qualitative
reasoning (QR) [31] which transforms qualitative descriptions of a system
into qualitative differential equations. These extend ordinary differential
equations in two ways: (1) functional relationships between variables can
be represented by functions that are monotonous, but do not have to be
completely specified; and (2) the variable values are described using a set of
landmark values rather than exact numerical values. QR was applied to sim-
ulate and analyze many real world systems. However, its applications seem
to suggest that it is suitable when modeling with accurate, yet incomplete,
knowledge rather than with a verbal description of the system.
2
To date, more than 30 books on system dynamics have appeared. A very readable
one is [48]. The journal Systems Dynamics Review contains up-to-date papers.

4
3.1 Natural language and fuzzy logic
As noted in [39], the world around us is plagued by various forms of uncer-
tainty, ambiguity, and vagueness. The latter property arises in the grouping
of elements. Suppose that D is some universe of discourse, P is some prop-
erty, and consider the set S := {x ∈ D : x has property P }. In many cases,
it is impossible to provide a precise binary definition of S. For example,
consider the case where D is the set of all the man-made items, and P is the
property: x is a chair. Any attempt to provide a clear cut definition of S
is doomed to fail. This is evident from some dictionary definitions for chair,
e.g., “any of various devices that hold up or support.”
A typical feature of vagueness is continuity: a small difference between
two objects should not lead to an abrupt change in the decision of whether
they belong to S or not. In other words, the transition from having prop-
erty P to not having it (or vice versa) is smooth.
One approach for modeling such continuity is based on characterizing the
relation between an object and its properties by means of a scale. Conse-
quently, the membership in the set S is no longer binary, but becomes a
matter of degree. The concept of a fuzzy set provides a useful mathematical
model for characterizing sets that allow graded membership.
The property of vagueness is most striking in the semantics of natu-
ral language. This suggests that an algorithmic or mathematical treatment
of natural language should include mechanisms that deal with the inherent
vagueness.
Zadeh laid down the foundations of fuzzy sets and fuzzy logic and linked
them to human linguistics [66, 67]. The pioneering work of Wenstop [61, 62]
was aimed at building verbal models (VMs) capable of representing and pro-
cessing information stated in natural language. A VM consists of three basic
components: (1) generative grammar–used for defining the semantics of the
verbal statements; (2) fuzzy logic-based inferencing–used in the deductive
process; and (3) linguistic approximation–used for attaching suitable linguis-
tic labels to the outputs. The entire process was implemented using the APL
computer language. Wenstop and Kickert developed VMs for several inter-
esting systems from the social sciences [62, 28][27, Ch. 7]. However, VMs are
not standard mathematical models, as their input and output are linguis-
tic values rather than numerical values. Consequently, they suffer from two
drawbacks: (1) there are no methods for analyzing the behavior of VMs, so
they can only be used for simulations; and (2) when modeling dynamic sys-

5
tems the degree of fuzziness (or uncertainty) increases with every iteration,
so the system can be simulated effectively only over relatively short time
spans.
Kosko [29] suggested fuzzy cognitive maps (FCMs) as a tool for represent-
ing causal relationships between various linguistic concepts. FCMs were used
to model several interesting real world phenomena (see [1] and the references
therein). However, the inferencing process used in FCMs yields a discrete-
time linear system. Such a simple mathematical model is inadequate for
many real world systems.
Mamdani [33] designed a fuzzy algorithm for regulating a laboratory-
built steam engine. The design was based on transforming the linguistic
control protocol, used by a skilled human operator, into a fuzzy controller.
The transformation consisted of the following steps. The verbal informa-
tion was stated as a set of If-Then fuzzy rules. Vague terms such as small
or large 3 were modeled using suitable fuzzy sets. Inferring the fuzzy rule-
base (FRB) led to a well-defined controller. This approach, although rather
ad-hoc, proved to be very successful and led the way to numerous applica-
tions (see, e.g., [12, 54, 55, 64]). However, most of these applications are
based not on modeling natural phenomena, but rather on transforming the
knowledge of a human expert into a fuzzy expert system.
In the last 15 years or so considerable research interest has been devoted
to establishing a solid mathematical foundation for fuzzy logic theory and
fuzzy modeling (see, e.g., [40, 21]). Yet, the success of these more theoretical
approaches in practical applications has yet to be demonstrated.

4 Fuzzy modeling of animal behavior

Recently, FM was applied successfully to transform verbal descriptions of
animal behavior into well-defined mathematical models. The method consists
of four steps: (1) identifying the state-variables; (2) restating the given verbal
descriptions as an FRB relating these variables; (3) defining the fuzzy terms
using suitable membership functions; and (4) inferring the FRB to obtain a
well-defined mathematical model.
This approach was used to derive mathematical models for: (1) the ter-
ritorial behavior of fish [57]; (2) the orientation of a planarian to light [58];
(3) the foraging behavior of ants [49]; and (4) the mechanisms regulating
3
Dvorak and Novak [13] refer to such terms as evaluating linguistic expressions.

6
the population size in flies [47]. In all these examples, the starting point
was a detailed verbal description of the natural phenomenon. Using FM this
was transformed into a well-defined mathematical model. Simulations and
rigorous analysis demonstrated the suitability of the mathematical model.
Independently, Lebar Bajec et al. [2] used fuzzy modeling to develop an
interesting algorithm for simulating the flocking behavior of birds.
We now briefly review two examples of deriving mathematical models for
animal behavior using FM. More details can be found in [57] and [58].

4.1 Territorial behavior in the stickleback

Territory plays a major role in social animal behavior [63] and results in a
rich set of phenomena, but how is territory created? Nobel Laureate Konrad
Lorenz describes a specific example [32, p. 47]:

“. . . a real stickleback fight can be seen only when two males are
kept together in a large tank where they are both building their
nests. The fighting inclinations of a stickleback, at any given
moment, are in direct proportion to his proximity to his nest.
. . . The vanquished fish invariably flees homeward and the victor
. . . chases the other furiously, far into its domain. The farther the
victor goes from home, the more his courage ebbs, while that of
the vanquished rises in proportion. Arrived in the precincts of
his nest, the fugitive gains new strength, turns right about and
dashes with gathering fury at his pursuer . . . ”

Note that Lorenz provided us with a complete verbal description of the

mechanisms underlying the dynamics. Furthermore, he also described the
resulting global patterns as observed in nature:

“The pursuit is repeated a few times in alternating directions,

swinging back and forth like a pendulum which at last reaches a
state of equilibrium at a certain point.”

4.1.1 Fuzzy modeling

The state of the system at time t is determined by the location and fighting
inclination of the fish. For fish i, let xi (t) ∈ Rn , wi (t) ∈ R, and ci ∈ Rn denote

7
the location, fighting inclination, and the location of the nest, respectively.
Here n is the dimension of the state-space.
Lorenz’ description of the change in fighting inclination is transformed
into the following fuzzy rules:

• if neari (xi , ci ) then ẇi = p

• if f ari (xi , ci ) then ẇi = −p,

where p > 0, that is, the fighting inclination increases (decreases) when the
fish is near (far) its nest. Similarly, the description of the movement of fish i
is transformed into:

• if near i (xi , xj ) and highi (wi ) then ẋi = xj − xi

• if near i (xi , xj ) and low i (wi ) then ẋi = ci − xi

where xj is the location of the other fish. That is, when the other fish is
near, and the fighting inclination is high (low), then move in the direction of
the other fish (nest).
Membership functions are used to model the fuzzy terms.4 The term
neari (x, y) is modeled using ni (x, y) = exp(−||x − y||2 /(ki2 )), with ki > 0.
Note that ni (x, y) ≤ 1 with equality only when y = x. The term highi
is defined using hi (w) = (1 + tanh(w/ai ))/2, where ai > 0 determines the
slope of hi . Note that lim hi (w) = 0 and lim hi (w) = 1. The opposite
w→−∞ w→+∞
terms f ari and lowi are defined using fi (x, y) = 1 − ni (x, y), and li (w) =
1 − hi (w), respectively.
Inferring the rules, using multiplication for the “and” operator, and center
of gravity defuzzification, yields:

ẇi = 2p exp(−||xi − ci ||2 /(ki2 )) − p,

ẋi = ci − xi + hi (wi )(xj − ci ), i = 1, 2. (1)

Summarizing, the FM approach transformed the verbal description into

the well-defined mathematical model (1).

4
For a discussion on the degrees of freedom in the FM approach, see Section 6 below.

8
 1

0.8

0.6

0.4

0.2

−0.2

−0.4

−0.6

−0.8

−1
0 5 10 15 20 25
t

Figure 1: Trajectories x1 (t) (solid line) and x2 (t) (dashed line) for initial
positions x1 (0) = −0.4 and x2 (0) = 0.8. The nests are located at c1 = −1
and c2 = 1.

4.1.2 Simulations
Fig. 1 depicts the results of simulations of (1) for the one-dimensional case
(n = 1), with: c1 = −1, c2 = 1, p = a1 = a2 = k1 = k2 = 1, and the
initial conditions: x1 (0) = −0.4, x2 (0) = 0.8, w1 (0) = w2 (0) = 1. It may be
seen that the fish follow an oscillatory movement, with one fish advancing,
the other retreating until a point is reached where they switch roles. Finally,
they converge to a steady state point at x1 = −0.1674 and x2 = −x1 . One
may then say that the territory of fish i is the region between its nest ci
and the point xi . Note that the model reproduces the oscillatory movement
described by Lorenz.
Once the verbal description is converted into a mathematical model, it is
of course also possible to rigorously analyze this model, use it for predictions
of new scenarios, etc. The interested reader is referred to [57] for more details
on this specific model.

4.2 Orientation to light in the Dendrocoleum lacteum

An animals life depends on oriented movements. Such movements guide
the animal into its normal habitat or into other situations which are of im-
portance to it. Various stimuli, such as light or smell, activate the living

9
mechanisms and lead to orientation.
Taxis is a mechanism of orientation by means of which an animal moves
in a direction related to a source of stimulation [19]. For example, positive
(negative) photo-taxis is the directed movement towards (away from) a source
of light. Taxes require sensory organs that can accurately detect the direction
of the stimulus, and a brain sophisticated enough to process the sensory data
and consequently determine the appropriate direction of movement.
Simple organisms, such as wood lice or flat-worms, do not necessarily
have the physiological equipment needed to perform taxes. Their eyes, for
example, do little more than indicate the general intensity of light, but not
its direction. In such organisms the locomotory action can be affected by
the intensity, but not the direction, of the stimulus. This type of response is
referred to as kinesis.5
Klino-kinesis6 is defined as a movement where the rate of turning, but
not the direction of turning, depends on the intensity of the stimulus. This
type of movement appears in many flat-worms: in regions with higher light
intensity their rate of turning increases. As a result, the animals eventually
aggregate in shadier parts of the available habitat.
In a classic paper, Philip Ullyott studied this type of behavior in the
planarian Dendrocoleum lacteum [59]. In order to determine whether the
reaction is simply to the intensity of the light falling on the animal, or to
its direction as well, he designed an apparatus insuring that the only object
visible to the animal is the single patch of light directly above it. Ullyott
analyzed the animals behavior, and found that the stimulus did not affect the
animals linear velocity. Instead, increased light intensity yielded an increase
in the rate of change of direction (RCD) (measured in angular degrees per
minute) in which the animal moved. Ullyott defined the RCD as the sum
of all the deviations in the animals path during one time unit, summing up
both right-hand and left-hand deflections as positive changes. As the light
was switched on, the RCD immediately increased but with time it fell off,
converging to a constant level, which Ullyott designated the basal RCD. This
decrease of response under constant stimulation suggested the existence of
an adaptation mechanism. Ullyott [59, p. 274] summarized his findings as
follows:
“(1) An increase in stimulating intensity produces an increase in RCD.

5
from the Greek kinesis, movement.
6
from the Greek klino, incline.

10
 y
A B
E

dim bright
light light

D C

Figure 2: Path of the “average” animal. Adapted from [17, p. 49].

(2) This initial increase in RCD falls off under constant stimulation owing to
adaptation.
(3) There is a basal RCD, which is an expression of the fact that turning
movements occur even in absolute darkness or at complete adaptation.”
Fraenkel and Gunn [17, Ch. V] reviewed and refined Ullyott’s work. They
developed a simplified and deterministic model for the “averaged” animals
movement. The most important simplification is the assumption that the
animal always turns to the right and always through exactly 90◦ . As the
RCD increases, the time between these right-hand turns decreases.
Following Ullyott, Fraenkel and Gunn furnished a heuristic explanation of
how this behavior drives the animal to the darker regions. Suppose that the
animal is placed in a plane (described by two coordinates (x, y)), where the
light intensity gradually increases along the positive direction of the x-axis
(see Fig. 2). Beginning at a point A, (and assuming that the animal is fully
adapted to the light at A), the animal continues in the positive x direction
until making a right-hand turn at point B, and so on. Along the segment AB,
the light intensity increases and the adaptation level lags behind, so the
RCD increases. Along the BC segment the light intensity is constant and
the RCD decreases back to its basal level. Along the CD segment the light
intensity decreases and the RCD remains constant (note that the RCD is
affected only by an increase in the light intensity). Finally, the behavior
along the DE segment is as in the BC segment. Since the RCD increases

11
along the AB segment and, due to the adaptation process, decreases along
the other segments (until it converges to the basal RCD), the segment AB
will be shorter than the segment CD. Hence, after a set of consecutive turns
the animal ends up at a point E, which might be closer to the darker part
of the plane than the initial point A. Note that the adaptation process plays
an essential role in this explanation, since it allows the animal to (indirectly)
compare a certain light intensity in the present with a previous light intensity.
Ullyott’s results were quite surprising at the time of their publication, as
he postulated that although the direction of the animals movement is not
oriented to the direction of the light at all, it still yields a result similar to
negative photo-taxis, (i.e., the animal eventually finds its way to the shadier
regions of its habitat): “This alternate stimulation and adaptation has an
effect on the RCD of such a kind that the animal is led automatically to the
place of minimal intensity.” [59, p. 277]
Both Ullyott and Fraenkel and Gunn provided only a verbal description
of the animals behavior and its outcome. Patlak [43] studied the behavior
of random particles affected by a force field. He derived a suitable Fokker-
Planck-type equation characterizing the particles movement, and used this
stochastic model to analyze klino-kinesis [42]. However, his model ignores
the adaptation process which plays a vital role in the description given by
Ullyott and Fraenkel and Gunn.

4.2.1 Fuzzy modeling

The verbal description given above can be transformed into a mathematical
model using the FM approach. The state-variables are: the animals location
in the plane (x(t), y(t)); the light intensity at each point in the plane l(x, y);
the light intensity that the animal is currently adapted to la (t); the RCD r(t);
the direction of movement θ(t); and the set of turning times: t1 , t2 , . . . . The
model also includes two constants: the animals linear velocity v, and the
basal RCD rb .
The verbal description is restated as an FRB relating the state-variables.
The adaptation process changes the level of adaptation to light in accordance
with intensity of light. We state this as two fuzzy rules:

• If l − la is positive, then l˙a = c1

• If l − la is negative, then l˙a = −c1

12
where c1 > 0. The RCD decreases when it is above the basal RCD, and
increases when the light stimulus is above the adaptation level. We state
this as:
• If r − rb is large, then ṙ = −c2
• If l − la is high, then ṙ = c3
where c2 , c3 > 0. Finally, following Fraenkel and Gunn’s model, we add the
crisp rule:
Rt
• If tl r(τ )dτ = q, then θ ← θ − π/2
Here, tl < t denotes the time when the last turn took place, and q > 0. In
other words, the RCD is constantly accumulated and whenever it reaches the
threshold q, the animal makes a right-hand turn.
The next step is to define the fuzzy membership functions. For our first
set of rules, we use µpositive (x) = ek1 x /(ek1 x + e−k1 x ), with k1 > 0, and
µnegative (x) = 1 − µpositive (x).
Let Sk be the piecewise linear function defined by: Sk (z) = 0 for z ≤ 0;
Sk (z) = z/k for 0 < z < k, and Sk (z) = 1 for z ≥ k. For the second set of
rules, we use µlarge (x) = Sk2 (x), µhigh (x) = Sk3 (x), with k2 , k3 > 0.
Using additive inferencing (see, e.g., [5]), the first set of rules yields
l˙a = c1 µpositive (l − la ) − c1 µnegative (l − la )
= c1 tanh(k1 (l − la )). (2)
Similarly, the second set of rules yields ṙ = −c2 Sk2 (r − rb ) + c3 Sk3 (l − la ). The
actual movement of the animal is given by ẋ = v cos(θ), and ẏ = v sin(θ).
The crisp rule
R ti+1implies that the value θ “jumps” at the discrete times t1 , t2 , . . .
satisfying ti r(τ )dτ = q, i = 0, 1, . . . (with t0 = 0). Thus, the model is
a hybrid system [60] combining continuous-time dynamics and discrete-time
events.
Fig. 3 summarizes the mathematical model. The upper arrow in this
figure corresponds to the conditional transition described by the crisp rule.
When the condition holds, θ is updated, and then the evolution in time
proceeds using the continuous-time dynamics.
The FM approach allowed us to transform the verbal description of the
behavior into a mathematical model. One criteria for evaluating the suit-
ability of this model is how well it mimics the patterns that were actually
observed in the natural system.

13
 Rt
ti r(τ )dτ = q
l˙a = c1 tanh(k1 (l − la )) θ ← θ − π/2
ṙ = −c2 Sk2 (r − rb ) + c3 Sk3 (l − la ) i←i+1

ẋ = v cos(θ) ẏ = v sin(θ) ti ← t

Figure 3: The hybrid model.

4.2.2 Simulations
In one of his experiments, Ullyott [59] placed the animals in total darkness
for three hours (to make them completely dark adapted). He then exposed
the animals to a sudden increase in light intensity, and measured the RCD
at different times. Fig. 4 depicts his results.
We simulated a similar scenario in our mathematical model with

c1 = 5, c2 = 1, c3 = 2, k1 = k2 = 1, k3 = 2, r(0) = rb = 2, la (0) = 0, (3)

and light intensity: l(t) = 0 for t ∈ [0, 1), and l(t) = 1 for t ≥ 1. Fig. 5
depicts la (t) and r(t) as a function of time. It may be seen that the adaptation
level increases once the light is switched on, and then converges to la (t) =
1 (= l(t)). Note the qualitative resemblance between the behavior of r(t)
and the RCD as actually measured by Ullyott (Fig. 4).

4.2.3 Analysis
One of the principle advantages of mathematical models is that they are
amenable to mathematical analysis. The next result analyzes the models
behavior for a step function in the light intensity.

Proposition 1 [58] Consider the hybrid model with initial conditions la (0) =
0, r(0) = r0 , and light intensity l(t) = 0 for t < 0, and l(t) = 1 for t ≥ 0. De-
note w := c1 k1 k2 − c2 , p1 := sinh(k1 ), and assume that the models parameters
satisfy:
k3 ≥ 1, w > 0, and k2 ≥ r0 − rb + (p1 k2 c3 )/(k1 k3 w). (4)
Then
la (t) = 1 − sinh-1 (p1 exp(−c1 k1 t))/k1 , (5)

14
 800

600

RCD (angular degrees per minute) 400

200

0
0 10 20 30 40 50
Time (minutes)
A B C

Figure 4: The relationship between RCD and duration of stimulus. AB,

RCD of the animal in darkness (basal RCD). At B a light intensity of 2500
ergs/(cm2 ·sec) was switched on. BC, adaptation to the stimulus. Each point
on the curve represents the average of fourteen experiments. (Reproduced
from [59] with permission.)

where sinh-1 denotes the inverse hyperbolic sine function, and

Z t
c2 c3 c2
r(t) = rb +exp(− t)(r0 −rb )+ exp( (τ −t)) sinh-1 (p1 exp(−c1 k1 τ ))dτ.
k2 k1 k3 0 k2
(6)
It is easy to verify that the specific parameter values used in the simula-
tion (3) indeed satisfy (4). Hence, (5) and (6) actually provide an analytical
description of the responses depicted in Fig. 5.
It is possible to show that (6) implies that r(t) decays exponentially. This
agrees with the observations of Ullyott, who states: “. . . it is possible to see
that the falling off of RCD with time is exponential...” [59, p. 270].
Extensive literature exists on developing mathematical models in biology
(see, e.g., [37, 38, 14]), using classic modeling techniques that have nothing

15
1.4 2.15

1.2

2.1
1

0.8

2.05

0.6

0.4
2

0.2

0 1.95
0 0.5 1 1.5 2 2.5 3 3.5 4 4.5 5 0 1 2 3 4 5 6 7 8
t t

Figure 5: la (t) (left) and r(t) (right) as a function of time. The light is
switched on at t = 1.

to do with FM. A natural question is: why use FM at all? We try to address
this question by describing some of the advantages of FM.

5 Advantages of the fuzzy modeling approach

The most important advantages of the FM approach stem from the close con-
nection between the initial verbal description and the resulting mathematical
model. In the FM approach the knowledge about the system is represented in
three different forms in parallel: (1) the initial verbal description and expla-
nation; (2) the FRB; and (3) the mathematical model obtained by inferring
the rules. These three representations provide a synergistic overview of the
system that may provide new insights on the studied phenomenon.
Fig. 6 summarizes the design cycle in the FM approach. The initial verbal
description is stated as an FRB. Defining the membership functions, and
inferencing yields the mathematical model. Simulations and analysis can be
used to check whether the models behavior is congruent with that actually
observed in nature. When this is not the case, it is sometimes possible,
due to the If-Then structure of the rules, to determine which fuzzy rule

16
 fuzzy
verbal modeling fuzzy inferencing mathematical
description rule base model

modification simulation and analysis

Figure 6: Synergy between three different forms of knowledge representation.

should be changed and how. Inferring the modified FRB yields a modified
mathematical model, and so on. Furthermore, any change in the FRB can
also be interpreted as a change in the initial verbal description, suggesting
directions for further research of the original natural phenomenon. In this
sense, the behavior of the mathematical model can be used, to some extent,
to prove or refute the modeler’s ideas as to how the natural system behaves.
We now demonstrate these issues using the mathematical models described
in Section 4.

5.1 Interpretability
A fuzzy model represents the real system in a form that corresponds closely
to the way humans perceive it. Thus, the model is understandable, even by
non-professionals, and each parameter has a readily perceivable meaning [20].
The next example demonstrates this.

Example 1 Consider the parameters k1 and k2 in the mathematical model (1).

To gain some intuitive understanding of these parameters, recall that the ki s
originate from the membership function ni (x, y) = exp(−||x−y||2 /(ki2 )) used
to define the term neari (x, y). As ki decreases, the Gaussian becomes more
peaked, so fish i will sense the other fish as “near” only when it is indeed
very close to it. In other words, a decrease in ki corresponds, in some sense,
to making fish i “less aggressive.” This provides a clear interpretation of ki
by relating it to the original verbal description.
This interpretation can be verified by analyzing (1). Assume, without

17
loss of generality, that c2 = c and c1 = −c, for some c > 0. In this case,
√ √
x1 = −c + k1 ln 2, x2 = c − k2 ln 2,
x1 + c x2 − c
w1 = a1 tanh (2 2
−1
− 1), w2 = a2 tanh (2 1
−1
− 1),
x +c x −c

where tanh−1 denotes the inverse hyperbolic tangent function, is a (locally

asymptotically stable) equilibrium point of (1). In other words, √ there exist
initial conditions for which fish i will come to rest at a distance ki ln 2 from
its nest, i = 1, 2.
If k1 > k2 then the equilibrium position is no longer symmetric with
respect to the nests and, eventually, fish 1 will guard a larger territory than
fish 2. This leads to a clear link between the relative aggressiveness of the
fish, as manifested by the values of the parameters, and the resulting sizes
of their territories. 2

5.2 Verifying the verbal description

Once the mathematical model is derived, its suitability can be examined
using both simulations and rigorous analysis. The close connection between
this mathematical model and the original verbal description can be used, to
some extent, to verify the correctness of the verbal description. The next
examples demonstrate this.

Example 2 The hybrid model for the planarian’s orientation to light in-
cludes the fuzzy rule: “If l − la is high, then ṙ = c3 .” If c3 = 0, then the
RCD will not increase when l − la is high. Since the increase in RCD plays a
crucial role in Ullyott’s explanation, we may expect that the models behavior
will change substantially.
To verify this, we substitute c3 = 0 in the mathematical model. Then

ṙ(t) = −c2 Sk2 (r − rb ),

and assuming that r(0) = rb yields r(t) = rb for all t ≥ 0. Recall that the
right-hand turns take place at times t1 , t2 , . . . such that
Z ti+1
r(τ )dτ = q,
ti

18
so ti+1 − ti = q/rb for all i. In other words, the right-hand turns take
place at regular intervals. Since the linear velocity v is constant, it is clear
that x(ti ) = x(ti+4 ) and y(ti ) = y(ti+4 ) for all i. In other words, the planarian
will periodically return to its initial position, and the process of gradually
moving toward the shadier parts of the habitat will not take place. 2

Example 1 (continued) Consider again the stickleback model. A closer

look shows that there is actually a discrepancy between Lorenz’ verbal de-
scription and the FRB. Indeed, Lorenz claimed that: “The fighting inclina-
tions of a stickleback, at any given moment, are in direct proportion to his
proximity to his nest,” whereas the FRB relates the proximity to the nest
with ẇi , that is, the derivative of the fighting inclination.
There is a good reason for this discrepancy. Initially, we actually did use
fuzzy rules with wi , and not ẇi , in the Then-part (e.g., if neari (xi , ci ) then
wi = p). Simulating the resulting mathematical model showed that the fish
move toward each other and then stop at an equilibrium point. There were
no periodic motions. This does not agree, of course, with the actual behavior
observed in nature.
A moment of reflection suggested that the problem was that the fish
have no “memory,” so they stop once they reach the edge of their territory.
To correct this, we changed the Then-part of the rules using ẇi instead
of wi . Then, when a fish approaches the border of its territory, the fighting
inclination starts decreasing, but it may still be sufficient to lead it across to
the other territory, and thus initiate the oscillatory behavior.
The change in the FRB can be naturally interpreted as a change in the
initial verbal description. The modified FRB actually corresponds to the
modified statement: The change in the fighting inclinations of a stickleback,
at any given moment, are in direct proportion to its proximity to its nest.
This is a new assertion that can be checked only by means of suitable ex-
periments on the real world system. Nevertheless, this demonstrates the
synergistic view provided by the interconnection of the verbal description,
the FRB, and the mathematical model. 2

Example 2 (continued) Consider again the hybrid model. Fig. 7 depicts

a simulation of this model for initial conditions: r(0) = rb , x(0) = y(0) =
la (0) = 0, and l(x, y) = x, that is, the light intensity increases linearly along
the x-axis.
Fig. 7 depicts the trajectory (x(t), y(t)). It may be seen that x(t0 ) >

19
 1

−1

y −2

−3

−4

−5
−2 −1 0 1 2 3 4 5
x

Figure 7: Trajectory (x(t), y(t)) for initial values x(0) = y(0) = 0, and light
intensity l(x, y) = x.

x(t4 ) > x(t8 ) > . . . , where ti are the turning times (with t0 = 0). Thus, the
trajectory indeed moves toward the darker region of the plane. This fact is in
agreement with Ullyott’s observations [59, p. 272]: “The shift of the position
of the animal towards the end of the gradient was rather a gradual process,
but in each case with the steep gradient, the animal was to be found at the
darker end within 2 hours from the beginning of the experiment.”
However, it may also be seen that y(t0 ) < y(t4 ) < y(t8 ) < . . . . This
feature is not described in Fraenkel and Gunn’s explanations (see Fig. 2).
A moments reflection suggests that this behavior is indeed what we should
expect. Indeed, reconsidering the discussion preceding Fig. 2 suggests that,
due to the adaptation process, the segment DE should be longer than the
segment BC. Although this is evident in retrospect, it is not necessarily
immediate when reading the verbal explanation. The simulations of the
corresponding mathematical model, derived based on this explanation, is
what makes this aspect of the verbal explanation apparent. 2
The examples above describe the case where the verbal description was
transformed into a mathematical model using FM. In the specific approach

20
used here, the first two stages in this transformation are [57]: (1) identify
the variables needed to define the systems state; and (2) re-state the given
verbal information as an FRB relating these variables.
It has been the author’s experience, in many examples, that by following
these stages, it is possible to detect deficiencies in the given verbal descrip-
tion. For example, it is sometimes possible to see that although some state-
variable, say x(t), is needed, the verbal description does not include sufficient
information on the relation of x(t) to the other variables.
The interesting thing is that detecting such a problem in the verbal de-
scription is not necessarily immediate. It is revealed only when one tries to
follow the FM approach. This is similar to discovering that an algorithm,
stated in pseudo-code, is not really complete while actually trying to encode
it in some computer language.
Scientists who provide verbal descriptions for various phenomena may
choose to use FM not only to derive a mathematical model, but simply
to verify that there are no unapparent missing parts or gaps in their ex-
planations. Verifying the verbal description using this approach forces the
scientist to provide a mechanistic and complete description of the relevant
phenomenon. This is congruent with the methodological aspect of logic as
a theory for analyzing the consistency and completeness of knowledge-based
systems [45].
The discussion will not be complete without describing some of the down-
sides of the FM approach.

6 Disadvantages of the fuzzy modeling ap-

proach
In the FM approach the fuzzy rules follow more or less directly from the
verbal description. However, there are many degrees of freedom, and a great
deal of subjectivity, in specifying the other components of the model: the
membership functions, logical operators, and inferencing method. A change
in any of these may yield a mathematical model with a rather different be-
havior. This is usually regarded as a disadvantage, as it suggests that we can
easily miss the “correct” mathematical model.
It should be noted however that this “fuzzines” is not really a feature of
the FM approach, rather it is a direct consequence of the inherent vague-

21
ness of verbal descriptions. Indeed, imagine a scenario in which two applied
mathematicians are given the same verbal description, and asked to develop
a suitable mathematical model for it. It seems reasonable to expect that the
two may end up with different models.
The conclusion is that one should not be tempted to believe that there
exists a single “correct” mathematical model. Any reasonable approach for
transforming words into equations must contain some degrees of freedom.
These should be addressed by a trial and error approach, hopefully converging
to a useful model.
The inherent vagueness of natural language is not a weakness, rather it
is this vagueness that allows us to communicate valuable information parsi-
moniously. This is congruent with Zadeh’s principle of incompatibility [67]:
“As the complexity of a system increases, our ability to make precise and
yet significant statements about its behavior diminishes until a threshold is
reached beyond which precision and significance (or relevance) become al-
most mutually exclusive characteristics.”
Note also that in the specific context of biomimcry, the goal is not neces-
sarily to obtain a mathematical model that reproduces the relevant natural
behavior precisely. Rather, the goal is to design an artificial system that is
inspired by the natural behavior and solves a given engineering problem.
In the examples described above the degrees of freedom in the FM ap-
proach were determined in attempt to reach a model that satisfies two re-
quirements. The first was, of course, that the model reproduces the described
behavior. The second requirement was that it be amenable to mathematical
analysis.
Extensive experience and information have accumulated over decades of
successful implementations of the FM approach. Consequently, there is con-
siderable literature on how the various elements in the fuzzy model influence
its behavior (see, e.g., [53, 20]). If the verbal description of the system is
accompanied by quantitative data, then it is possible to apply well-known
methods, such as fuzzy clustering, neural learning, and least squares approx-
imation [20, 7, 25], for learning the parameters of the fuzzy model.

7 Fuzzy modeling and biomimcry

FM may be suitable for addressing biomimicry in a systematic manner.
Namely, start with a verbal description of an animals behavior (e.g., for-

22
aging in ants), and apply FM to obtain a mathematical model of this behav-
ior that can be implemented by artificial systems (e.g., autonomous robots).
Although it is a natural idea, it seems that the computational intelligence
community is not aware of this possible application of FM in biomimcry.
It should be noted that in addition to the general advantages of FM
described above, there are at least two considerations that make FM par-
ticularly suitable for modeling biological behavior and, in particular, animal
behavior.
First, many animal (and human) actions are non-binary, that is, the
behavior itself is “fuzzy.” For example, the response to a (low intensity)
stimulus might be what Oskar Heinroth [23] called intention movements, that
is, a slight indication of what the animal is intending to do. Tinbergen [56,
Ch. IV] states: “As a rule, no sharp distinction is possible between intention
movements and more complete responses; they form a continuum.” It is
interesting to recall that Zadeh [66] defined a fuzzy set as “a class of objects
with a continuum of grades of membership.” Hence, FM seems the most
appropriate tool for studying such behaviors.
The second consideration is that studies of animal behavior often pro-
vide a verbal and, therefore, an inherently vague, description of both field
observations and interpretations. Darwin [9, Ch. 2] states: “Nor shall I here
discuss the various definitions which have been given of the term species. No
one definition has as yet satisfied all naturalists; yet every naturalist knows
vaguely what he means when he speaks of a species.” As another example,
Fraenkel and Gunn [17, p. 23] describe the behavior of a cockroach, that
tends to become stationary when its body surface is in contact with a solid
object, as: “A high degree of contact causes low activity . . . .” FM is a
suitable approach for addressing this vagueness. Indeed, note that the last
statement can be immediately stated in the form of a fuzzy If-Then rule: “If
degree of contact is high then activity is low.”

8 Concluding Remarks
Recently, the design of artificial algorithms and machines that imitate bi-
ological behavior, is attracting considerable interest. In some cases, there
exist verbal descriptions of the natural behavior we aim to mimic. The prob-
lem of mimicking the natural behavior is then reduced to transforming the
given verbal information into a well-defined mathematical formula or algo-

23
rithm that can be implemented in an artificial system. A natural approach
for addressing this is FM.
In this paper, we tried to summarize some of the lessons learned from
applying FM to develop mathematical models of animal behavior, and to
suggest the next natural step, that is, the use of FM in biomimcry. Special
attention was devoted to presenting the potential advantages of FM in this
specific context.
We believe that FM can and should be utilized as one of the main tools
in the field of biomimicry. We hope that this paper will stimulate more work
in this direction.

Acknowledgments
The author is grateful to the anonymous reviewers and the editor for their
detailed and constructive comments. The author thanks The Company of
Biologists Ltd. for permission to reproduce one of the figures from [59].

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