Nash equilibria for an evolutionary language game

We study an evolutionary language game that describes how signals become associated with meaning. In our context, a language, L, is described by two matrices: the P matrix contains the probabilities that for a speaker certain objects are associated with certain signals, while the Q matrix contains the probabilities that for a listener certain signals are associated with certain objects. We define the payoff in our evolutionary language game as the total amount of information exchanged between two individuals. We give a formal classification of all languages, L(P, Q), describing the conditions for Nash equilibria and evolutionarily stable strategies (ESS). We describe an algorithm for generating all languages that are Nash equilibria. Finally, we show that starting from any random language, there exists an evolutionary trajectory using selection and neutral drift that ends up with a strategy that is a strict Nash equilibrium (or very close to a strict Nash equilibrium). Received: 1 March 2000 / Published online: 3 August 2000     

The evolutionary language game.

We explore how evolutionary game dynamics have to be modified to accomodate a mathematical framework for the evolution of language. In particular, we are interested in the evolution of vocabulary, that is associations between signals and objects. We assume that successful communication contributes to biological fitness: individuals who communicate well leave more offspring. Children inherit from their parents a strategy for language learning (a language acquisition device). We consider three mechanisms whereby language is passed from one generation to the next: (i) parental learning: children learn the language of their parents; (ii) role model learning: children learn the language of individuals with a high payoff; and (iii) random learning: children learn the language of randomly chosen individuals. We show that parental and role model learning outperform random learning. Then we introduce mistakes in language learning and study how this process changes language over time. Mistakes increase the overall efficacy of parental and role model learning: in a world with errors evolutionary adaptation is more efficient. Our model also provides a simple explanation why homonomy is common while synonymy is rare.     

Yawning: An evolutionary perspective

Yawning is a ubiquitous activity among humans, nonhuman primates, mammals, birds and other vertebrates. Comparative analysis suggests that yawning has two major features: (1) communication — whereby the behavior of other individuals is affected, and (2) direct physiological benefit — whereby the organism is receiving some direct physiological benefit from yawning. Various functional hypotheses used to explain yawning in an evolutionary context are reviewed. The contagious nature in humans and the manifest lack of contagion in other species suggests that yawning in humans has a different and as yet poorly understood evolutionary history.     

A relationship between GC content and coding-sequence length

Since base composition of translational stop codons (TAG, TAA, and TGA) is biased toward a low G+C content, a differential density for these termination signals is expected in random DNA sequences of different base compositions. The expected length of reading frames (DNA segments of sense codons flanked by in-phase stop codons) in random sequences is thus a function of GC content. The analysis of DNA sequences from several genome databases stratified according to GC content reveals that the longest coding sequences—exons in vertebrates and genes in prokaryotes—are GC-rich, while the shortest ones are GC-poor. Exon lengthening in GC-rich vertebrate regions does not result, however, in longer vertebrate proteins, perhaps because of the lower number of exons in the genes located in these regions. The effects on coding-sequence lengths constitute a new evolutionary meaning for compositional variations in DNA GC content. Correspondence to: J. L. Oliver     

Firefly Femmes Fatales: A Case Study in the Semiotics of Deception

Mimicry and deception are two important issues in studies about animal communication. The reliability of animal signs and the problem of the benefits of deceiving in sign exchanges are interesting topics in the evolution of communication. In this paper, we intend to contribute to an understanding of deception by studying the case of aggressive signal mimicry in fireflies, investigated by James Lloyd. Firefly femmes fatales are specialized in mimicking the mating signals of other species of fireflies with the purpose of attracting responding males to become their prey. These aggressive mimics are a major factor in the survival and reproduction of both prey and predator. It is a case of deception through active falsification of information that leads to efficient predation by femmes fatales fireflies and triggered evolutionary processes in their preys’ communicative behaviors. There are even nested coevolutionary interactions between these fireflies, leading to a remarkable system of deceptive and counterdeceptive signaling behaviors. We develop here a semiotic model of firefly deception and also consider ideas advanced by Lloyd about the evolution of communication, acknowledging that deception can be part of the explanation of why communication evolves towards increasing complexity. Increasingly complex sign exchanges between fireflies evolve in an extremely slow pace. Even if deceptive maneuvers are played out time and time again between particular firefly individuals, the evolution of the next level of complexity—and thus the next utterance in the dialogue between species—is likely to take an immense amount of generations.     

Indirect reciprocity and the evolution of “moral signals”

Signals regarding the behavior of others are an essential element of human moral systems and there are important evolutionary connections between language and large-scale cooperation. In particular, social communication may be required for the reputation tracking needed to stabilize indirect reciprocity. Additionally, scholars have suggested that the benefits of indirect reciprocity may have been important for the evolution of language and that social signals may have coevolved with large-scale cooperation. This paper investigates the possibility of such a coevolution. Using the tools of evolutionary game theory, we present a model that incorporates primitive “moral signaling” into a simple setting of indirect reciprocity. This model reveals some potential difficulties for the evolution of “moral signals.” We find that it is possible for “moral signals” to evolve alongside indirect reciprocity, but without some external pressure aiding the evolution of a signaling system, such a coevolution is unlikely.     

Autocommunication and Perceptual Markers in Landscape: Japanese Examples

Juri Lotman distinguishes between two main types of communication. In addition to the classical I-YOU communication, he speaks about I-I communication, where both the addresser and the addressee are one and the same person. Contrary to how it sounds, autocommunication is not self-sufficient musing inside one’s self, it is remodelling oneself through a code from an entity outside oneself, be it animate or inanimate. According to Lotman, it is often the rhythmical phenomena like poetry, the rhythm of waves, etc. that lend themselves for the act of autocommunication as external codes. After having received the message one is not identical to the original oneself anymore. Perceptual markers of landscape—specific rhythms, ephemera, the rhythm of human everyday activities, bodily movement—can be considered as a secondary code leading to autocommunication in the person who contemplates the landscape. Looking at the landscape—which also implies the rhythmical movement of the eyes—one uses it as a code to reconstitute oneself. A person who has confronted a landscape does not leave it as the same person. The present article poses a definition of autocommunication in landscapes and discusses the way in which other sensorial information apart from the visual—smell, movement, rhythms etc—are used culturally to reinforce autocommunication with oneself. It can be said that several institutionalised religious and cultural practices expect the subject to reconstitute him- or herself mainly through the bodily landscape experience.     

Indiscriminate polyandry and male parental effort

Extrapair paternity involves cooperation between mated females and extrapair males. On the other hand, mated males exhibit a spectrum of anti-cuckolding strategies. Hence, extrapair attributes of diverse species and populations reported in the literature are particular solutions of evolutionary games involving gender-specific cuckolding/anti-cuckolding strategies. Here we use game theoretical methods to study the effect of male paternal effort conserving strategies in situations where females seek extrapair fertilizations (EPF) for reasons of genetic compatibility and/or in pursuit of genetic diversity for their offspring. In such cases, females cannot make a pre-copulatory selection of the optimal genetic partners, and therefore combine promiscuous copulation with the use of in copula and/or post-copulatory selection mechanisms to optimize the genetic endowment of their offspring—indiscriminate polyandry. Our results indicate that, when indiscriminate polygamy is constrained by the availability of extrapair male partners, there are three possible (parameter regime wise) evolutionary stable strategy solutions. (1) All females seek EPF, while all males restrict parental care. (2) All females seek EPF, while all males are unconditionally parental. (3) Females use a combination strategy where pursuit of EPF is mixed—on either a population, or an individual level—with genetic monogamy, while all males use a conditional paternal care strategy, which involves adjusting their parental efforts according to their certainty of paternity.     

Left-hand Preference for a Complex Manual Task in a Population of Wild Siamangs (<Emphasis Type="BoldItalic">Symphalangus syndactylus</Emphasis>)

Many hypotheses have been proposed linking the emergence of lateralized activities—such as handedness—in primates, with hemispheric specialization and the evolution of complex communication such as human language. Although data to test these ideas are rapidly accumulating for many primate taxa, some species are still largely unexplored, especially under natural conditions. I present the first data on a population of wild siamangs (Symphalangus syndactylus), highly arboreal small apes. Preference for the left hand is shown both at the individual and population levels for a complex manual task: collecting and drinking water from tree holes. There was no difference in hand preference between males and females, and immature individuals showed more variable patterns than adults. These results are consistent with the postural origins theory, allow a new interpretation of the findings of comparable studies, and indicate a useful behavior for future investigations of laterality in wild primates.     

A stochastic model for the synthesis and degradation of natural organic matter part II: molecular property distributions

A stochastic biogeochemical model has been developed to simulate the transformation and degradation of natural organic matter (NOM) using an agent-based algorithm which treats each molecule as a separate and potentially unique entity. Molecules react when a pseudo-random number is lower than the calculated reaction probability in a given time step; repeated time steps simulate the transformation of precursor molecules into a complex NOM assemblage. The data for each molecule—elemental and functional group composition—can be used to calculate many properties directly and exactly for each molecule in the assemblage, e.g., molecular weight (MW), fraction of aromatic C (Ar), and charge at pH 7 (Z). Empirical quantitative structure activity relationships (QSARs) are developed which permit the estimation of thermodynamic quantities K _ow (the octanol–water partition coefficient) and pK _a (acidity) for each molecule. Root mean square errors for these QSARs are 0.39 log units for log K _ow and 0.45 log units for pK _a. Distributions of both exactly calculated (MW, Ar, Z) and estimated thermodynamic (K _ow, pK _a) properties are examined and compared with published experimental data. Molecular weight distributions from size exclusion HPLC experiments on aquatic NOM are quantitatively similar to simulation results. pH titrations and polarity distributions from reversed-phase HPLC are qualitatively similar to simulation results. This agreement suggests that the agent-based model can be used to explore hypotheses regarding both compositional and thermodynamic properties of NOM. Robert Wetzel—deceased.     

A multifaceted framework to establish the presence of meaning in non-human communication

Does non-human communication, like language, involve meaning? This question guides our focus through an interdisciplinary review of the theories and terminology used to study meaning across disciplines and species. Until now, it has been difficult to apply the concept of meaning to communication in non-humans. This is partly because of the varied approaches to the study of meaning. Additionally, while there is a scholarly acknowledgement of potential meaning in non-human cognition, there is also scepticism when the topic of communication arises. We organise some of the key literature into a coherent framework that can bridge disciplines and species, to ensure that aspects of meaning are accurately and fairly compared. We clarify the growing view in the literature that, rather than requiring multiple definitions or being split into different types, meaning is a multifaceted yet still unified concept. In so doing, we propose that meaning is an umbrella term. Meaning cannot be summed up with a short definition or list of features, but involves multiple complexities that are outlined in our framework. Specifically, three global facets are needed to describe meaning: a Signal Meaning Facet, an Interactant Meaning Facet, and a Resultant Meaning Facet. Most importantly, we show that such analyses are possible to apply as much to non-humans as to humans. We also emphasise that meaning nuances differ among non-human species, making a dichotomous approach to meaning questionable. Instead, we show that a multifaceted approach to meaning establishes how meaning appears within highly diverse examples of non-human communication, in ways consistent with the phenomenon's presence in human non-verbal communication and language(s). Therefore, without further recourse to ‘functional’ approaches that circumvent the critical question of whether any non-human meaning exists, we show that the concept of meaning is suitable for evolutionary biologists, behavioural ecologists, and others to study, to establish exactly which species exhibit meaning in their communication and in what ways.     

Explanations in search of observations

The paper explores how, in economics and biology, theoretical models are used as explanatory devices. It focuses on a modelling strategy by which, instead of starting with an unexplained regularity in the world, the modeller begins by creating a credible model world. The model world exhibits a regularity, induced by a mechanism in that world. The modeller concludes that there may be a part of the real world in which a similar regularity occurs and that, were that the case, the model would offer an explanation. Little concrete guidance is given about where such a regularity might be found. Three modelling exercises in evolutionary game theory—one from economics and two from biology—are used as case studies. Two of these (one from each discipline) exemplify ‘explanation in search of observation’. The third goes a step further, analysing a regularity in a model world and treating it as informative about the real world, but without saying anything about real phenomena. The paper argues that if the relation between the model and real worlds is understood in terms of similarity, and if modelling is understood as an ongoing discovery process rather than as the demonstration of empirical truths, there can be value in creating explanations before finding the regularities that are to be explained.     

The poetics of babytalk

Caretaker-infant attachment is a complex but well-recognized adaptation in humans. An early instance of (or precursor to) attachment behavior is the dyadic interaction between adults and infants of 6 to 24 weeks, commonly called "babytalk." Detailed analysis of 1 minute of spontaneous babytalk with an 8-week infant shows that the poetic texture of the mother’s speech—specifically its use of metrics, phonetics, and foregrounding—helps to shape and direct the baby’s attention, as it also coordinates the partners’ emotional communication. We hypothesize that the ability to respond to poetic features of language is present as early as the first few weeks of life and that this ability attunes cognitive and affective capacities in ways that provide a foundation for the skills at work in later aesthetic production and response. By linking developmental social processes with formal cognitive aspects of art, we challenge predominant views in evolutionary psychology that literary art is a superfluous byproduct of adaptive evolutionary mechanisms or primarily an ornament created by sexual selection. David S. Miall is Professor of English at the University of Alberta in Canada. He is the author of essays on British Romantic writers, empirical studies of readers’ responses to literature, hypertext and literary computing. Ellen Dissanayake is Visiting Scholar at the Walter Chapin Simpson Center for the Humanities, University of Washington. Her most recent book is Art and Intimacy: How the Arts Began (2000).     

Mathematical modeling of evolution. Solved and open problems

Evolution is a highly complex multilevel process and mathematical modeling of evolutionary phenomenon requires proper abstraction and radical reduction to essential features. Examples are natural selection, Mendel’s laws of inheritance, optimization by mutation and selection, and neutral evolution. An attempt is made to describe the roots of evolutionary theory in mathematical terms. Evolution can be studied in vitro outside cells with polynucleotide molecules. Replication and mutation are visualized as chemical reactions that can be resolved, analyzed, and modeled at the molecular level, and straightforward extension eventually results in a theory of evolution based upon biochemical kinetics. Error propagation in replication commonly results in an error threshold that provides an upper bound for mutation rates. Appearance and sharpness of the error threshold depend on the fitness landscape, being the distribution of fitness values in genotype or sequence space. In molecular terms, fitness landscapes are the results of two consecutive mappings from sequences into structures and from structures into the (nonnegative) real numbers. Some properties of genotype–phenotype maps are illustrated well by means of sequence–structure relations of RNA molecules. Neutrality in the sense that many RNA sequences form the same (coarse grained) structure is one of these properties, and characteristic for such mappings. Evolution cannot be fully understood without considering fluctuations—each mutant originates form a single copy, after all. The existence of neutral sets of genotypes called neutral networks, in particular, necessitates stochastic modeling, which is introduced here by simulation of molecular evolution in a kind of flowreactor.     

Synthetic constructs functionally mimicking ribonuclease A

The mechanism of hydrolysis of RNA substrates—diribonucleoside monophosphate CpA and decaribonucleotide UUCAUGUAAA—by chemical constructs functionally mimicking ribonuclease A was studied. It is shown that RNA cleavage by chemical RNases 2L2 and 2D3 proceeds similar to the RNase A-induced RNA hydrolysis through 2′,3′-cyclophosphate as an intermediate product. A comparison of hydrolyses of CpA in water and D₂O revealed an isotope effect (K _H/K _D=2.28), which implies acid-base catalysis at the limiting stage of the reaction. Two feasible mechanisms of RNA hydrolysis by chemical RNases (linear and adjacent) are discussed.     

Fertility assurance through extrapair fertilization,and male parental effort

Extrapair paternity (EPP) has been observed in many formally monogamous species. Male pursuit of extrapair fertilizations (EPF) is explained by the advantages of having offspring that receive essential paternal care from other males. Because females are capable of exercising a degree of control over the post-copulatory sperm competition, EPP’s persistence indicates that females benefit from EPF. Thus, EPP involves cooperation between mated females and extrapair males. On the other hand, mated males exhibit a spectrum of anti-cuckolding strategies. Hence, extrapair attributes of diverse species and populations reported in the literature are particular solutions of evolutionary games involving gender-specific cuckolding/anti-cuckolding strategies. Here we use game theoretical methods to study the effect of male paternal effort conserving strategies in situations where females seek EPF for reasons of genetic compatibility and/or in pursuit of genetic diversity for their offspring. Our results indicate that in these circumstances pursuit of EPF is the only evolutionary stable female strategy. Males, on the other hand, have two, mutually exclusive, evolutionary stable strategies: males that restrict parental care regardless of their mate’s fidelity, and males that never restrict parental care. That is, when females seek EPF for reasons of fertility assurance and/or genetic diversity, the conditional male strategy—therein the male’s parental efforts are based on his certainty of paternity—loses in competition with the unconditional strategies.     

Dialog with black box: using Information Theory to study animal language behaviour

In this review, three main experimental approaches for studying animal language behaviour are compared: (1) direct decoding of animals’ communication, (2) the use of intermediary languages to communicate with animals and (3) application of ideas and methods of the Information Theory for studying quantitative characteristics of animal communication. Each of the three methodological approaches has its specific power and specific limitations. Deciphering animals’ signals reveals a complex picture of natural communication in its evolutionary perspective but only fragmentary because of many methodological barriers, among which low repeatability of standard living situations seems to be a bottleneck. Language-training experiments are of great help for discovering potentials of animal language behaviour but leaves characteristics of their natural communications unclear. The use of the methods of Information Theory is based on measuring the time duration that animals spend on transmitting messages of definite information content and complexity. This approach, although does not reveal the nature of animals’ signals, provides a new dimension for studying important characteristics of natural communication systems, which have not been available before. First of all, this approach enables explorers of animals’ language behaviour to obtain knowledge just about the ability of subjects for transferring meaningful messages. Besides, the important properties of animal communication and intelligence can be evaluated such as the rate of information transmission, the complexity of transferred information and potential flexibility of communication systems.     

Click-based echolocation in bats: not so primitive after all

Echolocating bats of the genus Rousettus produce click sonar signals, using their tongue (lingual echolocation). These signals are often considered rudimentary and are believed to enable only crude performance. However, the main argument supporting this belief, namely the click’s reported long duration, was recently shown to be an artifact. In fact, the sonar clicks of Rousettus bats are extremely short, ~50–100 μs, similar to dolphin vocalizations. Here, we present a comparison between the sonar systems of the ‘model species’ of laryngeal echolocation, the big brown bat (Eptesicus fuscus), and that of lingual echolocation, the Egyptian fruit bat (Rousettus aegyptiacus). We show experimentally that in tasks, such as accurate landing or detection of medium-sized objects, click-based echolocation enables performance similar to laryngeal echolocators. Further, we describe a sophisticated behavioral strategy for biosonar beam steering in clicking bats. Finally, theoretical analyses of the signal design—focusing on their autocorrelations and wideband ambiguity functions—predict that in some aspects, such as target ranging and Doppler-tolerance, click-based echolocation might outperform laryngeal echolocation. Therefore, we suggest that click-based echolocation in bats should be regarded as a viable echolocation strategy, which is in fact similar to the biosonar used by most echolocating animals, including whales and dolphins.     

Three reciprocally monophyletic mtDNA lineages elucidate the taxonomic status of Grant’s gazelles

The intraspecific phylogeography of Grant’s gazelles Nanger granti was assessed with mitochondrial DNA control region sequences. Samples of 177 individuals from 17 Kenyan and Tanzanian populations were analysed. Three highly divergent, reciprocally monophyletic lineages were found, with among group net nucleotide distances of 8–12%. The three lineages—notata, granti and petersii—grouped populations according to their geographic origin, encompassing populations in the north, southwest, and east, respectively. The mtDNA lineages reflected distinct evolutionary trajectories, and the data are discussed in reference to the four currently recognised subspecies. We suggest Grant’s gazelles be raised to the superspecies Nanger (granti) comprising three taxonomic units corresponding to the three mtDNA lineages. There was no evidence of gene flow between the notata and granti lineages, despite their geographic proximity, suggesting reproductive isolation. These constitute evolutionary significant units within the adaptive evolutionary framework. Due to its restricted geographic distribution and genetic and morphological distinctiveness, we suggest the petersii lineage be raised to the species Nanger (granti) petersii within the Grant’s gazelles superspecies.     

Primate Communication: By Nature Honest, or by Experience Wise?

We review evolutionary views on honesty and deception and their application to studies of nonhuman primate communication. There is evidence that some primate signals are likely to be accurate on the basis of costliness. They appear most often in contexts that include overtly competitive interactions in which unrelated individuals have limited access to information about one another. However, both game theoretic models and most empirical work suggest that costly signals are not often likely to be the basis for honest communication in nonhuman primates. Inexpensive signaling can exist in contexts wherein communication occurs among related animals, something common among many nonhuman primate societies. Another condition in which inexpensive signaling is possible and that is also typical of nonhuman primates, is when sender and receiver both benefit from coordinated interactions. Additionally, when individuals interact repeatedly and can use past interactions to assess the honesty of signals and to modify future response to signals, low-cost signals can evolve. Nonhuman primates appear to deal with the problem of deception via skeptical responding, which can be largely accounted for by learning rules and the fact that they live in stable social groups and can recognize one another and recall past interactions.