From emotional signals to symbols.

The quest for the origins of language is a diverse enterprise, where research from a variety of disciplines brings area-specific ideas and area-specific terminology to bear. This variety often results in misunderstandings and misconceptions about communication in various species. In the present paper, we argue for focus on emotional systems as the primary motivators for social signals in animals in general. This focus can help resolve discrepancies of interpretation among different areas of inquiry and can illuminate distinctions among different social signals as well as their phylogenetic origins in animals and especially in humans. We advocate, following Jaak Panksepp, a view wherein the Seeking System, the endogenous tendency to search and explore, is the most fundamental emotional motivation. The Seeking System forms the basis for flexible, voluntary, and exploratory control of motor systems and makes much of learning possible. The relative lack of vocal learning and expression in nonhuman primates contrasted with extensive vocal learning and expression in humans began, we propose, with the evolution in ancient hominins of a necessary foundation for the many subsequent capabilities required for language. That foundation was, according to the reasoning, naturally selected in the form of neurological connections between the Seeking System and mechanisms of glottal/phonatory control. The new connections allowed ancient hominins to develop flexible, endogenous vocal fitness signals produced at very high rates and including large numbers of discrete syllables, recombinable to form syllable combinations with many prosodic variations. The increasing sociality of hominins supported evolution of massive expansion in the utilization of these flexible vocal forms to allow development of words and sentences.

Sex differences in infant vocalization and the origin of language.

Seeking to discern the earliest sex differences in language-related activities, our focus is vocal activity in the first two years of life, following up on recent research that unexpectedly showed boys produced significantly more speech-like vocalizations (protophones) than girls during the first year of life.We now bring a much larger body of data to bear on the comparison of early sex differences in vocalization, data based on automated analysis of all-day recordings of infants in their homes. The new evidence, like that of the prior study, also suggests boys produce more protophones than girls in the first year and offers additional basis for informed speculation about biological reasons for these differences. More broadly, the work offers a basis for informed speculations about foundations of language that we propose to have evolved in our distant hominin ancestors, foundations also required in early vocal development of modern human infants.

Perspectives on the origin of language: Infants vocalize most during independent vocal play but produce their most speech-like vocalizations during turn taking.

A growing body of research emphasizes both endogenous and social motivations in human vocal development. Our own efforts seek to establish an evolutionary and developmental perspective on the existence and usage of speech-like vocalizations ("protophones") in the first year of life. We evaluated the relative occurrence of protophones in 40 typically developing infants across the second-half year based on longitudinal all-day recordings. Infants showed strong endogenous motivation to vocalize, producing vastly more protophones during independent vocal exploration and play than during vocal turn taking. Both periods of vocal play and periods of turn-taking corresponded to elevated levels of the most advanced protophones (canonical babbling) relative to periods without vocal play or without turn-taking. Notably, periods of turn taking showed even more canonical babbling than periods of vocal play. We conclude that endogenous motivation drives infants' tendencies to explore and display a great number of speech-like vocalizations, but that social interaction drives the production of the most speech-like forms. The results inform our previously published proposal that the human infant has been naturally selected to explore protophone production and that the exploratory inclination in our hominin ancestors formed a foundation for language.

Protophones, the precursors to speech, dominate the human infant vocal landscape.

Human infant vocalization is viewed as a critical foundation for vocal learning and language. All apes share distress sounds (shrieks and cries) and laughter. Another vocal type, speech-like sounds, common in human infants, is rare but not absent in other apes. These three vocal types form a basis for especially informative cross-species comparisons. To make such comparisons possible we need empirical research documenting the frequency of occurrence of all three. The present work provides a comprehensive portrayal of these three vocal types in the human infant from longitudinal research in various circumstances of recording. Recently, the predominant vocalizations of the human infant have been shown to be speech-like sounds, or 'protophones', including both canonical and non-canonical babbling. The research shows that protophones outnumber cries by a factor of at least five based on data from random-sampling of all-day recordings across the first year. The present work expands on the prior reports, showing the protophones vastly outnumber both cry and laughter in both all-day and laboratory recordings in various circumstances. The data provide new evidence of the predominance of protophones in the infant vocal landscape and illuminate their role in human vocal learning and the origin of language. This article is part of the theme issue 'Vocal learning in animals and humans'.

Infant boys are more vocal than infant girls.

Although it is generally assumed females have a language advantage over males, Oller et al., studying all-day recordings of 100 infants, found that boys in the first year of life produced more speech-like vocalizations than girls and that the effect size was more than four times larger than the commonly reported female language advantage.

Functionally Flexible Signaling and the Origin of Language.

At the earliest break of ancient hominins from their primate relatives in vocal communication, we propose a selection pressure on vocal fitness signaling by hominin infants. Exploratory vocalizations, not tied to expression of distress or immediate need, could have helped persuade parents of the wellness and viability of the infants who produced them. We hypothesize that hominin parents invested more in infants who produced such signals of fitness plentifully, neglecting or abandoning them less often than infants who produced the sounds less frequently. Selection for such exploratory vocalization provided a critically important inclination and capability relevant to language, we reason, because the system that encouraged spontaneous vocalization also made vocalization functionally flexible to an extent that has not been observed in any other animal. Although this vocal flexibility did not by itself create language, it provided an essential foundation upon which language would evolve through a variety of additional steps. In evaluating this speculation, we consider presumable barriers to evolving language that are thought to be implications of Darwinian Theory. It has been claimed that communication always involves sender self-interest and that self-interest leads to deceit, which is countered through clever detection by receivers. The constant battle of senders and receivers has been thought to pose an insuperable challenge to honest communication, which has been viewed as a requirement of language. To make communication honest, it has been proposed that stable signaling requires costly handicaps for the sender, and since language cannot entail high cost, the reasoning has suggested an insurmountable obstacle to the evolution of language. We think this presumed honesty barrier is an illusion that can be revealed by recognition of the fact that language is not inherently honest and in light of the distinction between illocutionary force and semantics. Our paper also considers barriers to the evolution of language (not having to do with honesty) that we think may have actually played important roles in preventing species other than humans from evolving language.

Language Origins Viewed in Spontaneous and Interactive Vocal Rates of Human and Bonobo Infants.

From the first months of life, human infants produce "protophones," speech-like, non-cry sounds, presumed absent, or only minimally present in other apes. But there have been no direct quantitative comparisons to support this presumption. In addition, by 2 months, human infants show sustained face-to-face interaction using protophones, a pattern thought also absent or very limited in other apes, but again, without quantitative comparison. Such comparison should provide evidence relevant to determining foundations of language, since substantially flexible vocalization, the inclination to explore vocalization, and the ability to interact socially by means of vocalization are foundations for language. Here we quantitatively compare data on vocalization rates in three captive bonobo (Pan paniscus) mother-infant pairs with various sources of data from our laboratories on human infant vocalization. Both humans and bonobos produced distress sounds (cries/screams) and laughter. The bonobo infants also produced sounds that were neither screams nor laughs and that showed acoustic similarities to the human protophones. These protophone-like sounds confirm that bonobo infants share with humans the capacity to produce vocalizations that appear foundational for language. Still, there were dramatic differences between the species in both quantity and function of the protophone and protophone-like sounds. The bonobo protophone-like sounds were far less frequent than the human protophones, and the human protophones were far less likely to be interpreted as complaints and more likely as vocal play. Moreover, we found extensive vocal interaction between human infants and mothers, but no vocal interaction in the bonobo mother-infant pairs-while bonobo mothers were physically responsive to their infants, we observed no case of a bonobo mother vocalization directed to her infant. Our cross-species comparison focuses on low- and moderate-arousal circumstances because we reason the roots of language entail vocalization not triggered by excitement, for example, during fighting or intense play. Language appears to be founded in flexible vocalization, used to regulate comfortable social interaction, to share variable affective states at various levels of arousal, and to explore vocalization itself.

New Frontiers in Language Evolution and Development.

This article introduces the Special Issue and its focus on research in language evolution with emphasis on theory as well as computational and robotic modeling. A key theme is based on the growth of evolutionary developmental biology or evo-devo. The Special Issue consists of 13 articles organized in two sections: A) Theoretical foundations and B) Modeling and simulation studies. All the papers are interdisciplinary in nature, encompassing work in biological and linguistic foundations for the study of language evolution as well as a variety of computational and robotic modeling efforts shedding light on how language may be developed and may have evolved.

Developmental Plasticity and Language: A Comparative Perspective.

The growing field of evo-devo is increasingly demonstrating the complexity of steps involved in genetic, intracellular regulatory, and extracellular environmental control of the development of phenotypes. A key result of such work is an account for the remarkable plasticity of organismal form in many species based on relatively minor changes in regulation of highly conserved genes and genetic processes. Accounting for behavioral plasticity is of similar potential interest but has received far less attention. Of particular interest is plasticity in communication systems, where human language represents an ultimate target for research. The present paper considers plasticity of language capabilities in a comparative framework, focusing attention on examples of a remarkable fact: Whereas there exist design features of mature human language that have never been observed to occur in non-humans in the wild, many of these features can be developed to notable extents when non-humans are enculturated through human training (especially with intensive social interaction). These examples of enculturated developmental plasticity across extremely diverse taxa suggest, consistent with the evo-devo theme of highly conserved processes in evolution, that human language is founded in part on cognitive capabilities that are indeed ancient and that even modern humans show self-organized emergence of many language capabilities in the context of rich enculturation, built on the special social/ecological history of the hominin line. Human culture can thus be seen as a regulatory system encouraging language development in the context of a cognitive background with many highly conserved features.

Vocal Development as a Guide to Modeling the Evolution of Language.

Modeling of evolution and development of language has principally utilized mature units of spoken language, phonemes and words, as both targets and inputs. This approach cannot address the earliest phases of development because young infants are unable to produce such language features. We argue that units of early vocal development-protophones and their primitive illocutionary/perlocutionary forces-should be targeted in evolutionary modeling because they suggest likely units of hominin vocalization/communication shortly after the split from the chimpanzee/bonobo lineage, and because early development of spontaneous vocal capability is a logically necessary step toward vocal language, a root capability without which other crucial steps toward vocal language capability are impossible. Modeling of language evolution/development must account for dynamic change in early communicative units of form/function across time. We argue for interactive contributions of sender/infants and receiver/caregivers in a feedback loop involving both development and evolution and propose to begin computational modeling at the hominin break from the primate communicative background.

On Quantitative Comparative Research in Communication and Language Evolution.

Quantitative comparison of human language and natural animal communication requires improved conceptualizations. We argue that an infrastructural approach to development and evolution incorporating an extended interpretation of the distinctions among illocution, perlocution, and meaning (Austin 1962; Oller and Griebel 2008) can help place the issues relevant to quantitative comparison in perspective. The approach can illuminate the controversy revolving around the notion of functional referentiality as applied to alarm calls, for example in the vervet monkey. We argue that referentiality offers a poor point of quantitative comparison across language and animal communication in the wild. Evidence shows that even newborn human cry could be deemed to show functional referentiality according to the criteria typically invoked by advocates of referentiality in animal communication. Exploring the essence of the idea of illocution, we illustrate an important realm of commonality among animal communication systems and human language, a commonality that opens the door to more productive, quantifiable comparisons. Finally, we delineate two examples of infrastructural communicative capabilities that should be particularly amenable to direct quantitative comparison across humans and our closest relatives.

Vocabulary learning in a Yorkshire terrier: slow mapping of spoken words.

Rapid vocabulary learning in children has been attributed to "fast mapping", with new words often claimed to be learned through a single presentation. As reported in 2004 in Science a border collie (Rico) not only learned to identify more than 200 words, but fast mapped the new words, remembering meanings after just one presentation. Our research tests the fast mapping interpretation of the Science paper based on Rico's results, while extending the demonstration of large vocabulary recognition to a lap dog. We tested a Yorkshire terrier (Bailey) with the same procedures as Rico, illustrating that Bailey accurately retrieved randomly selected toys from a set of 117 on voice command of the owner. Second we tested her retrieval based on two additional voices, one male, one female, with different accents that had never been involved in her training, again showing she was capable of recognition by voice command. Third, we did both exclusion-based training of new items (toys she had never seen before with names she had never heard before) embedded in a set of known items, with subsequent retention tests designed as in the Rico experiment. After Bailey succeeded on exclusion and retention tests, a crucial evaluation of true mapping tested items previously successfully retrieved in exclusion and retention, but now pitted against each other in a two-choice task. Bailey failed on the true mapping task repeatedly, illustrating that the claim of fast mapping in Rico had not been proven, because no true mapping task had ever been conducted with him. It appears that the task called retention in the Rico study only demonstrated success in retrieval by a process of extended exclusion.

Does Octopus vulgaris have preferred arms?

Previous behavioral studies in Octopus vulgaris revealed lateralization of eye use. In this study, the authors expanded the scope to investigate arm preferences. The octopus's generalist hunting lifestyle and the structure of their arms suggest that these animals have no need to designate specific arms for specific tasks. However, octopuses also show behaviors, like exploration, in which only single or small groups of arms are involved. Here the authors show that octopuses had a strong preference for anterior arm use to reach for and explore objects, which points toward a task division between anterior and posterior arms. Four out of 8 subjects also showed a lateral bias. In addition, octopuses had a preference for a specific arm to reach into a T maze to retrieve a food reward. These findings give evidence for limb-specialization in an animal whose 8 arms were believed to be equipotential.

Octopus arm choice is strongly influenced by eye use.

This study aims to investigate the octopus' eye and arm coordination and raises the question if visual guidance determines choice of arm use. Octopuses possess eight seemingly identical arms but have recently been reported to show a preference as to which arm they use to initiate contact with objects. These animals also exhibit lateralized eye use, therefore, a connection between eye and arm preference seems possible. Seven Octopus vulgaris were observed during approach, contact initiation and exploration of plastic objects that were positioned on three different levels in the water column. The subjects most commonly used an arm to initiate contact with an object that was in a direct line between the eye used to look at the object, and the object itself. This indicates that choice of arm use is spatially rather opportunistic when depending on visual guidance. Additionally, first contact with an object was usually established by the central third of the arm and in arm contact sequences neighboring arms were the most likely to follow an arm already touching the object. Although results point towards strong eye/arm coordination, we did not find lateralized behavior in this experiment. Results are discussed from a neuro-anatomical, behavioral and ecological perspective.

Brightness discrimination in the dog.

Almost nothing is known about brightness discrimination in animals and how this ability relates to their lifestyles. As arrhythmic visual generalists, three dogs, a German shepherd and two Belgian shepherds, were tested on their ability to discriminate brightness using a series of 30 shades of grey varying from white to black. The dogs were trained to discriminate between different shades of grey in a simultaneous two-choice situation. Weber's law can be correlated to their ability to discriminate brightness differences with a calculated Weber fraction of 0.22 for the German shepherd and 0.27 for the Belgian shepherds. Thus brightness discrimination in dogs is about 2 times worse than in humans, a diurnal species.