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• W4313639593 abstract "Symbiosis refers to intimate relations between life forms that involve strong interdependence. Some well-known examples are the associations between flowers and bees, fungi and tree roots, the lichen symbiosis between fungi and algae, and even the mitochondria that make chemical energy for eukaryotic cells. In this list, the examples go from relatively loose associations to ones that are ever more tightly coupled. While the flower and the bee are interdependent products of co-evolution, they are still distinct and distinguishable organisms. For some biologists, including Lynn Margulis and Dorion Sagan (2002, 18), they do not count as examples of symbiosis because they do not live in close or “obligate” contact. The wasp can find other sources of food; the orchid can be pollinated by other species. But mycorrhizae, lichens, and mitochondria are much more integrated. Lichens, the example for which biologists first used the term symbiosis in the late 19th century (Sapp 1994, 6), look like a single organism even if their fungal and algal partners can still be observed separately under a microscope. Mitochondria are fully integrated into cells, living inside their membranes and utterly dependent on their hosts. While the word symbiosis commonly refers to mutual benefit, its more specialized meaning in biology is a continuum from mutualism to one-sided parasitism.In addition to proximity and benefit, symbiosis is about evolution. As Angela Douglas (2010) puts it, evolution as descent with modification is the dominant theory of how organisms vary and acquire new traits. The assumption of both Darwinism and Neo-Darwinism is that descent with modification is gradual. But symbiosis can be its own source of “modification” or evolutionary variation: There is unambiguous evidence that some traits of great evolutionary and ecological importance have been gained laterally from different, often phylogenetically distant, taxa. Some laterally acquired traits are novel for the recipient organism and they can be evolutionary innovations, i.e., “new acquired structures or properties which permit the assumption of a new function.” (Douglas 2010, 1) The term for evolution through symbiosis is symbiogenesis, which refers to a theory first outlined by the Russian biologist and lichenologist Konstantin Mereschkowski in papers from 1905 and 1910. The famous image of the tree of life, which fails to capture this reintegration of lineages noted by Douglas, gives way to a network. The tree does not only branch in one temporal direction, so that all branches lead back to common ancestors. The branches also fuse and reticulate like a mycelium. These two powerful images, tree and mycelium, illustrate two distinct processes of evolution (and there may be more, such as self-organization [Depew and Weber 1994], which no doubt plays a role in symbiosis as well). They are complementary, not mutually exclusive.Theoretical biology has seen many explanations of symbiosis accompanied by symptomatic figures of speech. Even though Margulis and Sagan (2002, 15–17) are quick to denounce the idea that Darwinian competition is “more natural” and less anthropomorphic than cooperation, they go on to criticize anthropomorphism in biology as such—and then to rely heavily on social figures of cooperation. And why not, when scholars have established so many times that metaphor is inseparable from scientific reasoning? But the stakes of anthropomorphism change if I am unsure where to draw the line between humans and the rest of life, whether I even should, and how widely the qualities that anthropomorphism ostensibly projects are already distributed among species. Jan Sapp (1994) details the wide range of social analogies that biologists have used to describe symbiosis, which include friendship, hospitality, neighborliness, oppression, slavery, imprisonment, love, games, cost–benefit economics, and more. As he argues, “‘commonsense’ views of human social relations” are often used “to explain relations between microbes and their hosts,” as well as between organisms of the same scale” (135).Recent explanations of symbiosis have sought to move beyond this language of human social relation that assumes a kind of origin story: Individuals and species preexist separately; only later do they evolve into close symbioses. For example, Donna Haraway (2008; 2016) points to a conflict between symbiogenesis and another concept from theoretical biology: Humberto Maturana and Francisco Varela's concept of autopoiesis, or the recursive self-production whereby life forms make themselves out of the raw materials of their environments. Autopoiesis entails not ecological closure but “operational closure” because life can be defined by “the network of processes of production of components . . . that continuously regenerate and realize the network of processes (relations) that produced them” (Maturana and Varela 1972/1980, 79). While both autopoiesis and symbiogenesis have been important for antireductionist approaches to thinking biology in systems rather than neo-Darwinian selfish genes, Haraway sees them as contradictory. For her, there can be no self-enclosed units of life. The recursive closure demanded by autopoiesis is incompatible with a biosphere composed of relations all the way down, a biosphere in which life forms are constantly enfolded and merged with one another—in other words, a biosphere in which symbiosis is so constitutive that it would be impossible to spot the chimeras and distinguish them from basic singular beings. Haraway offers “sympoiesis” as a way to move us past the impasse of autopoiesis and symbiogenesis.As an attempted synthesis of these two concepts, the problem with sympoiesis is that it provides no replacement for the bootstrapping abilities of life—abilities that are not about some kind of anti-ecological isolation from the environment, but rather present a compelling answer to the question of what makes life distinct and irreducible to the chemistry of replicators. My essay intervenes in the critical and scientific discourse about how to interpret symbiosis by arguing for an alternative resolution of the conflict between autopoiesis and symbiogenesis. I look back to work on animal artifacts (Gould and Gould 2012) and extended phenotypes (Dawkins 1982/1999) and seek to push it further. Symbiosis is a kind of prosthesis or technological process. Lichens are nonhuman technologies. The fungus is a greenhouse for the alga and the alga is a solar panel for the fungus. The alga can live in drier conditions because of the fungus; the fungus can live an autotrophic idyll rather than decomposing dead organisms as most fungi do. The two or more systems in symbiosis, according to this logic, remain autopoietic. But they are functionalized by the new lichen-like symbiotic unit.Symbiogenesis is not a matter of organisms using one another like nonliving tools or machines, but a fundamental technical process in which one autopoietic life form externalizes functions into another. Life forms in mutualism supplement one another to open new functions and relations to their environments. Over evolutionary time, however, there is an ambivalent and (to use Derrida's language) pharmacological play along the continuum between parasitism and mutualism. Symbiosis comes into relief as a technical process when we see it as unconscious, mutual prosthesis. In other words, explaining symbiosis in terms of technology is only viable through a theory of technics that rejects instrumentality and leaves us with a very different perspective on technology from that of a human individual intentionally using a tool. To understand symbiosis and especially symbiogenesis in technical terms, I argue for the value of critical and deconstructive theories of technology that think technics as irreducible to the use of objects to realize purposes intended in advance.“Prosthetic symbiosis” is both an explanation of symbiosis and a way of situating it within the theory and politics of ecology in the 21st century. In this context, symbiosis is increasingly a concept that crystalizes general ecology in an affirmative mode—a role that the word's etymology as “living-together process” certainly invites. This is what Erich Hörl (2017, 31) calls the “general ecological principle of symbiosis.” For him, symbiosis is part and parcel of “profound epistemological, media-historical, and technological-historical changes” in the concept of ecology, changes that militate toward “primordial participation as essential relation, which precedes the constitution of its terms, namely participating entities” (2017, 52). He argues that symbiosis, symbiogenesis, and sympoiesis are all crucial examples of this new general ecology. They are grist for the mill of a relationality more fundamental than any substance.There are good things about this trend toward infinite ecologies whereby any phenomenon can be framed in ecological terms, so that “eco” modifies x or y in a constitutively relational and postnatural light. One downside is that symbiosis ends up applied widely across scales and registers, as when sympoiesis, for Haraway, can be general concept relevant to ecological thought at the planetary scale. Some of the fascinating specificity of symbiosis as something life offers to ecotheory is lost or vitiated. To correct for this movement of generalization, I elaborate prosthetic symbiosis by situating it as a kind of minimal ecological relation: a threshold between ecological and biological registers and a technical process that mediates between ecological relation and biological self-relation. With this specificity, we can better understand the relation of symbiosis to the complex politics of the life sciences.In the summer of 2017, wildfires driven by global warming cut through British Columbia and the western United States. Vancouver, BC, was opaque with smoke when I heard that my friend Trevor Goward was under evacuation alert. As a radical localist, he was planning to stay in his home. But as a lichenologist, he hoped to save his thousands of dried, identified, and labeled lichens that make up western Canada's biggest research collection by moving them from the Clearwater Valley, near the center of the province, to the University of British Columbia. Years before, I had worked as a research assistant for Goward, but that story is too long to tell. The fire risk was too high for the UBC herbarium to officially send an employee to collect the specimens, but still low enough to feel safe. With a friend, I rented a moving truck and drove six hours to the interior. We picked up the lichens and delivered them to the Vancouver campus.The next day, I woke to an e-mail from CBC radio. They wanted an interview about the lichen “rescue” they had heard about on social media. They had been looking for fire stories all summer, but by this point the fires had been burning for so long that they seemed to want content other than livelihoods destroyed and experts who could patiently link the weather to global warming. The CBC wanted to know why these lichens needed rescuing, why Goward's collection mattered, and what lichens even are. During the interview, I explained the trip and the lichen collection. I wanted to say something more philosophical as well: some comment about the theories I care about, despite the odds it would please the interviewer. What came out was that lichens are more than one life form among others. They are special because of their status as symbiotic organisms. They are symbols of future coexistence.Indeed, lichens have an ecotopian meaning that has attracted some cultural attention in recent years (i.e., Robinson 1992; Gilbert et al., 2012; Griffiths 2015; Haraway 2016; Milne 2017; Grantina 2020; Sheldrake 2020; Palmer 2023), lifting them from obscurity and the injustice of being mistaken for moss. But will symbiosis save us from Earth system mutation? Does mutual aid point the way for life forms to coexist in finitude? For many, the answer is yes. In a course I taught about environmental literature in 2021, my students arrived with the view that symbiosis is a core principle for deciding what counts as green utopian and what does not. I had not taught them the term. Their ability to use it suggests that formulas like “we have upset the delicate symbiosis between ourselves and nature” are broadly legible (Pindar and Sutton in Guattari 1989/2019, 4). Symbiosis between organisms, between humans and nature, between cities and ecosystems, among trees in the forest: All seem ways of living in mutual benefit and reciprocity rather than selfishness and harm.The full spectrum of political values has been applied to this idea of natural mutualism. From a right-environmentalist perspective, humans need to stop being parasites on Gaia and learn to live in reciprocity with our environment in case the earth decides to manage our population in its own vengeful way. From a centrist point of view, symbiosis is an idea that mediates technocratic goals of energy efficiency, reduced emissions, or waste management. For anarchists and Marxists (Kropotkin 1902; Milne 2017; Morton 2017), symbiosis is a principle of solidarity with other species, or a theory of evolution that naturalizes cooperation rather than competitive individualism and self-organizing markets. For posthumanists, symbiosis is a project of abandoning the human exception; our future is like the future of Octavia Butler's Xenogenesis novels (1989/2000): to merge with other species in new ways—and to do so while realizing that we were never strictly human in the first place, but rather messy, lichen-like consortia of species (Clarke 2008). These interpretations of symbiosis have been argued in both general and specialist texts and across the deep, persistent split between the “two cultures” of the arts and sciences.When contemporary ecological theorists draw upon the discourse of symbiosis in its trajectory from Simon Schwedener and Albert Bernhard Frank's work on lichens in the 1860s, they often use it as raw material for an affirmative biopolitics. Symbiosis becomes a general condition of the biosphere that humans, in our delusions of autonomy, tend to deny. Symbiosis becomes a normative example of the form of relation we should aspire to or mimic.Famously, Donna Haraway's late work reads microbiologist Lynn Margulis toward a posthumanism grounded in symbiotic entanglement. But not the entanglement of previously separate beings: symbiosis goes all the way down in evolutionary time; we were never individuals on the first place; and all eukaryotic life forms are constituted by relations with other life. This is knowledge of how things are, but also a political desire for how they could be made better through what Haraway (2016) calls “the layered, curious practice of becoming with others for a habitable, flourishing world” (168). Her reading of Margulis has been influential, echoing across a wide range of posthumanist work and finding its way into science writing such as Merlin Sheldrake's Entangled Life: How Fungi Make Our Worlds, Change Our Minds, and Shape Our Futures (2020). Haraway's (2016) understanding of symbiosis also shifts toward the planetary concerns of climate change and the Anthropocene. Her alternative, the Chthulucene, asks readers to learn how to live with our inevitably decentered position in the biosphere. Through symbiosis and by making queer kin with nonhumans, we can give “a fierce reply to the dictates of both Anthropos and Capital” (2). In this way, the small scale of intimate symbiotic relation ascends to the pantheon of planetary ecopolitics.Haraway is only one of the highest profile scholars to make hay with symbiosis as an anticapitalist alternative to neoliberal Darwinism and ecologies of the invisible hand. Other uses of symbiosis include Luciana Parisi's (2004) posthuman, Deleuzean, bacteria-inspired theory of sex and Bruce Clarke's (2020) exhaustive study of Margulis's role in the emergence of Gaia theory throughout the late 20th century. Writing about eco-Marxism and its need to incorporate “the symbiotic real” and “solidarity with nonhuman people,” Timothy Morton (2017) claims that communism can only work at the planetary scale, which aligns him with Haraway's Chthulucene (1). For Morton, “humankind” does not mean our species in the hypostatized anthropological sense of the term but rather “an ecological being that can be found in the symbiotic real” (3). Building on Peter Kropotkin's Mutual Aid: A Factor of Evolution (1902), an anarchist response to Darwin's On the Origin of Species (1859), Morton does not make the predictable gesture of embracing symbiotic cooperation, but rather argues for the need to “deconstruct the selfishness-altruism binary” (171). They draw on Kropotkin's comparison between symbiosis and neighbor, the idea that neighbors will help each other put out a fire even if they lack kinship and friendship: “Solidarity is the noise that the symbiotic real makes . . . . the zero-degree, cheapest coexistence mode, something you rely on when all else fails” (174). Such examples suggest that life has an instinctual tendency toward help and generosity, one that should not be understood sentimentally but rather as a widespread unconscious process.Despite this ongoing cultural work, the scope and significance of symbiosis remain controversial in the sciences. Symbiogenesis is often put forward as a new and revolutionary idea despite its long history. Over the past 10 years, the most viral example has been the wood wide web. The work of its biggest proponent, the scientist Suzanne Simard, has given symbiosis new purchase on the public imagination (i.e. Macfarlane 2016). “Wood wide web” refers to networks of fungi, or mycelium, that grow beneath the soil and connect the roots of plants. This symbiosis was originally thought to be a mutualism between plants and fungi: In exchange for energy generated by photosynthesis, these “mycorrhizal” fungi extend the roots of plants to an exquisite degree of ramification, increasing their ability to absorb nutrients. But Simard (1997; 2021) argues that the mycorrhizal relationship is about more than extended roots. These mycelia also form expansive, underground networks that connect multiple species of plants and fungi. Even trees of different species (in Simard's original study they were birch and Douglas fir) seem to share food and communicate with one another. Mycorrhizal fungi are thus being rethought, often in the fraught speculative frame of plant sociality and sentience. The wood wide web also imbues symbiosis with digital utopianism and the troubled legacies (Thacker and Galloway 2007; Neyrat 2016/2019; Woods 2022) of networks and distributed agency. While such examples have an extraordinary cultural cachet that calls for separate discussion (see Nixon 2021), biologists are far from consensus about the scope and significance of symbiosis as either an ecological or evolutionary mechanism.The theoretical stakes are high. Many of the same scholars who argue that there is no nature opposed to society would do away with the idea of individual organic integrity. For Scott F. Gilbert, Jan Sapp, and Alfred I. Tauber (2012), symbiosis is one among several aspects of 21st-century biology that show that “we have never been individuals”—and, better, that “we are all lichens” (336). If we are all lichens, perhaps corporeal boundaries are illusions. Such arguments have by now given lichens a prominent and surprising role in the critical humanities and social sciences, where the significance of lichens and symbiosis stretches beyond resistance to reductionist genetic determinism.In “Queer Theory for Lichens” (2015), for example, David Griffiths begins by citing Gilbert, Sapp, and Tauber's line about universal lichenization. He draws on insights from symbiosis research to theorize sexuality. Individuality is Griffiths's target because it connotes a certain bodily autonomy that has been associated with binary gender essentialism and ways of policing sexuality to maintain bodily purity. Symbiosis here becomes another way to disrupt the normalization of “sexual reproduction and vertical inheritance at the expense of many other forms of production and reproduction” (36). Because examples such as lichens are “natural” and widespread, they make it easy to question how straight the nature that queer and trans sexualities are often attacked as against was in the first place. Nature's unnatural reproductions affirm the critique of heteronormativity and offer new ways of “denaturalizing the primacy of heterosexuality and sexual reproduction in defining and legitimating bodies, practices, and communities” (36). In queer ecology, we are always already lichens because this term abstracts to the idea that all eukaryote bodies are symbiotic, multispecies communities.If I understand bodies in this way, then the question becomes what to do with a feature that finite bodies do seem to have: their provisional, relative, and permeable boundaries. Griffiths is right to say that examples like symbiogenesis militate against the idea that biological nature is nothing more than closed bodies and vertical lineages. But if nature has always been queer in this broad sense, this does not explain why so many societies have come to repress this fact and enforce or normalize heteronormativity. People who identify as queer might ask what their desires and relationships with others have to do with nonhuman symbioses, and whether their lives need any kind of biological justification in the first place. The idea that we are all lichens does useful work by making nature strange and denaturalizing widely held ideologies of sex. But we may demand too much from symbiosis by turning it into a general philosophy of nature. When it comes to understanding what nonhuman life means for topics like culturally mediated sexuality, what seems to be missing is a way to talk about a dynamic movement back and forth between individuality and symbiosis, relative closure and relative openness, the negativity of distance as well as the affirmation of relation. This gap speaks to the downside, as critics respond to reductive biological essentialism, of elevating symbiosis while leaving aside the conceptual affordances of autopoiesis.To move forward with symbiosis in critical theory, we should avoid overstating its ethical and political potential by moving too quickly from biological observation to normative value. For example, there may be scales where symbiosis matters and scales where it does not. Cats who depend on microbes in their digestive tract can playfully torture mice to death. There may also be “symbiotic” processes that work against themselves. A tree can share nutrients, via mycorrhizae, with the same seedlings that die from its shade. It is true that individuality is no absolute boundary, and as Margulis and Sagan (2002) point out, traditional zoocentric concepts of the body's individuality and autonomy have been applied far too widely (21). But even if it is difficult to demarcate the boundaries of individuals, they remain individuals to the extent that they can die. Even if species are really multispecies assemblages, fuzzily bounded “species” continue to exist insofar as they can go extinct. From an ecological perspective, death is integral to function of the Earth system. The biosphere is also a necrosphere. As Jonathan Basile argues (2021) in “Symbioautothanatosis,” this ambivalence of symbiosis needs more attention. It shows the need for caution when we affirm symbiosis for a politics of coexistence or make it scalable for planetary justice. With the concept of prosthetic symbiosis, I mean to embrace this ambivalence by thinking symbiosis through the logic of the pharmakon: as both “poison and cure” and, beyond this overused gloss, as a dynamic of functional supplementarity and externalized memory.Along with the time I spent working as a research assistant for Trevor Goward, one starting point for my work on symbiosis has been the conflict in theoretical biology I mentioned at the beginning of this essay. To return to Haraway (2008, 32–33; 2016, 33), there is a contradiction between the concepts of autopoiesis and symbiogenesis: there can be no self-organization without selves, no self-production in a biosphere so thoroughly symbiogenetic that units are unable to evolve to be separate. The story that has separate organisms “choosing” symbiosis for reasons of game-theoretic benefit is moot without initially separate units.Haraway's attempt to radicalize Margulis by dropping autopoiesis leads her to the concept of “sympoiesis.” Sympoiesis is meant to “enfold, unfurl, and extend autopoiesis” (2016, 58), but drop its recursive closure in favor of “collectively-produced systems that do not have self-defined spatial or temporal boundaries” (33). As Bruce Clarke and Scott Gilbert (2022, forthcoming) have argued, there are good reasons to see autopoiesis and symbiogenesis as compatible; Margulis herself “observed no contradiction” between them (7). While I am in broad agreement with these authors, my concern is that Sagan and Margulis's (1991) language of “merger” and “integration” that leads to “autopoietic entities of still greater complexity” glosses over something crucial about how it is possible for the fundamentally internal self-relation of autopoiesis to relay “itself” through another system (368–69).Prosthetic symbiosis is an alternative way to address the apparent conflict between autopoiesis and symbiogenesis, one that embraces recursive autopoietic closure but deconstructs Maturana and Varela's reliance on systemic unity. Perhaps symbiosis and symbiogenesis are neither generalized webs of reciprocity and interdependence nor the merger of previous separate species but instead a primordial form of technics. Instead of merging, life forms “use” one another as technologies. This argument is not entirely new. There is already a theory of symbiosis as technology implicit in the metaphors of a small group of critical theorists and biologists, as when Margulis and Sagan (2002) describe bacteria as the first genetic engineers (41; see also Clarke 2022 on a different kind of “natural technicity”).For his part, Sheldrake (2020) evokes the wood wide web with a crucial chiasmic metaphor. In the mycorrhizal symbiosis, “plants gain a prosthetic fungus, and fungi gain a prosthetic plant” (125). As he writes elsewhere in the same book, “both use the other to extend their reach” (125). In other examples, he repeats the term prosthesis to describe how fungi take over the bodies of ants (96) and the “prosthetic metabolisms” at work when termites cultivate fungi in their mounds to digest cellulose (190). In so doing, Sheldrake grafts his argument to Ernst Kapp (1877/2018) and Marshall McLuhan's (1964/1994) famous theories of media as extensions of the human body. Sheldrake's metaphor can also be placed in the tradition of theorizing technical prosthesis that passes through the work of Martin Heidegger, Gilbert Simondon, Jacques Derrida, Bernard Stiegler, David Wills, and others.Where Sheldrake uses prosthesis as an occasional metaphor, I hope to lay the conceptual groundwork for a theory of symbiosis as nonhuman technics. It could be that the theory of symbiosis that has received the least attention explains it best: as a radicalized form of technology, as biotechnics, or as a variation on what Eugene Thacker calls biomedia (2004). Prosthesis is no metaphor but actually how symbiosis works.Species use one another as tools to open new relationships to their milieus. They extend their functions, or add new ones, through their relationships with other species. What these simplifications reach to describe in everyday language is something that cannot be described accurately without concepts such language struggles to capture. The concept of prosthetic symbiosis can only be viable through a process of critique and revision that moves past the problems inherent in understanding any technical relation in terms of intentional tool use.Thinking in terms of technics rather than technology is shorthand for my argument that the theory of prosthetic symbiosis should work with critical theories of technology that avoid the logic of instrumentality but retain prosthesis. As Stiegler (1994/1998) writes early in his Technics and Time project, “technics constitutes a system to the extent that it cannot be understood as a means—as in Saussure the evolution of language, which forms a system of extreme complexity, escapes the will of those who speak it” (24). Even in the relation that forms between fungi and algae or the biochemical signaling of bacteria (which I could not confidently compare to the complexity of language), even in symbiosis between “simple” life forms, there is a technical system that cannot be understood as a means to an end. If this is a negative definition that only says what technics is not, it has the advantage of reminding us that symbiosis thought as technics is irreducible to instrumentality. By distinguishing a system that cannot be understood as means, Stiegler demarcates the aspect of technics that needs to be rethought.The problem with instrumentality is about more than general concepts of technology. With its implication of slavery, the image of one species using another as a tool is wrong for understanding the technical character of symbiosis. In a discussion of parasitism and symbiogenesis in the work of Octavia Butler and the history of science, Zakiyyah Iman Jackson (2020) points out that the lichen symbiosis was seen by Simon Schwedener, when he first argued that lichens are dual organisms, as the enslavement of algae by fungi. Jackson points out that Margulis, Sagan, and Haraway's recourse to metaphors of “mutual enslavement” and “miscegenat" @default.
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