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Biosemiotic Criticism

Abstract and Keywords

This article examines the concept of biosemiotic criticism. It contrasts biosemiotics with the semiology of Ferdinand de Saussure and provides an overview of biosemiotics as a synthetic biological discipline. It describes how the emergence of biosemiotics widened the sphere of semiotic processes to embrace all living organisms on Earth and offers a perspective of what biosemiotic criticism might be. The article also considers attempts to develop models that would bridge biosemiotics (or semiotic thinking more generally) and cultural or literary criticism.

Keywords: biosemiotic criticism, semiology, Ferdinand de Saussure, synthetic biological discipline, semiotic process, cultural criticism, literary criticism

Biosemiotics, described in the most general way, is a discipline that examines sign processes, meanings, and communication in and between living organisms. Biosemiotic criticism could be defined as the study of literature and other manifestations of human culture with an emphasis on the biosemiotic understanding that life is, down to its most fundamental levels, organised by sign processes. A rather similar term—literary biosemiotics—was proposed by W. John Coletta in 1999,1 but I prefer biosemiotic criticism for the same reason that I believe ecocriticism should be favored over literary ecology: both paradigms strive to account for the environmental aspect of various cultural phenomena, not only literary works. The present essay gives a brief overview of biosemiotics as a synthetic biological discipline, draws up a list of possibilities for describing humans’ semiotic relations with their environment, and discusses some synthetic applications and models. It must be noted, however, that biosemiotics is a recent and quickly developing discipline that is still negotiating its theoretical base and conceptual framework. The present essay develops a perspective of what biosemiotic criticism might be, but the reader should be aware that alternative possibilities exist.

Introduction to Biosemiotics

For most people in the humanities, at least in Europe, the word semiotics is associated, first of all, with the structuralist tradition, the semiology of Ferdinand de Saussure, the Prague linguistic circle, Louis Hjemslev, Claude Lévi-Strauss, Roland Barthes, and other representatives of the same tradition of thought. On the other hand, biosemiotics relates to another tradition of thought that, inside of semiotics, has become more and more eminent in the recent decades. This tradition proceeds from the semiotics of the American philosopher Charles Sanders Peirce and has been elaborated by his students or followers Charles W. Morris, Thomas A. Sebeok, Jesper Hoffmeyer, John Deely, and many others. The central concept for this tradition (or semiotics proper as (p. 261) opposed to semiology) is the concept of sign, or its dynamical aspect—semiosis, that in the most general way can be described as a mediated relation or mediated change. To pick one among the many definitions of sign by Peirce, a sign is “something which stands to somebody for something in some respect or capacity.”2 As to semiosis, Peirce has described it as “an action, or influence, which is, or involves, a cooperation of three subjects, such as a sign, its object, and its interpretant, this tri-relative influence not being in any way resolvable into actions between pairs.”3 Peirce’s complex terminology remains outside the scope of the present article; let it simply be noted that Peirce’s semiotics deals with the mediated or triadic relations (as being opposed to dyadic or physical relations between objects). Unlike European semiology, which focuses on sign structures or systems, Peircean semiotics is also capable of dealing with various local sign relations in nature. There are various examples of such relations in nature that have sign property: for instance, courtship feeding where a female passerine displays a begging behaviour with flapping wings, head, and neck bended down and beak opened that may be a sign of dependency relationship and relates to the feeding behavior of chicks; or replacement behavior in which a kitten who plays with a ball of yarn and bites it as if it were a prey animal is at the same time also aware of the difference and does not try to eat the yarn.

Another important source for biosemiotic paradigms has been the meaning-centred Umwelt theory of the Baltic-German biologist Jakob von Uexküll (1982, 1992).4 He has described an animal and that part of the environment it lives in as mutually coupled through meaning relations containing perception and action and through correspondence between animal body forms and environmental objects. Uexküll argued that those and only those parts of the environment meaningfully linked with an animal are present for it and are contained in its subjective universe or Umwelt. Nature, in Uexküll’s view, is construed by meaningfully connected perception and action points of different animal species, or points and counterpoints, as he borrows musical terms for expressing this holistic understanding. While Peircean semiotics provides biosemiotics with a view of relational signs capable of connecting organisms and objects in the environment, Uexküll’s theory of meaning allows sign processes to be grounded in bodily and biological organization but to be also seen in the framework of ecological relations that connect animal species and bind together ecosystems.

The meeting of Peircean semiotics and Uexküllian biology has probably been a prerequisite for the biosemiotic maxim, according to which semiosis is intrinsically connected with life. Thomas A. Sebeok, a Hungarian-born founder of biosemiotics, linguist, and semiotician has expressed this thought as follows: “the process of message exchanges, or semiosis, is an indispensable characteristic of all terrestrial life forms,”5 and also: “semiosis, independent of form or substance, is thus seen as a universal, criterial property of animate existence.”6 This idea has later been repeated in different wordings by Jesper Hoffmeyer, Kalevi Kull, Marcello Barbieri, and other leading biosemioticians. Biosemiotics has examined various processes in living systems as semiotic, from the transmission of genetic and molecular information on the cellular level up to intra- and interspecific communication in animals.7 Central principles of biosemiotics (p. 262) include self-organization and self-regulation of living systems; code-duality, such as the parallelism of digital and analogic information in the development of life; embodiment of communicative and interpretive processes; the organism’s inside–outside boundary as a semiotic filter or translation mechanism;8 sign processes as regulators of ecological relations and ecological communities; organisms as active shapers of their semiotic niches or Umwelten; and the growth of complexity of semiotic processes in biological evolution.9 Additionally, the history of biosemiotics can be interpreted in different ways as well as ramified by influences from phenomenology, hermeneutics, cybernetics, system theory, genetics, molecular biology, system biology, evolutionary developmental biology, and other fields.10 In general, the present essay focuses on two of several subfields of biosemiotics: zoosemiotics, the study of semiosis, communication and representation of animals;11 and ecosemiotics, semiotic relations between culture or organisms carrying it and the natural environment.12 These two subfields seem the most relevant for considering the connections between semiotic processes in nature and in human culture.

For the humanities, the emergence of biosemiotics widens the sphere of semiotic processes to embrace all living organisms on Earth, thereby ensuring that human cultural and semiotic activities cannot be treated as a semiotic island in the vast ocean of unsemiotic void. Rather, human culture should be considered as being surrounded by a multitude of other semiotic systems, some partly accessible, some rather different from ours. The issues that biosemiotics can bring to the attention of the humanities would include: (1) communicative and sign relations between human cultural activities and other semiotic subjects and their representations in literature and other cultural texts; (2) interrelations between environmental information and literary texts or other human cultural representations and the question of whether the latter may be motivated by the former; (3) the presence and traces of human bodily perception, sensations, and biological organization in literary texts and other human cultural representations; and (4) resemblances and analogies between literary texts or other cultural representations and elements of nature as such and the use of biosemiotic research models in the study of human culture in this aspect. These different possibilities and their practical applicability will be discussed more systematically in the following pages. In general, biosemiotic criticism emphasizes contextual and ecological reading and interpretation of the manifestations of the human culture and their natural surroundings.

Semiotics of Human–Nature Relations

One central question for discussion between biosemiotics and ecocritical studies on both the object and the paradigmatic levels is the relatedness of human cultural activities and nonhuman nature through semiotic means. The success in answering this question will settle whether biosemiotics as a paradigm of natural science has anything to offer to ecocriticism and whether the project of biosemiotic criticism is viable at all. (p. 263) Monist–dualist debates run deep in Western philosophy, the humanities and biological science13, but it seems that biosemiotics—by considering the capacity to interpret as an intrinsic property of the living matter—can add some fresh arguments to this discussion. I propose that the existing biosemiotic landscape can be organized around five types of relations bridging the human–nature divide: evolutionary, communicative, hierarchical, significational, and analogical. This typology should be taken as a tentative attempt to conceptualize the field of biosemiotic criticism.

1. Among these five possibilities, the evolutionary approach of relating humans with nonhuman nature is indeed most common in the humanities. It is based on the understanding of humans being descendants of the animal world, and it is applied by many Darwinian schools of humanities, such as literary Darwinism and evolutionary psychology, that seek to explain human culture through its evolutionary origin.14 Such approaches have often been accused of inclining towards biological determinism and reductionist descriptions as they may, for instance, try to explain human cultural texts by animal instincts and motivations. Although interest in the biological origins of human language and other semiotic systems is definitely present also in biosemiotics,15 the Darwinist way of relating humans with nonhuman nature is generally not characteristic of biosemiotics. What could be, however, a fruitful approach for biosemiotic criticism, would be to search for and identify such bases of similarities, provided by biological evolution, that function as points of departure both for zoosemiotic and linguistic modelling (on these concepts see below). Examples of such basic similarities include orientation on the vertical bottom-up axis that is a connecting feature for most animal and plants;16 the baby-schema (“Kindchen-schema,”17 or in English, “neotenic features”), a complex of face proportions characteristic of juvenile animals that is feature common to most vertebrates; or group relations and group hierarchies that is a connecting feature for most mammals. A resemblance on a certain level of biological organization provides the common ground on which interspecific communication and interpretation can be built (e.g., anthropomorphic depiction often uses and exaggerates the baby-schema features). The evolutionary connectedness of humans and animals can also be used as an argument in the humanities in a new way: instead of considering literary expression a means to maximize authors’ reproductive success, it is possible to study, for instance, the aesthetical and artistic behavior of other animal species.18

2. The communicative approach makes an attempt to widen the sphere of subjects that have culture or communication ability, in other words, to extend this sphere over the borders of our species, including at least some higher social mammals and birds. Such approach is characteristic of, for instance, cognitive ethology, cultural biology and some schools of environmental philosophy dealing with animal rights issues. In biosemiotics (or in zoosemiotics, to be exact), the communicative approach is present in debates as to whether any other animal species besides humans have language-like communication or to what extent humans can decode the communication systems of other species. These questions relate to the topic of the fundamental features of human language itself. In this discussion, Charles F. Hockett’s list of design features of human language and its (p. 264) applications in other communication systems remains a classical and still actively used source.19 There are also many ethological research studies, such as descriptions of ethograms or vocabularies of different species (a list of vocalizations of chaffinch by William H. Thorpe being an elegant example20). In essence, the communicative approach argues that instead of talking about the opposition between the humans and the environment, the environment itself can be seen as twofold, including the physical environment as well as semiotically competent animals who are much more similar to humans than we are to rocks or rivers. Animals’ actions have at least local intentionality, and this may turn them into an active party in the communicative relations with humans. At the same time, attributing language to other species besides humans is rarely done in biosemiotics, as the structural complexity of human language (especially because of lexical syntax) appears to far surpass any other communication system on Earth.

An interesting compromise and concept in this respect is that of “primary modelling system” or “zoosemiotic modelling system,” introduced by Sebeok, who argued that human capacity for linguistic communication is both ontogenetically and phylogenetically preceded by yet another modeling system—the-world-as-perceived—, where signs are distinguished by the organism’s species-specific sensory apparatus and nervous system and aligned with its behavioural resources and motor events.21 According to Sebeok, we possess at least two mutually sustaining modeling systems: the anthroposemiotic verbal, which is unique to the human species, and the zoosemiotic nonverbal, which unites us with the world of other animals. Verbal modeling may link further to higher ideological, poetic, artistic, or religious forms of modeling.22 Direct and spatial perceptions, tactile and smelling sensations, as well as many occurrences of nonverbal communication23 belong to the sphere of nonverbal modeling. Language does not have good resources for describing these kinds of phenomena, although it is certainly possible to express them. A particular target for biosemiotic criticism would be the manifestations of zoosemiotic modeling in a range of cultural artefacts.

  1. 3. The approach that I describe as hierarchical questions how we humans understand ourselves. The hierarchical approach argues that we are not uniform subjects, but rather hierarchical structures that contain many interacting layers of organization, all of which have their own subjectivity, memory, and semiotic competence. These claims are often supported by studies from neurology and molecular biology. For instance, Jesper Hoffmeyer has described the human immune system as a semi-autonomic agency with its own memory and activity and argued that the structure and functioning of the human nervous system is closer to swarm intelligence than a singular subject.24 Similarly, Sebeok has introduced the concept of a semiotic self: it is a multilayered structure, based on all the memory-capable codes in the body,25 including at least immunological, neurological, cognitive, and, in the case of human animals, also verbal and narrative layers. In her book The Whole Creature: Complexity, Biosemiotics and the Evolution of Culture,26 Wendy Wheeler argues for the interrelations between social, psychological, neurological and immunological systems in humans, referring to psychoneuroimmunological (p. 265) (PNI) studies of Paul Martin, Candace Pert, and others. Emphasizing the interactional nature of the human subject on different levels may also open biosemiotics up to issues of social criticism and theories of education and development in this context.27

  2. 4. The next way for human cultural activities to be related with nonhuman nature could be called significational, and this proceeds from the very heart of the logic of semiotics. The concept of significationality refers here to the works of the German semiotician Winfried Nöth,28 who has used this term to denote semiotic processes involving natural signs. For St. Augustine, from whom the concept of natural signs derives, natural signs lead to the knowledge of something else, but they are not used intentionally for communication (the footprints indicating the presence of an animal being a classical example). Natural signs can function because there are some correspondences or structural relations present in the natural world; for example, there can hardly be any animal footprints present in the landscape if no actual animal has visited this spot.

In arguing for the accessibility of natural environment for the living organism, biosemiotics may rely on some psychological or philosophic theory, such as James J. Gibson’s concept of environmental affordance29 or Michel Polanji’s philosophy of tacit knowledge.30 In its philosophical grounding, biosemiotics more often relates to pragmatism or scholastic realism (sensu John Deely31). For natural signs to function, the natural environment and the realm of representations need to be bridged, which presumes developed structurality in both. On the level of practical analysis, this may also mean juxtaposing literary representations of nature with collateral sources of knowledge such as scientific, folkloristic, or common-sense understandings of nature, as undertaken, for instance, by Kadri Tüür in her analysis of bird sounds in nature writing.32

To understand the functioning of natural signs, Peircean semiotic theory, which underlies much of contemporary biosemiotics, can also be helpful. As I mentioned before, a Peircean sign is essentially tripartite, consisting, first, of something that enters the attention of an individual—this would be a sign in the narrow sense or a representamen; second, of some object that this sign refers to; and third, of an interpretant that is some further thought, reaction or application related to this object. In such a tripartite structure, every sign is temporally organized, as it relates what is perceived to some object that has been before and leads to some future activity. Related to this, tripartite signs may induce motivatedness and intentionality into semiosis, as objects generate interpretants, whereas their properties remain constraints for possible interpretation.33 Peirce explains, “I define a Sign as anything which is so determined by something else, called its Object, and so determines an effect upon a person, which effect I call its Interpretant, that the latter is thereby mediately determined by the former.”34 At the same time, Peirce’s view is not deterministic, as an interpreter also needs to be capable of regarding a sign as a representamen of something else, and different individuals may interpret the same physical aspect of the sign (or the so-called sign vehicle) differently.

In principle, Peirce distinguishes three possibilities for how a representamen or sign in the narrow sense can be related to the object. In symbols, the signs (p. 266) characteristic of human linguistic communication, the relation is based on habit or convention. In addition to symbolic signs, Peirce speaks about icons, in which the relation between the representamen and the object is based on resemblance, and about indexes, where the relation is physical or spatial. We can see that in icons and indexes the motivatedness of the sign arises from the things present in the environment, whereas in the case of symbols it is induced because of human conventions. This rather complex semiotic vocabulary is introduced in order to show that for Peircean semiotics, human cultural artefacts can be immediately related to the environment if they include iconical or indexical aspects. Examples of iconical sign relations on the lexemic level of human language are metaphors and onomatopoeia;35 examples of indexical signs are deictic words. There are probably more possibilities for iconic and indexical relations in sentence structure and on the narrative level. We can also think of the natural environment inspiring us to create a piece of nature writing as a certain type of motivated sign activity.

Biosemiotics, stemming from the Peircean tradition, when applied to nature–culture relations, appears to establish an ecological perspective by emphasizing relations between nature writings, films, art, and so on, and the natural environment. This is especially so if one considers that also ecological relations between species have an essentially semiotic nature (forming what might be called semethic interactions36) by being shaped by processes of recognition and communication. A significational approach may lead to describing literature or other human cultural representations, human experience and the nonhuman environment as a nonhierarchical complex bounded together by sign relations, which is a local memory tradition or a local semiosphere. It may, for instance, deserve attention and reconsideration from this specific semiotic perspective that some particular landscapes and geographic areas have inspired their peculiar traditions of nature writing, with common motives and implicit references between the works of different authors.

  1. 5. The fifth possibility argues for the existence of deep structural parallels between the communication within and amongst living systems and those of human cultural manifestations, and suggests the possibility of using biosemiotic methods for describing literature or other representations in human culture on this basis. Such a possibility is historically influenced by studies of the genetic code and its possible analogy to human language, noted by linguists in the 1960s.37 Recent developments of this line of thinking include the Prague school of biohermeneutics38 that argues for the hermeneutic nature of the living world, life’s own role in creating the world in the evolutionary process and the possibility of using narrative strategies in understanding it. Similar ideas may depart from Peircean semiotics. For instance, W. John Coletta has studied the necessary conditions in nature for sign relations to emerge and underlined several parallels between the functioning of language and nature.39 Following the Peircean tradition, he has written about what he calls the “literary dimensions of nature”: nature’s embeddedness in language, its ironic and agentive features, its self-organizing and self-maintaining properties, its emergent properties, and its semiosis or signing action.40 His topics include resemblances between the functioning of language in predication and ecological (p. 267) relations, such as predation; he has also discussed human metaphors with reference to biological adaptations or natural metaphorics already present in the environment. For instance, Coletta argues that the specific resemblance between the coloration of a prey species and its natural background may produce a relationship similar to predication in human language. In the case of the fish Rock Beauty or Holocantus tricolor, with a yellow head and dark rock-like body, “the relationship between the indexical, specifying head of the fish and the iconic, complementary rock-like body of the same fish […] produces at least the effect of the phrase ‘this yellow fish behind the rock’ in the mind of interpretant or predator…”41

A practical research method inspired by a possible analogy in the functioning of nature and cultural representations would be to use Uexküll’s Umwelt analysis in the study of literary texts. Uexküll used this analysis to describing different animal Umwelten in comparison and in relation to each other. The research method allows for the formation of hypotheses about how the Umwelten of two species interact, as well as what the meaning of the characteristic behaviors of one animal might be for another, and vice versa. The groundwork for the Umwelt analysis is laid by von Uexküll in the book The Theory of Meaning,42 in which he analyses the Umwelt of a tick and the mammals’ place in it. The analysis consists of three parts. At first Uexküll determines the carrier and the receiver of meaning, thus establishing the position of the subject for the analysis. Then he describes the links between one animal’s sensory organs and the activities or features of another as corresponding points and counterpoints. Both parties are enclosed in their species-specific Umwelten and have their species-specific bodies and functionalities to use. As the third step of the analysis, on the basis of correspondence between different meaning points, Uexküll infers a common meaning rule connecting both organisms. For example, in the relation between the tick and an unspecified mammal, the meaning relation for the tick could be expressed as: “recognition and attack of the prey and extraction of blood.”43 We can assume that such method could also be used in the analysis of some literary novels, in which the different position and relations of the protagonists are the central theme. The Uexküllian approach would emphasize the subjective worlds of the protagonists, invite the discovery of correspondences in their characters, dialogues, and deeds (that could be either affirming or destructive), and examine the meaning of such relations on a higher level, either for the rest of the community in the narrative or for the structure of the book.

Towards Hybrid Research Models

When we look at the works of contemporary authors who are trying to bridge biosemiotics (or semiotic thinking more generally) and cultural or literary criticism, there appears to be a common endeavor to undermine the dualistic pairs of culture–nature, text–world and to develop a new framework that would connect semiotic processes (p. 268) in human culture with those outside it. Often, writings in biosemiotic criticism arrive at an understanding that this work cannot be completed only on the level of applied research but that a more general epistemological or paradigmatic shift is needed.44 This positional change would include a new understanding of the relations between the text and the world, the text and its reader, and the text and the researcher as semiosis-based. The natural environment, both in its animate and its physical existence, needs to be reinterpreted as holding semiotic potential. And as is accepted in the semiotics of culture, eventually the activity of the researcher can be interpreted as a continuation of the sign process that led to the writer’s inspiration, the creation of a text, and its interpretation by the reader; and the resulting review or analysis can be regarded as yet another layer of signs or texts in the same semiotic series.

For literary analysis, especially regarding nature writing as a research object, developing synthetic research models that could account for a written text and the natural environment in the same framework is an essential task. There exist several models, attempting to relate culture to the environment that it represents or is otherwise connected to, that are focused either on the natural or cultural component and that are either more or less explicitly semiotic than others. For instance, the British education theorist and semiotician Andrew Stables has introduced the notion of “landscape as text” and argued that the blurring of the concept of author in modern literary theory makes it possible to open the concept of text to natural phenomena. Stables notes that in landscapes the network of shared meanings extends beyond the human sphere and that it is difficult to draw a dividing line between the creative activities of humans, other life-forms, and natural forces.45 This parallel allows us to introduce as well the concept of environmental literacy as a natural equivalent to the competence of orienting in literary realms. A rather similar approach is introduced by an American cultural geographer Anne W. Spirn who has described the physical environment using language-related terminology. In her view, landscape contains “patterns of shape, structure, material, formation, and function”; it is “pragmatic, poetic, rhetorical, polemical” and can be “spoken, written, read or imagined.”46

In addition to arguing that nature or some of its elements have text-like characteristics, it is also possible to describe human cultural texts and nature-as-text as mutually intertwined. David Abram, in his investigation into the phenomenology of the more-than-human world,47 has described the mutual coupledness of texts and landscapes in many Native American (Amahuaca, Apache, Koyukon) and Australian traditional cultures. In these cultures, songs and tales help members of that culture to remember the properties, resources, and dynamics of the landscape as well as the proper behavior toward it, whereas the variability of landscape acts as a visual mnemonic pointing to the stories, teachings, and traditions of the culture. The same would seem to apply at least to some extent also to modern nature writing, in which the written texts and the natural environment connect and intertwine in complicated ways. Relying on the use of the concept of text in the Tartu-Moscow semiotic school, I have proposed the concept of “nature-text” as one way to integrate in a single research model representations of nature in culture and nature in its own semiotic activity.48 “Nature-text” refers to the unit (p. 269) that is formed through meaning relations between the written text that speaks about nature and points to nature and the depicted part of the natural environment itself. Such interaction can significantly shape possible interpretations of the text, especially in cases when sign relations with the local environment are more intense than cultural meanings. It is remarkable that in the case of “nature-text,” the written text does not need to convey all meanings, as they are present in the environment and familiar to the reader. Think for instance about geographies or climate conditions of a particular place: quite often nature essays leave these unspecified, assuming that reader has some experience of the place. Pointing to them is often enough, and the gaps in the fabric of the text may be as important as the explicitly expressed meanings.49 Such claims could also be supported by Gregory Bateson’s observations on the redundancy present on different levels between a unit of meaning (text) and its surroundings that together constitute a cybernetic system regulated by redundancy restraints and feedback loops.50 In such a case, the reading of a cultural text becomes a doubly interpretative activity, where the written text is read in relation to personal experience of the environment.

An understanding of the interwovenness of the natural environment and symbolic or literary expressions may also rise at a higher level of complexity of culture. For instance, Alfred K. Siewers, a medievalist at Bucknell University, has found use for the semiotic approach in his studies of Celtic literature.51 He has introduced the concept of eco-semiosphere to describe a fantasy world (for instance Celtic Otherworld) or culture that is closely and reciprocally related to an eco-region of the Earth, each forming the other in a locality. In eco-semiosphere, there is a rich engagement of cultural narrative in an overlap between regional semiosphere and biosphere. The concept itself is adapted from Juri Lotman’s “semiosphere,” but with an emphasis on the role of physical landscape and geography. It is remarkable that Lotman, in his turn, derived the concept of semiosphere from a biological source—from Vladimir Vernadsky’s “biosphere,” and the original meaning of both concepts emphasizes the place for life or semiosis that is inhabited but at the same time influenced by the living or semiotic activities. In other words, a semiosphere is wider than just a collection of written texts, and it extends as far as any possible influences of semiotic processes reach in the world and perhaps also as far as any possible influences of the world reach in the texts.

These different attempts to propose new research models appear to have some common traits: the role of the nonhuman environment is emphasized, the environment is considered in most cases to be an active and dynamical player, and human cultural phenomena are regarded as open to the environment or as intertwined with the environment. Such approach does not attempt to hierarchize the nonhuman environment, human experience, and representations, but rather regards these as a complex bounded together by sign relations. Accordingly, biosemiotic criticism can also be understood as a truly ecological approach towards literary texts, in the sense that it tries to consider the texts themselves in their creation and interpretation in the context of a wider environment that is not only textual or cultural but includes also animate subjects and the physical realm as well as interpreters and protagonists as embodied biological creatures with their own environmental relations and being.

(p. 270) Acknowledgments

The research has been supported by the European Union through the European Regional Development Fund (Centre of Excellence CECT, Estonia), by Estonian Science Foundation Grant No. 7790.

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                                                                                                                    Notes:

                                                                                                                    (1) . W. John Coletta, “Literary Biosemiotics and the Postmodern Ecology of John Clare,” Semiotica 127, no. 1–4 (1999).

                                                                                                                    (2) . Charles Sanders Peirce, The Collected Papers of Charles Sanders Peirce (Cambridge: Harvard University Press, 1994), CP 2.228.

                                                                                                                    (3) . Charles Sanders Peirce, The Essential Peirce. Selected Philosophical Writings II, ed. Nathan Houser, et al. (Bloomington, IN: Indiana University Press, 1998), EP 2.411.

                                                                                                                    (4) . Jakob von Uexküll, “The Theory of Meaning,” Semiotica 42, no. 1 (1982); Jakob von Uexküll, “A Stroll through the Worlds of Animals and Men: A Picture Book of Invisible Worlds,” Semiotica 89, no. 4 (1992).

                                                                                                                    (5) . Thomas A. Sebeok, “Communication” in Thomas A. Sebeok. A Sign is Just a Sign (Bloomington, IN: Indiana University Press, 1991), 22.

                                                                                                                    (6) . Thomas A. Sebeok, “Zoosemiotics: At the Intersection of Nature and Culture,” in Thomas A. Sebeok.Essays in Zoosemiotics (= Monograph Series of the TSC 5) (Toronto: Toronto Semiotic Circle; Victoria College in the University of Toronto, 1990), 47.

                                                                                                                    (7) . In the case of interest for more specific topics, see Springer’s book series and journal both under the title “Biosemiotics”, such as Donald Favareau, ed., Essential Readings in Biosemiotics. Anthology and Commentary (Dordrecht: Springer, 2009).

                                                                                                                    (8) . For an introduction to these principles, see Jesper Hoffmeyer. Biosemiotics: An Examination into the Signs of Life and the Life of Signs, Donald Favareau, ed. (Scranton, PA: University of Scranton Press, 2008), esp. ch. 2, 4, 7.

                                                                                                                    (9) . For various interpretations in biosemiotics, see Myrdene Anderson, “Biology and Semiotics,” in Semiotics in the Individual Sciences, ed. W. A. Koch (Bochum: Brockmeyer, 1990); Claus Emmeche, Kalevi Kull, and Frederik Stjernfelt, Reading Hoffmeyer, Rethinking Biology (=Tartu Semiotics Library 3) (Tartu: Tartu University Press, 2002); Frederik Stjernfelt, “Tractatus Hoffmeyerensis: Biosemiotics as Expressed in 22 Basic Hypotheses,” Sign Systems Studies 30, no. 1 (2002); Kalevi Kull, Claus Emmeche, and Donald Favareau, “Biosemiotic Questions,” Biosemiotics 1, no. 1 (2008); Kalevi Kull et al., “Theses on Biosemiotics: Prolegomena to a Theoretical Biology.” Biological Theory: Integrating Development, Evolution, and Cognition 4, no. 2 (2009); and Kaveli Kull and Claus Emmeche, eds, Towards a Semiotic Biology. Life Is the Action of Signs. (Singapore: World Scientific, 2011).

                                                                                                                    (10) . See Donald Favareau, “The Evolutionary History of Biosemiotics,” in Introduction to Biosemiotics: The New Biological Synthesis, ed. Marcello Barbieri (Berlin: Springer, 2006); Timo Maran, “Why Was Thomas A. Sebeok Not a Cognitive Ethologist? From ‘Animal Mind’ to ‘Semiotic Self,’” Biosemiotics 3 no. 3, 315–29 (2010); Kalevi Kull, “Biosemiotics in the Twentieth Century: A View from Biology,” Semiotica 127, no. 1/4 (1999); and Marcello Barbieri, “A Short History of Biosemiotics,” Biosemiotics 2, no.2 (2009).

                                                                                                                    (11) . Timo Maran, Dario Martinelli, and Aleksei Turovski, “Introduction,” in Readings in Zoosemiotics, ed. Timo Maran, Dario Martinelli and Aleksei Turovski (Berlin: Mouton de Gruyter, 2011. In press).

                                                                                                                    (12) . Winfred Nöth, “Ökosemiotik,” Zeitschrift für Semiotik 18, no. 1 (1996); and Kalevi Kull, “Semiotic Ecology: Different Natures in the Semiosphere,” Sign Systems Studies 26 (1998).

                                                                                                                    (13) . I refer here mostly to questions of continuity or discontinuity of mind and matter, humans and other animals, and culture and nature.

                                                                                                                    (14) . See, for example, Robert Storey, Mimesis and the Human Animal: On the Biogenetic Foundations of Literary Representation (Evanston, IL: Northwestern University Press, 1996); and Joseph Carroll, Literary Darwinism: Evolution, Human Nature, and Literature (New York: Routledge, 2004).

                                                                                                                    (15) . See, for example, Wendy Wheeler. The Whole Creature: Complexity, Biosemiotics and the Evolution of Culture (London: Lawrence and Wishart, 2006); and Terrence W. Deacon, The Symbolic Species. The Co-Evolution of Language and the Brain (New York: W. W. Norton, 1997).

                                                                                                                    (16) . The essential connection between the biological organization and language in humans has been argued by cognitive linguistics to be embedded in conceptual metaphors (for up–down organization of so-called orientational metaphors: good–bad, life–death, more–less, and so on. See George Lakoff and Mark Johnson, Metaphors We Live By (London: The University of Chicago Press, 1980), 1522.

                                                                                                                    (17) . Konrad Lorenz, “Die angeborenen Formen möglicher Erfahrung,” Zeitschrift für Tierpsychologie 5, no. 2 (1943), 278.

                                                                                                                    (18) . Thomas A. Sebeok, “Prefigurements of Art,” Semiotica 27, no. 1–3 (1979); Dario Martinelli, Zoosemiotics.Proposals for a Handbook (= Acta Semiotica Fennica 26) (Imatra: Finnish Network University of Semiotics; Imatra: International Semiotics Institute; Helsinki: Semiotic Society of Finland, 2007).

                                                                                                                    (19) . Charles F. Hockett, “Logical Considerations in the Study of Animal Communication,” in Animal Sounds and Communication, ed. Wesley E. Lanyon and William N. Tavolga (Washington: American Institute of Biological Sciences, 1960).

                                                                                                                    (20) . William H. Thorpe, “The Learning of Song Patterns by Birds, with Especial Reference to the Song of the Chaffinch Fringilla Coelebs,” Ibis 100, no. 4 (1958).

                                                                                                                    (21) . Thomas A. Sebeok, “In What Sense Is Language a ‘Primary Modeling System’?” in Thomas A. Sebeok, A Sign Is Just a Sign, 49–58. Bloomington, IN: Indiana University Press, 1991.

                                                                                                                    (22) . See Thomas A. Sebeok and Marcel Danesi, The Forms of Meaning: Modeling Systems Theory and Semiotic Analysis (Berlin: Mouton de Gruyter, 2000), but also Juri M. Lotman, “Primary and Secondary Communication-Modeling Systems,” in Soviet Semiotics: An Anthology, transl. and ed., D. P. Lucid (Baltimore: The Johns Hopkins University Press, 1977) for possible elaborations of modeling systems theory.

                                                                                                                    (23) . Alf Hornborg, “Vital Signs: An Ecosemiotic Perspective on the Human ecology of Amazonia,” Sign Systems Studies 29, no. 1 (2001): 128.

                                                                                                                    (24) . Jesper Hoffmeyer, Signs of Meaning in the Universe. Transl. B. J. Haveland (Bloomington, IN: Indiana University Press, 1996): 83–88, 113–120.

                                                                                                                    (25) . Thomas A. Sebeok, “Tell Me, Where is Fancy Bred?” The Biosemiotic Self,” In Thomas A. Sebeok, Global Semiotics (Bloomington: Indiana University Press, 2001), 124.

                                                                                                                    (27) . See Andrew Stables, Living and Learning as Semiotic Engagement. A New Theory of Education (Lewiston: The Edwin Mellen Press, 2005); and Wendy Wheeler, The Whole Creature.

                                                                                                                    (28) . Winfred Nöth, “Ecosemiotics and the Semiotics of Nature,” Sign Systems Studies 29, no. 1 (2001): 72.

                                                                                                                    (29) . James J. Gibson, The Ecological Approach to Visual Perception (Hillsdale, NJ: Lawrence Erlbaum, 1986).

                                                                                                                    (30) . Michael Polanyi, Personal Knowledge. Towards a Post-Critical Philosophy (London: Routledge, 1962).

                                                                                                                    (31) . John Deely, Four Ages of Understanding: The First Postmodern Survey of Philosophy from Ancient Times to the Turn of the Twenty-First Century (Toronto: University of Toronto Press, 2001).

                                                                                                                    (32) . Kadri Tüür, “Bird Sounds in Nature Writing: Human Perspective on Animal Communication,” Sign Systems Studies 37 no. 3/4 (2009).

                                                                                                                    (33) . Here, motivatedness should be understood as limited (because of constrained relations of object-sign-interpretant) possibilities for interpretation, and intentionality as directedness of semiosis because of these limitations.

                                                                                                                    (34) . Charles Sanders Peirce, Semiotic and Significs: The Correspondence between Charles S. Peirce and Victoria Lady Welby, ed. Charles S. Hardwick and J. Cook (Bloomington, IN: Indiana University Press, 1977), 80–81.

                                                                                                                    (35) . Metaphors and onomatopoetic words can be considered iconic to the extent that the perception of the relation to their referent is based on similarity.

                                                                                                                    (36) . Jesper Hoffmeyer, Biosemiotics: An Examination into the Signs of Life and the Life of Signs, ed. Donald Favareau (Scranton, PA: University of Scranton Press, 2008), 189.

                                                                                                                    (37) . Roman Jakobson, “Linguistics in Relation to Other Sciences,” in Roman Jakobson. Selected Writings II. Word and Language (The Hague: Mouton, 1971), 677–81.

                                                                                                                    (38) . Anton Markoš, Readers of the Book of Life: Conceptualizing Developmental Evolutionary Biology (Oxford: Oxford University Press, 2002); and Anton Markoš, Filip Grygar, László Hajnal, Karel Kleisner, Zdenek Kratochvíl, and Zdenek Neubauer, Life as Its Own Designer: Darwin’s Origin and Western Thought (Dordrecht: Springer, 2009).

                                                                                                                    (39) . Coletta, “Literary Biosemiotics”; W. John Coletta, “Predation as Predication: Toward an Ecology of Semiosis and Syntax,” Semiotica 109, no. 3–4 (1996); W. John. Coletta, “The Semiosis of Nature: Towards an Ecology of Metaphor and a Biology of Mathematics,” The American Journal of Semiotics 10, no. 3–4 (1993).

                                                                                                                    (44) . See, for example, Wendy Wheeler, “Postscript on Biosemiotics: Reading beyond Words—and Ecocriticism,” New Formations 64 (2008): 137–54; Coletta, “Literary Biosemiotics.”

                                                                                                                    (45) . Andrew Stables, “The Landscape and the ‘Death of the Author,’” Canadian Journal of Environmental Education 2, no. 1 (1997).

                                                                                                                    (46) . Anne W. Spirn, The Language of Landscape (New Haven: Yale University Press, 1998), 15.

                                                                                                                    (47) . David Abram, The Spell of the Sensuous. Perception and Language in a More-than-Human World (New York: Vintage Book, 1996).

                                                                                                                    (48) . Timo Maran, “Towards an Integrated Methodology of Ecosemiotics: The Concept of Nature-Text,” Sign Systems Studies 35, no. 1/2 (2007).

                                                                                                                    (50) . Gregory Bateson, Steps to an Ecology of Mind (Chicago: The University of Chicago Press, 1973), 405–31.

                                                                                                                    (51) . Alfred K. Siewers, Strange Beauty. Ecocritical Approaches to Early Medieval Landscape, (New York: Palgrave Macmillan, 2009).