Anda di halaman 1dari 13

The Semantic Morphology of Adolf Portmann:

A Starting Point for the Biosemiotics of Organic Form?


Karel Kleisner
Received: 23 March 2008 / Accepted: 16 April 2008 /
Published online: 16 July 2008
#
Springer Science + Business Media B.V. 2008
Abstract This paper develops the ideas of the Swiss zoologist Adolf Portmann
or, more precisely, his concept of organic self-representation, wherein Portmann
considered the outer surface of living organisms as a specific organ that serves
in a self-representational role. This idea is taken as a starting point from which
to elaborate Portmans ideas, so as to make them compatible with the theoretical
framework of biosemiotics. Today, despite the many theories that help us understand
aposematism, camouflage, deception and other phenomena related to the category of
mimicry, there still is a need for a general theory of self-representation that would
re-synthesize evolutionary, morphogenetic and semiotic aspects of the surface of
organisms. Here, Adolf Portmanns concept of self-representation is considered as
an important step towards the biosemiotics of animal form.
Keywords Semantic organs
.
Adolf Portmann
.
Homosemiosis
.
Self-representation
.
Mimicry
.
Imitation
.
Display
.
Homology
.
Organic form
Introduction
Adolf Portmann developed an original approach to the phenomenon of life, with
special emphasis on its representational aspects (Portmann 1960a, 1960b, 1969).
This paper re-introduces and elaborates his basic ideas, in the belief that they can
be a source of inspiration for modern biosemiotics. The evolution of particular
constituents of the body may be driven, not only by selective pressures that increase
their functional utility, but also by their ability to interact with the umwelten of other
living beings in a meaningful and contextual way. This is to say that without these
interactions, the functioning of any sexual display, mimicry or deceptive behavior
can hardly be understood. Organs of display (i.e., semantic organs) act through the
meaning that they acquire during umwelt-specific interpretations.
Biosemiotics (2008) 1:207219
DOI 10.1007/s12304-008-9014-4
K. Kleisner (*)
Department of Philosophy and History of Science, Charles University, Vinin 7, 128 44 Prague,
Czech Republic
e-mail: kleisner@seznam.cz
Portmanns investigations were strongly influenced by the theoretical biology and
Umweltforschung of Jakob von Uexkll (1921, 1928), as well as by his comparative
studies on morphology and behavior. In practice, Portmanns contribution was
overshadowed by the success of ethology, on the one hand, and by the general
enthusiasm for genetic and molecular methods on the other. The increasing popularity
of genetic research was accompanied by a devaluation of morphology, which lost
its status as a realm from which scientific explanations could be derived. Conversely,
the activity of genes is at least partially assumed to explain morphology. In such an
intellectual climate, Adolf Portmann was a proponent of an old-fashioned tradition
of continental biology, and while he may look rather anachronistic in his strong
focus on the realm of the visible, he frequently criticized the prevailing research
of his period for its orientation on the realm of the invisible, and its neglect of
lifes most spectacular aspects (Portmann 1960a, 1965). Biosemiotics now brings a
theoretical account of life that seems sufficiently open to embrace the ideas of
Adolf Portmann.
Explaining Organic Form?
Organic form can be explained by two kinds of causes: extrinsic and intrinsic.
Reasoning from extrinsic causation emphasizes the influence of various forms of
selective pressures caused either by other organisms or by environmental conditions.
This way of reasoning is entirely at the service of survival. Every part of an
organism must be formed in such a way as to fulfill its life-sustaining function; from
this it follows that reproduction is the sole and ultimate purpose of all formal-
structural representations of life. Consequently, reproduction is the sole referential
frame that is adequate to explain all manifestations of life.
On the other hand, reasoning from intrinsic causation accents the importance of
the inner potentialities of organisms that generate a particular form. In modern
biology, the activity of developmental genes is usually assumed to underlie the
emergence of particular morphologies. Thanks to the success of bottomup
methodologies in biotechnology, molecular and developmental genetics etc., biologists
ascribe to genes the meaning of causal function, and morphology has acquired the
practical role of testing the causal functions of the genes. Consider, for instance, the
intertwined studies on gene activation/inactivation, the mutation and the regulation of
gene expression via the monitoring of changes in morphology. On second thought, we
knowthat the results obtained at the genetic level are often used to explain the evolution
of forms (morphologies). However, do these attempts really seek to explain the
evolution of organic form? Or is it rather the logic of the interactions between
constituents of different levels that is revealed? Is it ever possible to understand the
transformations of constituents at different higher levels via functional causations
between constituents of the lower level?
It is well known that the mapping of genes onto morphologies, and vice versa,
is not an easy task: the inherent problem stems from the so-called evolutionary
dissociations, that is, from incongruent relationships among various levels of biological
organization. In principle, two kinds of dissociations are imaginable between genes and
morphologies: (1) homologous genes are involved in the formation of nonhomologous
208 K. Kleisner
structures, and (2) nonhomologous genes are recruited to serve the formation of
homologous morphologies (Wray and Abouheif 1998).
The results obtained at the genetic level are useful for studying the relationships
between genes and morphologies, but what is actually revealed here is the complexity
of those relationships, and the obtained results provide a rather restricted account of
the real transformations of organic form. This is so because behavior at the form-
structure level somehow transcends the functional causation of constituents at any
underlying level. And so we address the morphological aspects of an organism neither
from a strictly intrinsic nor from a purely extrinsic perspective, but attempt to explain
some of their evolutionary transformations via the potential of a morphology that
exceeds its functional necessity. The form (Gestalt) of organisms should be studied for
its own sake (Portmann 1990: 138). In this framework, the formal aspects of living
beings, such as shape or surface patterns, coloration or ornamentation, should be
approached neither as a useful control in genetic experiments, nor as a possible
source of data for phylogenetic inference.
The uniqueness of Adolf Portmanns approach may be found in his explanation
that organic form is in itself something valuable. Portmann acknowledged that the
external surface of an organism has its own formal value and a certain kind of
autonomy over other life-sustaining functions. He was convinced that this outermost
aspect of an organism opens a way to the innermost dimensions, because surface
manifestation reflects inner self-experience and the very selfhood of every organic
being, thus bringing us closer to understanding the existence of living creatures
(Portmann 1969: 315). For these reasons he developed a terminology, or rather a
conceptual framework, specific to such a purpose.
Portmanns Conceptual Framework and Self-Representation of Organisms
Portmans central idea can be formulated as follows: the outermost surface of an
organism is a manifestation of phenomena proper that arises within the process of self-
representation of organic inwardness. Phenomena proper (eigentliche Erscheinungen)
represent the entire outer aspect of an organism that may potentially stimulate the
senses of another living being, that is, those aspects of the organism which are exposed
to the perception of another subject in a natural, non-invasive, way. For example, the
various colorations, ornaments, coat patterns and other morphological features of the
outermost organismal surface are considered to be manifestations of phenomena
proper within the realm of sight. Although Portmann focused primarily on vision,
phenomena proper comprise many other expressions of life (i.e., behavioral, olfactory,
acoustic, tactile etc.). Nevertheless, there are also other dimensions of an organism
which are naturally concealed from the senses. All organs and organ systems that are
not immediately apparent from the outsidetypically inner structures that ensure the
energetic and the mechanical functioning of the organic wholebelong to this
category and were marked by Portmann as phenomena improper (uneigentliche
Erscheinungen). In contrast to phenomena proper, improper phenomena are not
primarily intended to arouse the sensation of another organic subject. All phenomena
proper serve to present the self; rising up from the inwardness of a living being, which
can be understood as a particular form of organic self-experience. In contrast,
The semantic morphology of Adolf Portmann 209
phenomena improper do not normally attain any self-representational role (Portmann
1960a: 102f., 1960b, 1965, 1969).
Portmann distinguished between two types of phenomena proper: addressed and
non-addressed (adressierten und unadressierten Erscheinungen; Portmann 1960a:
109f., 1965). Addressed phenomena represent a largely unproblematic category that
is frequently well explained within contemporary biological theory. Addressed
phenomena function as signals assigned to a particular receiver and thus partake in
the survival of the bearer. Cases such as warning coloration, camouflage, sexual
display etc. may serve as good examples. Non-addressed phenomena, on the other
hand, do not serve any practical purpose; they do not play any functional role related
to survival. Yet, non-addressed manifestations represent indispensable features that
characterize the members of a particular evolutionary lineage. Moreover, it was non-
addressed phenomena that led transcendental morphologists to the idea of the type.
The very existence of non-addressed phenomena proper stands for evidence that the
richness of life-expressions cannot be resolved solely by functional explanations in
the context of self-preservation and reproduction of organic beings. Non-addressed
displays represent those phenomena of life, those peculiarities, which are not
conditioned by any functional necessity. This non-addressed domain of all life may
thus be meaningfully embodied in the concept of self-representation.
At first sight, Portmans conceptual framework may be regarded as rather
unusual, even somewhat extraordinary, but still it remains within common biological
practice. Although it emphasizes aspects of living beings that cannot be fully
explained in terms of function, selection and survival, this approach remains to some
extent compatible with the traditional way of functionalist and technologist
reasoning, which does not necessarily stand in contradiction to the idea of organic
self-representation.
This may be demonstrated by the corporeal transparency of some sea creatures
such as, for example, salps, some species of fish, or some nudibranchiate mollusks.
Using such examples, Portmann demonstrates the limits of a technical explanation of
animal form. Normally, in a majority of animal forms, the asymmetry of the inner
organs is hidden beneath an opaque surface that is symmetrically arranged.
However, we can observe a peculiar solution to inner asymmetry in transparent
and semi-transparent living forms, wherein the symmetry is preserved by centering
all asymmetrical inner organ complexes into a single opaque visceral cluster
(nucleus vegetativus; Portmann 1960a: 104). Inner structures that are not centered
into an opaque visceral cluster are usually arranged symmetrically along the main
body axis. It is also worthy of note that a symmetrical arrangement of inner organs
in life forms with jellylike transparent bodiesas is the case of transparent
nudibranchiates from the genus Phyllirrhoe
1
, to use Portmanns examplemay also
1
A description of the species Phyllirrho bucephal from over 150 years ago makes us wonder about the
intriguing architecture of the symmetrical corporeal composition of this animal: The outer integument is
perfectly transparent and lined by muscular bundles, disposed longitudinally, and somewhat more than
their own breadth apart. These communicate with one another by oblique branching lips, which thus form
a kind of network enclosing long lozenge-shaped spaces. Here and there nerve-trunks of considerable size
accompany the longitudinal bundles, dividing off into smaller twigs, which distribute themselves at pretty
equal distances in a direction more or less perpendicular to that of the muscular fibres. (McDonald 1854:
363).
210 K. Kleisner
be explained in technical terms as a stabilizer during swimming or floating in
the water column. Transparency itself can be explained in technical terms, as a
secondary effect of a high water content in gelatinous tissues which ensures
buoyancy of the body in analogy to the swim bladder of fish. Even the concentration
of asymmetrical visceral structures into a single centered cluster in many fish may be
seen as a constructional solution that increases stability and maneuverability of the
body. This is probably true, but it does not tell the whole story. From the perspective
of technology, there is no reason why entrails organized in a visceral cluster should
be divided by an opaque pigment.
Portmann demonstrates that many life forms tend to preserve their optical symmetry
despite the asymmetrical nature of their inner organs, which are, architectonically, not to
be seen. In this respect, the concept of phenomena proper (eigentliche Erscheinungen)
as manifested by formal aspects of an exposed surface, exhibits superiority over purely
functional explanations. Portmann does not reject functional explanations as such;
nevertheless, he shows that function-based reasoning is unable to answer all questions
related to organic form.
One could admit that self-representation of phenomena proper is nothing more
than a kind of function. In other words, the concept of function might also be
extended to the realm of organic self-representation whose particular manifestations
(i.e., phenomena proper) bring some selective advantage to its bearer. However, by
placing phenomena proper into the context of function, only addressed phenomena
proper might be supposed to have a functional role (e.g. to communicate, to warn,
etc.). In accordance with this kind of function, organs of display of an animal are
thought to have a particular evolutionary purpose for which they have been selected
in the course of evolution.
2
In contrast, non-addressed phenomena proper certainly
do not fit this scenario; they are not deliberately applied to the sensory organ of a
particular receiver, so there is no adaptive purpose for which they have evolved.
Within this category one often finds cases in which it is hard to imagine the reasons
why they should be privileged by natural/sexual selection. For instance, why are
some beetles neutrally brownish while others are metallic, or why do some
butterflies expose intricate, bizarre, albeit species-specific, color patterns with no
striking adaptive role? Interestingly, Zdenk Neubauer (personal communication)
considers these taxon-specific appearances as heraldic signs that provide evidence of
ontological authenticity and sovereignty. A coat of arms, or blazon of arms, often
appears depicted on a shield (of a lord), which is a device with a protective function,
but heraldic signs themselves surpass any notion of functional role because they
serve as self-representations of the bearer and his lineage.
One may wonder how these intricate taxon-specific patterns might have evolved,
how they were retained in evolution when they carry no adaptive function.
According to Portmann, these non-addressed phenomena proper are ascribable to
externalization of organic inwardness through the process of self-representation;
these phenomena are always of a taxon-specific character, having their source in the
inner causes of an organism. Even though this internalistic position of Portmanns
would not satisfy most biologists, there still is good reason to acknowledge the idea
2
Such explanation of function is termed selected effect (SE) analysis of function (Amundson and
Lauder 1994).
The semantic morphology of Adolf Portmann 211
of the representation of selfhood of organisms as a peculiar property of living beings,
at least because extending the concept of function does not always solve the problem
of organic form entirely. In principle, and irrespective of whether they are addressed
or non-addressed, all visual phenomena proper presume the existence of a seeing
eye (consider the specifically shaped and patterned surfaces that contradict the
non-aesthetic functionality of inner organs). Again, the term non-addressed
means that these phenomena proper have no particular addressee, but it does not
mean that no other being is there to perceive them as somewhat meaningful. As an
example, imagine the black and yellow coloration of wasps. There probably is no
single and sole addressee of this black and yellow pattern at which it was originally
aimed; instead, many different living beings that act as addressees are afraid of it.
From the functional point of view, it is quite understandable how the wasp-like
pattern was adopted in evolution by different imitators e.g. other hymenoptera,
various hoverflies, long horn beetles etc., but the real problem is to explain how and
why this pattern appeared in wasps themselves. In other words, when we ask as to
why an organism resembles a wasp, we simultaneously pose the question: why is
a wasp wasp-like? Such questions lead us back into the realm of the self-
representational meaning of organisms.
Form, Function and Meaning
In the previous section, we have seen that it is impossible to characterize non-
addressed phenomena, i.e. neutral taxa-specific appearances, in the context of
function. However, does this apply to addressed phenomena as well? One may
rightly admit that the black and yellow color pattern of wasps, and of their imitators,
has, in the course of evolution, acquired a warning function, though its original
purpose, if there was one, could have been quite different. Nevertheless, not even the
effectiveness of a warning coloration can be properly understood within the realm of
function, because the cogency of any display is conditioned by circumstances that
exceed any concept of function. This brings us to the role of meaning. How does the
question of the function of a structure differ from the question of the meaning of the
same structure? For example, if we ask about the function of eyespots on the wings
of a butterfly, we may answer that they serve to intimidate avian predators. This tells
us that the structure under investigation functions as a warning signal with a
protective function for the bearer. Nevertheless, such a question does not draw
attention to the subject affected (in this case intimidated) by the eyespots. Nothing is
told about the receivers perception and interpretation of these structures. In contrast,
if we ask about the meaning of the eyespots, the question is put as to whether these
eyespots are interpreted as real eyes within the umwelt of some animal, or whether
they relate to some unspecified dangerous thing (see abstract mimicry sensu Maran
2007: 239). In other words, the question regarding the function of the eyespots can
be satisfactorily answered by describing how the eyespots affect the survival of the
prey in a preypredator interaction. However, to ask about the meaning of the
eyespots is an attempt to answer the question as to what do these structures signify to
an addressee, what do they represent in the perceptual world (umwelt) of a receiver,
and to what extent is the receivers signification of these structures comparable to
212 K. Kleisner
that of ours. Investigating functionality is the domain of the mechanistic approach
and belongs to the program of adaptationism, whereas asking about meaning falls
within the province of biosemiotics.
Semantic Organs: Concept and Examples
Traditionally, the morphological and anatomical constituents of animal bodies have
been conceptualized in terms of form or function. Differing scholars have valued
either the first or the second as the best explanation of the reasons that shape and
transform organic matter (Russel 1916; Kleisner 2007). Herein, we attempt to
introduce the idea of semantic organs, that is, organs that are defined neither by their
form nor by their function, but by their meaning in the umwelt of an organism, or a
group of organisms, under investigation (cf. Kleisner 2008; Kleisner and Marko
2005). In the following discussion, we focus on optical representations, although the
existence of semantic organs that are based on olfactory or acoustic representations
is clearly imaginable within the same theoretical framework.
This attempt stemmed from Portmanns idea that the opaque surfaces of
organisms represent a new specific kind of organ, and not merely a mechanical
barrier that binds together inner structures and metabolic processes in order to
protect them from external influences. Presumably, the appearance of ornamented
opaque surfaces is associated with the origin of vision. It is highly probable that
prior to the emergence of vision, most organisms had a semitransparent surface, not
dissimilar from the milky whitish semitransparent coloring of vertebrate embryos. A
certain comeback of such neutral and indistinct surface colorations is apparent in
organisms dwelling in a dark environment, especially in parasitic forms or in
inhabitants of caves or of the earth underground. On the contrary, some organisms
have developed incredible transparency; consider the naturally transparent frog
Hyalinobatrachium bergeri (Fig. 1, Castroviejo-Fisher et al. 2007).
In these rather rare cases, the entrails and other visible inner structures tend to be
arranged symmetrically, so the entire display is not disturbed by non-aesthetic
asymmetry of non-paired organs. In any case, the outer surface of most organisms
that enterdue to their way of lifethe visual sensing of other beings is usually
opaque and often intricately patterned. It is also highly probable that, in some groups
of animals, the evolutionary origin and further development of exposed organic
surfaces went hand in hand with the perfection of optic perception. This does not
mean, however, that pigmentation did not exist before the emergence of visual
sensing. In very early life forms, various pigments were certainly present and had a
functional role such as protection against ultraviolet radiation. Such a life-sustaining
function may also be used to explain the presence of pigment in modern organisms.
Nevertheless, specificity and peculiarity of various sophisticated ornaments as
well as intriguing color patterns cannot be explained by such functioning; hence,
one should presuppose the existence of some representational purpose of these
phenomena proper.
The perfection of sight in animals led to the fact that some of these non-addressed
phenomena proper have acquired a specific meaning in the umwelten of some
animals and thus have become addressed. The origin of vision was the crucial event
The semantic morphology of Adolf Portmann 213
that increased communicative abilities of organisms; and, again, it was exactly this
event in which many life-forms attained their very appearance.
In the most general sense, semantic organs are characterized as phenomena
proper that are meaningful within the umwelten of particular living beings. This
category is comprised of, not only everything that is topologically present on the
surface, but all entitiesexternal or internal, irrespectivelythat are exposed, and
therefore perceivable, on the surface. Haemoglobin, for example, has a clear function
in binding and distributing oxygen in animal bodies, but this does not exhaust the
biological meaning that comes from the red color of its blood pigment (Portmann
1960b: 222). The redness of blood often serves as a signal, usually with a meaning
that is alarming to other animals in their surroundings, regardless of whether they are
friends or predators. Consider, for instance, the dentition that usually serves as an
effective hunting and food processing structure, which, however, may also be
interpreted as a semantic organ that clearly states the inner-attitude of the bearer. The
next important property of such organs is that (1) semantic organs that have the same
meaning need not necessarily be generated by the same morphogenetic pathways;
(2) the same phenomena proper may acquire different meanings depending on the
inner attitude of an animal interpreter, or on the environmental context, and be thus
considered different semantic organs. These aspects may be readily demonstrated by
the example of the eyespots, the false eyes that are exposed on the surface of animals
from many different groups.
The wings of nymphalid and satyrid butterflies often bear a conspicuous row of
eyespots that are homologous to the element of border ocelii within the generalized
wing pattern of the nymphalid ground plan (Sffert 1927, Nijhout 1991). In contrast,
Fig. 1 The naturally transparent
frog Hyalinobatrachium bergeri
(redrawn after Castroviejo-
Fisher et al. 2007)
214 K. Kleisner
the eyespots on the hindwings of some sphingid moths originate from elements of
the central symmetry system, that is, from a different part of the nymphalid
groundplan. There are several hypotheses that try to explain the role of eyespots,
describing them, for example, as marks of direction with a deflective function, or as
signifiers of intimidation or sexual selection (Stevens 2005). In the satyrid butterfly
Bicyclus anynana, different roles are hypothesized for the eyespots that are on the
dorsal and on the ventral side of the wing. Whereas ventral eyespots are supposed to
act as antipredatory deflective marks meant to divert an attack towards the wing
margin away from the body itself (Blest 1957; Lyytinen et al. 2004), the dorsal ones
play a role in sexual selection by female choice acting on size and UV-reflectivity of
the white central pupil of dorsal eyespots (Robertson and Monteiro 2005).
Because the element of border ocelii consists of an array of eyespots, the whole
system has a modular character similar to that of the vertebrate backbone. In the
ancestral state, the eyespots were, presumably, highly coupled, both genetically and
developmentally. However, during the course of evolution, acting adaptive forces led
to their genetic decoupling, favoring individuation and independent diversification of
single eyespots (Beldade and Brakefield 2003). Artificial selection has been applied
to eyespots to explore their phenotypic plasticity. The results of selection experi-
ments on Bicyclus anynana show that the genetic and the morphological parameters
of the eyespots are positively correlated (Monteiro et al. 1994). Nevertheless, there
still is a significant potential for individual changes despite developmental coupling
within serially homologous eyespots (Beldade et al. 2002a; Beldade et al. 2002b).
Importantly, artificial selection had a rapid effect on size, location and color
composition of the eyespots, whereas much lower heritability was reported for
eyespot shape during selection for elliptical eyespots in the antero-posterior and
proximo-distal axes (Monteiro et al. 1997a; Monteiro et al. 1997b). As a result, it
seems evident that when compared to other parameters, the circular shape of the
eyespots is strongly constrained. This is probably because of developmental-
mechanistic reasons during the formation of the eyespot, such as the response of the
surrounding epidermis to radial diffusion of a signal from a central focus point (e.g.,
Nijhout 1980, 1991; French and Monteiro 1994). Alternatively, shared developmen-
tal constraints that retain the circularity of these elements are favored because they
increase the probability that the eyespots will be perceived as eye-like by a
majority of possible receivers and thus attract the sight of predators.
Most studies on the wing patterns of butterflies use eyespots as a model system of
development, wherein phenotypic changes in eyespot parameters provide informa-
tion on the functioning of causal mechanisms and underlying genetic factors. For
such approaches, the eyespots are an excellent experimental control, but do not help
us understand their very resemblance to true eyes. One should ask how these
eyespots became eye-like and how are they maintained as such during evolution.
Various groups of animals frequently imitate the eyes of vertebrates, a phenomenon
also known as partial mimicry. Different types of false eyes and false heads are
especially numerous; one can find such imitations of body parts in both vertebrate and
invertebrate clades (Komrek 2003: 73). Real eyes are subject to imitation because
they themselves function as semantic organs intended to bewilder potential
aggressors. The aposematic eyes of the red-eyed tree frog (Agalychnis callidryas)
are a striking example (see Fig. 2.).
The semantic morphology of Adolf Portmann 215
Note that there is no reason, stemming from the anatomy and physiology of sight,
why the sclera and iris should differ in color in the eyes of many vertebrates.
Consider also that patterned eyes are found among life forms that orientate visually,
such as many birds, humans and even nocturnal fruit bats, especially the later in
comparison with the atrophied eyes of most other bats that orientate acoustically.
This brings us to the idea that eyes often do not represent mere organs of sight, but
also serve as semantic organs that have a communicative or signaling role (see, e.g.,
Tomasello et al. 2007).
The semantic character of the eyes also explains similar properties of various false
eyes imitated in irrelevant positions on the body of a mimic. Therefore, we are often
able to recognize the (false) head positioned on the body opposite the real one; false
eyes are often placed in the occipital area of the head instead of the facial part, and
the eyes of vertebrates are often pictured on bodies of invertebrates. From such
perspective, there is no difference between real vertebrate eyes and fake eyes, or
between the black and yellow coloration of wasps and the wasp-like pattern of a
hover fly, because all these structures retain their iconic character irrespective of
whether they are original or mere imitation. These imitative representations have one
thing in common: they bear the same meaning in the umwelt of a particular perceiver.
In this respect, it does not matter whether some organs are identical (homologous)
from an evolutionary perspective or not, but it is important that they are considered
identical. From this type of reasoning comes the need for a comparative term in
biosemiotics that will stand for congruencies in the representation of sameness in the
umwelten of particular life forms. For these reasons the concept of homosemiosis
has been proposed (homosemiosis=correspondence of entities, parts and organs that
are interpreted as the same objects in the umwelt of a particular organism or group of
organisms under investigation; Kleisner 2008). Again, if eyespots are perceived as
real eyes, within the umwelt of a particular animal-interpreter, then we can say that
eyespots (i.e., images of eyes) are homosemiotic to eyes (i.e., organs of vision). The
same applies to all other semantic organs.
Fig. 2 The conspicuous eyes
of the red-eyed tree frog
(Agalychnis callidryas) do not
serve as mere organs of sight but
probably also as semantic organs
with a warning meaning
216 K. Kleisner
Conclusions
Adolf Portmann suggested that the opaque, shaped and patterned surfaces of
organisms do not represent a mere mechanical, chemical or informational interface
that demarcates the border between an organism and its outer world (Portmann
1960a: 102f., 1960b, 1969, 1990: 205). Rather, this outermost dimension of an
organism should be regarded as a peculiar kind of organ that finds its particular role
in the representation of the selfhood of a living being. This manifestation of
phenomena proper is no less important than the inward life-sustaining systems, e.g.
the nervous, the vascular, the digestive system etc. If we compare the arrangement of
the inner structures with the morphology of the outer surface, we recognize that the
visible surface is often independent of the underlying anatomy in terms of symmetry,
coloration and spatial pattern. This fact reveals the quasi-autonomous character of
the outer surface, which finds its unique role in expressing phenomena proper by
the process of self-representation of organic inwardness. When we focus more
intimately on the richness of appearance and behavior in and of organisms, we must
alsoamong other thingsconsider those behavioral acts and morphological
ornaments that represent the result of a particular inner attitude. As cases of self-
representation, these acts and ornaments can never be fully reducible to their mere
functionality for survival.
Phenomena proper, which meaningfully enter the umwelt of a particular
interpreter, may be considered semantic organs. Thus, it is true that every semantic
organ is a phenomenon proper but not vice versa. Semantic organs are semi-
autonomous entities that can be unambiguously derived from neither their anatomical,
morphological and genetic nature, nor from the standpoint of the physiology of a
perceiver. Rather, every semantic organ exists at the interface between the expression
of physical features and their interpretation. Therefore, semantic organs work thanks
to a meaningful interaction of embodied perceivable traits with the structure of the
umwelt of a particular perceiver.
Semantic organs may originate from an umwelt-specific interpretation of surface
patterns that takes place by morphogenetic processes during ontogenesis. However,
what is it that connects the genetic and the developmental potency of an organism
with its own umwelt? What powerful interactions make semantic organs exist and
persist? Presumably, here at the interface between the act of self-representation and
umwelt-specific interpretation, we should think about a set of conventions that
meaningfully connect two independent worlds (organic codes sensu Barbieri 1998,
2003, 2007: 188f.). On the other hand, these two dimensions of self-expression and
selfhood (inwardness) may eventually fall into unity because of the interdependence
of conventions that were negotiated during the evolution of umwelt-umwelt
interactions (as a result of biohermeneutic acts sensu Marko 2002; Marko et al.
2007). One way or another, if such conventions are disrupted, a particular semantic
organ will atrophy and degenerate.
Presumably, the evolutionary origin and maintenance of semantic organs depends,
first, on the heritability of the genetic and developmental precursors that underlie the
appearance of a specific perceivable surface in a particular bearer and, second, on
the composition of perceptual and cognitive properties (umwelt constitution) of the
The semantic morphology of Adolf Portmann 217
addressee. The former facilitates the emergence of patterns of structure and color,
while the latter ensures that such patterns are perceived within the umwelt of a
particular addressee and are thereby subject to selection processes. Selection may
further contribute both to the alteration and to the conservation of a particular
semantic organ, depending on the kind of interaction between the bearer and the
addressee. Significantly, the role of selection is defined by signification within the
umwelt of the interpreter, and not vice versa.
Acknowledgements I wish to thank Anton Marko for his comments on the text. I also owe my thanks
to Stanislav Komrek, who introduced me to the writings of Adolf Portmann, and to Zdenk Neubauer,
who prepared the first Czech translation of Portmanns book during the time of communist repression and
thus sowed the seeds of future interest in Portmanns ideas. Last but not least, I thank Lucie ermkov
for the beautiful drawings published in this paper. This work has been supported by the Research Program
CTS MSM 0021620845 and the GPSS Major Awards Program, a joint program of the Interdisciplinary
University of Paris and Elon University.
References
Amundson, R., & Lauder, G. V. (1994). Function without purpose: The uses of causal role function in
evolutionary biology. Biology & Philosophy, 9, 443469.
Barbieri, M. (1998). The organic codes. The basic mechanism of macroevolution. Rivista di Biologia-
Biology Forum, 91, 481514.
Barbieri, M. (2003). The organic codes: An introduction to semantic biology. Cambridge: University
Cambridge Press.
Barbieri, M. (2007). Is the cell a semiotic system? In M. Barbieri (Ed.), Introduction to biosemiotics (pp.
235255). Dordrecht: Springer.
Beldade, P., & Brakefield, P. M. (2003). Concerted evolution and developmental integration in modular
butterfly wing patterns. Evolution & Development, 5, 169179.
Beldade, P., Koops, K., & Brakefield, P. M. (2002a). Developmental constraints versus flexibility in
morphological evolution. Nature, 416, 844847.
Beldade, P., Koops, K., & Brakefield, P. M. (2002b). Modularity, individuality, and evo-devo in butterfly
wings. Proceedings of the National Academy of Sciences, 99, 1426214267.
Blest, A. D. (1957). The function of eyespot patterns in the Lepidoptera. Behavior, 11, 209256.
Castroviejo-Fisher, S., De la Riva, I., & Vil, C. (2007). Transparent frogs show potential of natural world.
Nature, 449, 972.
French, V., & Monteiro, A. (1994). Butterfly wings: Colour patterns and new gene expression patterns.
Bioessays, 16, 789791.
Kleisner, K. (2008). Homosemiosis, Mimicry, and Superficial Similarity: Notes on the conceptualization
of independent emergence of similarity in biology. Theory in Biosciences, 127, 1521.
Kleisner, K. (2007). The formation of the theory of homology in biological sciences. Acta Biotheoretica,
55, 317340.
Kleisner, K., & Marko, M. (2005). The semetic rings: Towards the new concept of mimetic resemblances.
Theory in Biosciences, 123, 209222.
Komrek, S. (2003). Mimicry, aposematism and related phenomena. Mimetism in nature and the history
of its study. Mnchen: Lincolm Europa.
Lyytinen, A., Brakefield, P. M., Lindstrm, L., & Mappes, J. (2004). Does predation maintain eyespot
plasticity in Bicyclus anynana? Proceedings of the Royal Society of London B, 271, 279283.
Macdonald, J. D. (18541855). Observations on the anatomy and affinities of the Phyllirrho bucephala
(Peron). Proceedings of the Royal Society of London, 7, 363368.
Maran, T. (2007). Semiotic interpretations of biological mimicry. Semiotica, 167, 223248.
Marko, A. (2002). Readers of the book of life: Contextualizing developmental evolutionary biology.
Oxford: Oxford University Press.
Marko, A., Grygar, F., Kleisner, K., & Neubauer, Z. (2007). Towards a Darwinian biosemiotics. Life as
mutual understanding. In Introduction to biosemiotics (pp. 235255). Dordrecht: Springer.
218 K. Kleisner
Monteiro, A., Brakefield, P. M., & French, V. (1997a). Butterfly eyespots: The genetics and development
of the color rings. Evolution, 51, 12071216.
Monteiro, A., Brakefield, P. M., & French, V. (1994). The evolutionary genetics and developmental basis
of wing pattern variation in the butterfly Bicyclus anynana. Evolution, 48, 11471157.
Monteiro, A., Brakefield, P. M., & French, V. (1997b). The genetics and development of an eyespot
pattern in the butterfly Bicyclus anynana: Response to selection for eyespot shape. Genetics, 146,
287294.
Nijhout, H. F. (1980). Pattern formation on lepidoptera wings: Determination of an eyespot.
Developmental Biology, 80, 287305.
Nijhout, H. F. (1991). The Development and Evolution of Butterfly Wing Patterns, Smithsonian Institution
Press, Washington, D.C.
Portmann, A. (1960a). Neue Wege der Biologie. Mnchen: Piper.
Portmann, A. (1960b). Die Tiergestalt. Studien ber die Bedeutung der tierischen Ercheinung. Basel:
Friedrich Reinhardt.
Portmann, A. (1965). Neue Fronten der biologischen Arbeit. In: G. Schulz (Ed.), Transparente Welt.
Festschrift zum 60. Geburtstag von Jean Gebser. Bern: Huber: 2337.
Portmann, A. (1969). Einfhrung in die vergleichende Morphologie der Wirbeltiere. Basel: Schwabe & Co.
Portmann, A. (1990). Essays in philosophical zoology by Adolf Portmann. The living form and seeing eye.
Lewiston: Edwin Mellen.
Robertson, K. A., & Monteiro, A. (2005). Female Bicyclus anynana butterflies choose males on the basis
of their dorsal UV-reflective eyespot pupils. Proceedings of the Royal Society of London B, 272,
15411546.
Russell, E. S. (1916). Form and function: A contribution to the history of animal morphology. London:
Murray.
Stevens, M. (2005). The role of eyespots as anti-predator mechanisms, principally demonstrated in the
Lepidoptera. Biological Reviews, 80, 573588.
Sffert, F. (1927). Zur vergleichenden Analyse der Schmetterlingzeichnung. Biologisches Zentralblatt, 47,
385413.
Tomasello, M., Hare, B., Lehmann, H., & Call, J. (2007). Reliance on head versus eyes in the gaze
following of great apes and human infants: The cooperative eye hypothesis. Journal of Human
Evolution, 52, 314320.
von Uexkll, J. (1921). Umwelt und Innenwelt der Tiere. Berlin: Springer.
von Uexkll, J. (1928). Theoretische Biologie. Berlin: Springer.
Wray, G. A., & Abouheif, E. (1998). When homology is not homology. Current Opinion in Genetics &
Development, 8, 67568.
The semantic morphology of Adolf Portmann 219