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 Portmannor, more precisely, his concept of organic self-representation, wherein Portmannconsidered the outer surface of living organisms as a specific organ that servesin a self-representational role. This idea is taken as a starting point from whichto elaborate Portman’s ideas, so as to make them compatible with the theoreticalframework of biosemiotics. Today, despite the many theories that help us understandaposematism, camouflage, deception and other phenomena related to the category ofmimicry, there still is a need for a general theory of self-representation that wouldre-synthesize evolutionary, morphogenetic and semiotic aspects of the surface oforganisms. Here, Adolf Portmann’s concept of self-representation is considered asan 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, withspecial emphasis on its representational aspects (Portmann 1960a, 1960b, 1969).This paper re-introduces and elaborates his basic ideas, in the belief that they canbe a source of inspiration for modern biosemiotics. The evolution of particularconstituents of the body may be driven, not only by selective pressures that increasetheir functional utility, but also by their ability to interact with the umwelten of otherliving beings in a meaningful and contextual way. This is to say that without theseinteractions, the functioning of any sexual display, mimicry or deceptive behaviorcan hardly be understood. Organs of display (i.e., semantic organs) act through themeaning that they acquire during umwelt-specific interpretations.

Biosemiotics (2008) 1:207–219DOI 10.1007/s12304-008-9014-4

K. Kleisner (*)Department of Philosophy and History of Science, Charles University, Viničná 7, 128 44 Prague,Czech Republice-mail: kleisner@seznam.cz

Portmann’s investigations were strongly influenced by the theoretical biology andUmweltforschung of Jakob von Uexküll (1921, 1928), as well as by his comparativestudies on morphology and behavior. In practice, Portmann’s contribution wasovershadowed by the success of ethology, on the one hand, and by the generalenthusiasm for genetic and molecular methods on the other. The increasing popularityof genetic research was accompanied by a devaluation of morphology, which lostits 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 anintellectual climate, Adolf Portmann was a proponent of an old-fashioned traditionof continental biology, and while he may look rather anachronistic in his strongfocus on the “realm of the visible”, he frequently criticized the prevailing researchof his period for its orientation on the realm of the invisible, and its neglect oflife’s most spectacular aspects (Portmann 1960a, 1965). Biosemiotics now brings atheoretical account of life that seems sufficiently open to embrace the ideas ofAdolf 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 ofselective pressures caused either by other organisms or by environmental conditions.This way of reasoning is entirely at the service of survival. Every part of anorganism must be formed in such a way as to fulfill its life-sustaining function; fromthis it follows that reproduction is the sole and ultimate purpose of all formal-structural representations of life. Consequently, reproduction is the sole referentialframe that is adequate to explain all manifestations of life.

On the other hand, reasoning from intrinsic causation accents the importance ofthe inner potentialities of organisms that generate a particular form. In modernbiology, the activity of developmental genes is usually assumed to underlie theemergence of particular morphologies. Thanks to the success of bottom–upmethodologies in biotechnology, molecular and developmental genetics etc., biologistsascribe to genes the meaning of causal function, and morphology has acquired thepractical role of testing the causal functions of the genes. Consider, for instance, theintertwined studies on gene activation/inactivation, the mutation and the regulation ofgene expression via the monitoring of changes in morphology. On second thought, weknow that the results obtained at the genetic level are often used to explain the evolutionof forms (morphologies). However, do these attempts really seek to explain theevolution of organic form? Or is it rather the logic of the interactions betweenconstituents of different levels that is revealed? Is it ever possible to understand thetransformations of constituents at different higher levels via functional causationsbetween 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 evolutionarydissociations, that is, from incongruent relationships among various levels of biologicalorganization. In principle, two kinds of dissociations are imaginable between genes andmorphologies: (1) homologous genes are involved in the formation of nonhomologous

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structures, and (2) nonhomologous genes are recruited to serve the formation ofhomologous morphologies (Wray and Abouheif 1998).

The results obtained at the genetic level are useful for studying the relationshipsbetween genes and morphologies, but what is actually revealed here is the complexityof those relationships, and the obtained results provide a rather restricted account ofthe real transformations of organic form. This is so because behavior at the form-structure level somehow transcends the functional causation of constituents at anyunderlying level. And so we address the morphological aspects of an organism neitherfrom a strictly intrinsic nor from a purely extrinsic perspective, but attempt to explainsome of their evolutionary transformations via the potential of a morphology thatexceeds its functional necessity. The form (Gestalt) of organisms should be studied forits own sake (Portmann 1990: 138). In this framework, the formal aspects of livingbeings, such as shape or surface patterns, coloration or ornamentation, should beapproached neither as a useful control in genetic experiments, nor as a possiblesource of data for phylogenetic inference.

The uniqueness of Adolf Portmann’s approach may be found in his explanationthat organic form is in itself something valuable. Portmann acknowledged that theexternal surface of an organism has its own formal value and a certain kind ofautonomy over other life-sustaining functions. He was convinced that this outermostaspect of an organism opens a way to the innermost dimensions, because surfacemanifestation reflects inner self-experience and the very selfhood of every organicbeing, thus bringing us closer to understanding the existence of living creatures(Portmann 1969: 315). For these reasons he developed a terminology, or rather aconceptual framework, specific to such a purpose.

Portmann’s Conceptual Framework and Self-Representation of Organisms

Portman’s central idea can be formulated as follows: the outermost surface of anorganism 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 thesenses of another living being, that is, those aspects of the organism which are exposedto the perception of another subject in a natural, non-invasive, way. For example, thevarious colorations, ornaments, coat patterns and other morphological features of theoutermost organismal surface are considered to be manifestations of phenomenaproper 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 organismwhich are naturally concealed from the senses. All organs and organ systems that arenot immediately apparent from the outside—typically inner structures that ensure theenergetic and the mechanical functioning of the organic whole—belong to thiscategory and were marked by Portmann as phenomena improper (uneigentlicheErscheinungen). In contrast to phenomena proper, improper phenomena are notprimarily intended to arouse the sensation of another organic subject. All phenomenaproper serve to present the self; rising up from the inwardness of a living being, whichcan be understood as a particular form of organic self-experience. In contrast,

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phenomena improper do not normally attain any self-representational role (Portmann1960a: 102f., 1960b, 1965, 1969).

Portmann distinguished between two types of phenomena proper: addressed andnon-addressed (adressierten und unadressierten Erscheinungen; Portmann 1960a:109f., 1965). Addressed phenomena represent a largely unproblematic category thatis frequently well explained within contemporary biological theory. Addressedphenomena function as signals assigned to a particular receiver and thus partake inthe survival of the bearer. Cases such as warning coloration, camouflage, sexualdisplay etc. may serve as good examples. Non-addressed phenomena, on the otherhand, do not serve any practical purpose; they do not play any functional role relatedto survival. Yet, non-addressed manifestations represent indispensable features thatcharacterize 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 therichness of life-expressions cannot be resolved solely by functional explanations inthe context of self-preservation and reproduction of organic beings. Non-addresseddisplays represent those phenomena of life, those peculiarities, which are notconditioned by any functional necessity. This non-addressed domain of all life maythus be meaningfully embodied in the concept of self-representation.

At first sight, Portman’s conceptual framework may be regarded as ratherunusual, even somewhat extraordinary, but still it remains within common biologicalpractice. Although it emphasizes aspects of living beings that cannot be fullyexplained in terms of function, selection and survival, this approach remains to someextent compatible with the traditional way of functionalist and technologistreasoning, which does not necessarily stand in contradiction to the idea of organicself-representation.

This may be demonstrated by the corporeal transparency of some sea creaturessuch as, for example, salps, some species of fish, or some nudibranchiate mollusks.Using such examples, Portmann demonstrates the limits of a technical explanation ofanimal form. Normally, in a majority of animal forms, the asymmetry of the innerorgans is hidden beneath an opaque surface that is symmetrically arranged.However, we can observe a peculiar solution to inner asymmetry in transparentand semi-transparent living forms, wherein the symmetry is preserved by centeringall asymmetrical inner organ complexes into a single opaque visceral cluster(nucleus vegetativus; Portmann 1960a: 104). Inner structures that are not centeredinto an opaque visceral cluster are usually arranged symmetrically along the mainbody axis. It is also worthy of note that a symmetrical arrangement of inner organsin life forms with jellylike transparent bodies—as is the case of transparentnudibranchiates from the genus Phyllirrhoe1, to use Portmann’s example—may also

1 A description of the species Phyllirrhoë bucephal from over 150 years ago makes us wonder about theintriguing architecture of the symmetrical corporeal composition of this animal: “The outer integument isperfectly transparent and lined by muscular bundles, disposed longitudinally, and somewhat more thantheir own breadth apart. These communicate with one another by oblique branching lips, which thus forma kind of network enclosing long lozenge-shaped spaces. Here and there nerve-trunks of considerable sizeaccompany the longitudinal bundles, dividing off into smaller twigs, which distribute themselves at prettyequal distances in a direction more or less perpendicular to that of the muscular fibres.” (McDonald 1854:363).

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be explained in technical terms as a stabilizer during swimming or floating inthe water column. Transparency itself can be explained in technical terms, as asecondary effect of a high water content in gelatinous tissues which ensuresbuoyancy of the body in analogy to the swim bladder of fish. Even the concentrationof asymmetrical visceral structures into a single centered cluster in many fish may beseen as a constructional solution that increases stability and maneuverability of thebody. This is probably true, but it does not tell the whole story. From the perspectiveof technology, there is no reason why entrails organized in a visceral cluster shouldbe divided by an opaque pigment.

Portmann demonstrates that many life forms tend to preserve their optical symmetrydespite the asymmetrical nature of their inner organs, which are, architectonically, not tobe seen. In this respect, the concept of “phenomena proper” (eigentliche Erscheinungen)as manifested by formal aspects of an exposed surface, exhibits superiority over purelyfunctional explanations. Portmann does not reject functional explanations as such;nevertheless, he shows that function-based reasoning is unable to answer all questionsrelated to organic form.

One could admit that self-representation of phenomena proper is nothing morethan a kind of function. In other words, the concept of function might also beextended to the realm of organic self-representation whose particular manifestations(i.e., phenomena proper) bring some selective advantage to its bearer. However, byplacing phenomena proper into the context of function, only addressed phenomenaproper 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 arethought to have a particular evolutionary purpose for which they have been selectedin the course of evolution.2 In contrast, non-addressed phenomena proper certainlydo not fit this scenario; they are not “deliberately” applied to the sensory organ of aparticular 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 reasonswhy they should be privileged by natural/sexual selection. For instance, why aresome beetles neutrally brownish while others are metallic, or why do somebutterflies expose intricate, bizarre, albeit species-specific, color patterns with nostriking adaptive role? Interestingly, Zdeněk Neubauer (personal communication)considers these taxon-specific appearances as heraldic signs that provide evidence ofontological authenticity and sovereignty. A coat of arms, or blazon of arms, oftenappears 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 theyserve 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 toexternalization of organic inwardness through the process of self-representation;these phenomena are always of a taxon-specific character, having their source in theinner causes of an organism. Even though this “internalistic” position of Portmann’swould 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 andLauder 1994).

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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 problemof organic form entirely. In principle, and irrespective of whether they are addressedor non-addressed, all visual phenomena proper presume the existence of a “seeingeye” (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 notmean that no other being is there to perceive them as somewhat meaningful. As anexample, imagine the black and yellow coloration of wasps. There probably is nosingle and sole addressee of this black and yellow pattern at which it was originallyaimed; 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-likepattern 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 andwhy this pattern appeared in wasps themselves. In other words, when we ask as towhy an organism resembles a wasp, we simultaneously pose the question: “why isa 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 offunction. However, does this apply to addressed phenomena as well? One mayrightly 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 originalpurpose, if there was one, could have been quite different. Nevertheless, not even theeffectiveness of a warning coloration can be properly understood within the realm offunction, because the cogency of any display is conditioned by circumstances thatexceed any concept of function. This brings us to the role of meaning. How does thequestion of the function of a structure differ from the question of the meaning of thesame structure? For example, if we ask about the function of eyespots on the wingsof a butterfly, we may answer that they serve to intimidate avian predators. This tellsus that the structure under investigation functions as a warning signal with aprotective function for the bearer. Nevertheless, such a question does not drawattention to the subject affected (in this case intimidated) by the eyespots. Nothing istold about the receiver’s perception and interpretation of these structures. In contrast,if we ask about the meaning of the eyespots, the question is put as to whether theseeyespots are interpreted as real eyes within the umwelt of some animal, or whetherthey relate to some unspecified dangerous thing (see abstract mimicry sensu Maran2007: 239). In other words, the question regarding the function of the eyespots canbe satisfactorily answered by describing how the eyespots affect the survival of theprey in a prey–predator interaction. However, to ask about the meaning of theeyespots is an attempt to answer the question as to what do these structures signify toan addressee, what do they represent in the perceptual world (umwelt) of a receiver,and to what extent is the receiver’s signification of these structures comparable to

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that of ours. Investigating functionality is the domain of the mechanistic approachand belongs to the program of adaptationism, whereas asking about meaning fallswithin the province of biosemiotics.

Semantic Organs: Concept and Examples

Traditionally, the morphological and anatomical constituents of animal bodies havebeen conceptualized in terms of form or function. Differing scholars have valuedeither the first or the second as the best explanation of the reasons that shape andtransform organic matter (Russel 1916; Kleisner 2007). Herein, we attempt tointroduce the idea of semantic organs, that is, organs that are defined neither by theirform nor by their function, but by their meaning in the umwelt of an organism, or agroup of organisms, under investigation (cf. Kleisner 2008; Kleisner and Markoš2005). In the following discussion, we focus on optical representations, although theexistence of semantic organs that are based on olfactory or acoustic representationsis clearly imaginable within the same theoretical framework.

This attempt stemmed from Portmann’s idea that the opaque surfaces oforganisms represent a new specific kind of organ, and not merely a mechanicalbarrier that binds together inner structures and metabolic processes in order toprotect them from external influences. Presumably, the appearance of ornamentedopaque surfaces is associated with the origin of vision. It is highly probable thatprior to the emergence of vision, most organisms had a semitransparent surface, notdissimilar from the milky whitish semitransparent coloring of vertebrate embryos. Acertain comeback of such neutral and indistinct surface colorations is apparent inorganisms dwelling in a dark environment, especially in parasitic forms or ininhabitants of caves or of the earth underground. On the contrary, some organismshave developed incredible transparency; consider the naturally transparent frogHyalinobatrachium bergeri (Fig. 1, Castroviejo-Fisher et al. 2007).

In these rather rare cases, the entrails and other visible inner structures tend to bearranged symmetrically, so the entire display is not disturbed by non-aestheticasymmetry of non-paired organs. In any case, the outer surface of most organismsthat enter—due to their way of life—the visual sensing of other beings is usuallyopaque and often intricately patterned. It is also highly probable that, in some groupsof animals, the evolutionary origin and further development of exposed organicsurfaces went hand in hand with the perfection of optic perception. This does notmean, however, that pigmentation did not exist before the emergence of visualsensing. In very early life forms, various pigments were certainly present and had afunctional role such as protection against ultraviolet radiation. Such a life-sustainingfunction may also be used to explain the presence of pigment in modern organisms.Nevertheless, specificity and peculiarity of various sophisticated ornaments aswell as intriguing color patterns cannot be explained by such functioning; hence,one should presuppose the existence of some representational purpose of thesephenomena proper.

The perfection of sight in animals led to the fact that some of these non-addressedphenomena proper have acquired a specific meaning in the umwelten of someanimals and thus have become addressed. The origin of vision was the crucial event

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that increased communicative abilities of organisms; and, again, it was exactly thisevent in which many life-forms attained their very appearance.

In the most general sense, semantic organs are characterized as phenomenaproper that are meaningful within the umwelten of particular living beings. Thiscategory is comprised of, not only everything that is topologically present on thesurface, but all entities—external or internal, irrespectively—that are exposed, andtherefore perceivable, on the surface. Haemoglobin, for example, has a clear functionin binding and distributing oxygen in animal bodies, but this does not exhaust thebiological meaning that comes from the red color of its blood pigment (Portmann1960b: 222). The redness of blood often serves as a signal, usually with a meaningthat is alarming to other animals in their surroundings, regardless of whether they arefriends or predators. Consider, for instance, the dentition that usually serves as aneffective hunting and food processing structure, which, however, may also beinterpreted as a semantic organ that clearly states the inner-attitude of the bearer. Thenext important property of such organs is that (1) semantic organs that have the samemeaning need not necessarily be generated by the same morphogenetic pathways;(2) the same phenomena proper may acquire different meanings depending on theinner attitude of an animal interpreter, or on the environmental context, and be thusconsidered different semantic organs. These aspects may be readily demonstrated bythe example of the eyespots, the false eyes that are exposed on the surface of animalsfrom many different groups.

The wings of nymphalid and satyrid butterflies often bear a conspicuous row ofeyespots that are homologous to the element of border ocelii within the generalizedwing pattern of the nymphalid ground plan (Süffert 1927, Nijhout 1991). In contrast,

Fig. 1 The naturally transparentfrog Hyalinobatrachium bergeri(redrawn after Castroviejo-Fisher et al. 2007)

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the eyespots on the hindwings of some sphingid moths originate from elements ofthe central symmetry system, that is, from a different part of the nymphalidgroundplan. 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 assignifiers of intimidation or sexual selection (Stevens 2005). In the satyrid butterflyBicyclus anynana, different roles are hypothesized for the eyespots that are on thedorsal and on the ventral side of the wing. Whereas ventral eyespots are supposed toact as antipredatory deflective marks meant to divert an attack towards the wingmargin away from the body itself (Blest 1957; Lyytinen et al. 2004), the dorsal onesplay a role in sexual selection by female choice acting on size and UV-reflectivity ofthe white central pupil of dorsal eyespots (Robertson and Monteiro 2005).

Because the element of border ocelii consists of an array of eyespots, the wholesystem has a modular character similar to that of the vertebrate backbone. In theancestral state, the eyespots were, presumably, highly coupled, both genetically anddevelopmentally. However, during the course of evolution, acting adaptive forces ledto their genetic decoupling, favoring individuation and independent diversification ofsingle eyespots (Beldade and Brakefield 2003). Artificial selection has been appliedto eyespots to explore their phenotypic plasticity. The results of selection experi-ments on Bicyclus anynana show that the genetic and the morphological parametersof the eyespots are positively correlated (Monteiro et al. 1994). Nevertheless, therestill is a significant potential for individual changes despite developmental couplingwithin serially homologous eyespots (Beldade et al. 2002a; Beldade et al. 2002b).

Importantly, artificial selection had a rapid effect on size, location and colorcomposition of the eyespots, whereas much lower heritability was reported foreyespot shape during selection for elliptical eyespots in the antero-posterior andproximo-distal axes (Monteiro et al. 1997a; Monteiro et al. 1997b). As a result, itseems evident that when compared to other parameters, the circular shape of theeyespots is strongly constrained. This is probably because of developmental-mechanistic reasons during the formation of the eyespot, such as the response of thesurrounding 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 theyincrease the probability that the eyespots will be perceived as “eye-like” by amajority of possible receivers and thus attract the sight of predators.

Most studies on the wing patterns of butterflies use eyespots as a model system ofdevelopment, wherein phenotypic changes in eyespot parameters provide informa-tion on the functioning of causal mechanisms and underlying genetic factors. Forsuch approaches, the eyespots are an excellent experimental control, but do not helpus understand their very resemblance to true eyes. One should ask how theseeyespots became “eye-like” and how are they maintained as such during evolution.

Various groups of animals frequently imitate the eyes of vertebrates, a phenomenonalso known as partial mimicry. Different types of false eyes and false heads areespecially numerous; one can find such imitations of body parts in both vertebrate andinvertebrate clades (Komárek 2003: 73). Real eyes are subject to imitation becausethey themselves function as semantic organs intended to bewilder potentialaggressors. The “aposematic” eyes of the red-eyed tree frog (Agalychnis callidryas)are a striking example (see Fig. 2.).

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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 incomparison 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, butalso 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 falseeyes imitated in irrelevant positions on the body of a mimic. Therefore, we are oftenable to recognize the (false) head positioned on the body opposite the real one; falseeyes are often placed in the occipital area of the head instead of the facial part, andthe eyes of vertebrates are often pictured on bodies of invertebrates. From suchperspective, there is no difference between real vertebrate eyes and fake eyes, orbetween the black and yellow coloration of wasps and the wasp-like pattern of ahover fly, because all these structures retain their iconic character irrespective ofwhether they are original or mere imitation. These imitative representations have onething 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 consideredidentical. From this type of reasoning comes the need for a comparative term inbiosemiotics that will stand for congruencies in the representation of sameness in theumwelten of particular life forms. For these reasons the concept of “homosemiosis”has been proposed (homosemiosis=correspondence of entities, parts and organs thatare interpreted as the same objects in the umwelt of a particular organism or group oforganisms under investigation”; Kleisner 2008). Again, if eyespots are perceived asreal eyes, within the umwelt of a particular animal-interpreter, then we can say thateyespots (i.e., images of eyes) are homosemiotic to eyes (i.e., organs of vision). Thesame applies to all other semantic organs.

Fig. 2 The conspicuous eyesof the red-eyed tree frog(Agalychnis callidryas) do notserve as mere organs of sight butprobably also as semantic organswith a warning meaning

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Conclusions

Adolf Portmann suggested that the opaque, shaped and patterned surfaces oforganisms do not represent a mere mechanical, chemical or informational interfacethat demarcates the border between an organism and its outer world (Portmann1960a: 102f., 1960b, 1969, 1990: 205). Rather, this outermost dimension of anorganism should be regarded as a peculiar kind of organ that finds its particular rolein the representation of the selfhood of a living being. This manifestation ofphenomena 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 ofthe inner structures with the morphology of the outer surface, we recognize that thevisible surface is often independent of the underlying anatomy in terms of symmetry,coloration and spatial pattern. This fact reveals the quasi-autonomous character ofthe outer surface, which finds its unique role in expressing phenomena proper bythe process of self-representation of organic inwardness. When we focus moreintimately on the richness of appearance and behavior in and of organisms, we mustalso—among other things—consider those behavioral acts and morphologicalornaments 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 merefunctionality for survival.

Phenomena proper, which meaningfully enter the umwelt of a particularinterpreter, may be considered semantic organs. Thus, it is true that every semanticorgan 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 aperceiver. Rather, every semantic organ exists at the interface between the expressionof physical features and their interpretation. Therefore, semantic organs work thanksto a meaningful interaction of embodied perceivable traits with the structure of theumwelt of a particular perceiver.

Semantic organs may originate from an umwelt-specific interpretation of surfacepatterns that takes place by morphogenetic processes during ontogenesis. However,what is it that connects the genetic and the developmental potency of an organismwith its own umwelt? What powerful interactions make semantic organs exist andpersist? Presumably, here at the interface between the act of self-representation andumwelt-specific interpretation, we should think about a set of conventions thatmeaningfully connect two independent worlds (organic codes sensu Barbieri 1998,2003, 2007: 188f.). On the other hand, these two dimensions of self-expression andselfhood (inwardness) may eventually fall into unity because of the interdependenceof conventions that were negotiated during the evolution of umwelt-umweltinteractions (as a result of biohermeneutic acts sensu Markoš 2002; Markoš et al.2007). One way or another, if such conventions are disrupted, a particular semanticorgan 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 theappearance of a specific perceivable surface in a particular bearer and, second, onthe composition of perceptual and cognitive properties (umwelt constitution) of the

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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 aparticular addressee and are thereby subject to selection processes. Selection mayfurther contribute both to the alteration and to the conservation of a particularsemantic organ, depending on the kind of interaction between the bearer and theaddressee. Significantly, the role of selection is defined by signification within theumwelt of the interpreter, and not vice versa.

Acknowledgements I wish to thank Anton Markoš for his comments on the text. I also owe my thanksto Stanislav Komárek, who introduced me to the writings of Adolf Portmann, and to Zdeněk Neubauer,who prepared the first Czech translation of Portmann’s book during the time of communist repression andthus sowed the seeds of future interest in Portmann’s ideas. Last but not least, I thank Lucie Čermákováfor the beautiful drawings published in this paper. This work has been supported by the Research ProgramCTS MSM 0021620845 and the GPSS Major Awards Program, a joint program of the InterdisciplinaryUniversity of Paris and Elon University.

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