A chimpanzee recognizes varied acoustical versions of sine-wave and noise-vocoded speech.

Previous research demonstrated that a language-trained chimpanzee recognized familiar English words in sine-wave and noise-vocoded forms (Heimbauer et al. Curr Biol 21:1210-1214, 2011). However, those results did not provide information regarding processing strategies of the specific acoustic cues to which the chimpanzee may have attended. The current experiments tested this chimpanzee and adult humans using sine-wave and noise-vocoded speech manipulated using specific sine-waves and a different number of noise bands, respectively. Similar to humans tested with the same stimuli, the chimpanzee was more successful identifying sine-wave speech when both SW1 and SW2 were present - the components that are modeled on formants F1 and F2 in the natural speech signal. Results with noise-vocoded speech revealed that the chimpanzee and humans performed best with stimuli that included four or five noise bands, as compared to those with three and two. Overall, amplitude and frequency modulation over time were important for identification of sine-wave and noise-vocoded speech, with further evidence that a nonhuman primate is capable of using top-down processes for speech perception when the signal is altered and incomplete.

Visual artificial grammar learning by rhesus macaques (Macaca mulatta): exploring the role of grammar complexity and sequence length.

Humans and nonhuman primates can learn about the organization of stimuli in the environment using implicit sequential pattern learning capabilities. However, most previous artificial grammar learning studies with nonhuman primates have involved relatively simple grammars and short input sequences. The goal in the current experiments was to assess the learning capabilities of monkeys on an artificial grammar-learning task that was more complex than most others previously used with nonhumans. Three experiments were conducted using a joystick-based, symmetrical-response serial reaction time task in which two monkeys were exposed to grammar-generated sequences at sequence lengths of four in Experiment 1, six in Experiment 2, and eight in Experiment 3. Over time, the monkeys came to respond faster to the sequences generated from the artificial grammar compared to random versions. In a subsequent generalization phase, subjects generalized their knowledge to novel sequences, responding significantly faster to novel instances of sequences produced using the familiar grammar compared to those constructed using an unfamiliar grammar. These results reveal that rhesus monkeys can learn and generalize the statistical structure inherent in an artificial grammar that is as complex as some used with humans, for sequences up to eight items long. These findings are discussed in relation to whether or not rhesus macaques and other primate species possess implicit sequence learning abilities that are similar to those that humans draw upon to learn natural language grammar.

Sexual selection on male vocal fundamental frequency in humans and other anthropoids.

In many primates, including humans, the vocalizations of males and females differ dramatically, with male vocalizations and vocal anatomy often seeming to exaggerate apparent body size. These traits may be favoured by sexual selection because low-frequency male vocalizations intimidate rivals and/or attract females, but this hypothesis has not been systematically tested across primates, nor is it clear why competitors and potential mates should attend to vocalization frequencies. Here we show across anthropoids that sexual dimorphism in fundamental frequency (F0) increased during evolutionary transitions towards polygyny, and decreased during transitions towards monogamy. Surprisingly, humans exhibit greater F0 sexual dimorphism than any other ape. We also show that low-F0 vocalizations predict perceptions of men's dominance and attractiveness, and predict hormone profiles (low cortisol and high testosterone) related to immune function. These results suggest that low male F0 signals condition to competitors and mates, and evolved in male anthropoids in response to the intensity of mating competition.

Silky sifaka (Propithecus candidus) "zzuss" vocalizations: sexual dimorphism, individuality, and function in the alarm call of a monomorphic lemur.

Vocalizations of Madagascar's lemurs have generally been less investigated than those of other primate groups, with virtually no information available about calling in the silky sifaka (Propithecus candidus), a large rainforest species. Current work examined the "zzuss" vocalization, one of the most common and loudest sounds produced by this monomorphic species, and included 160 calls from nine adults (five males, four females) in three groups. Analyses focused on overall acoustic features, individual and sex differences, call usage, and likely function. Acoustically, the calls included separable turbulent noise and tonal components, with the later often marked by frequency jumps and dramatic frequency modulation. Male and female zzuss calls differed most in F0- and amplitude-related features, characteristics that are relatively unconstrained by overall body size. All measures differed among individual callers, with F0-related variables again playing the largest role. Based on usage, these calls most likely function both as generalized alarm and group-coordination signals. The sounds were thus of interest in several regards, including showing sexual differentiation in the absence of other dimorphisms, exhibiting primarily F0-based differentiation in both sex- and individual-based comparisons, and combining apparent alarm and coordination functions across a variety of contexts.

Capuchin monkeys (Cebus apella) use positive, but not negative, auditory cues to infer food location.

Nonhuman primates appear to capitalize more effectively on visual cues than corresponding auditory versions. For example, studies of inferential reasoning have shown that monkeys and apes readily respond to seeing that food is present ("positive" cuing) or absent ("negative" cuing). Performance is markedly less effective with auditory cues, with many subjects failing to use this input. Extending recent work, we tested eight captive tufted capuchins (Cebus apella) in locating food using positive and negative cues in visual and auditory domains. The monkeys chose between two opaque cups to receive food contained in one of them. Cup contents were either shown or shaken, providing location cues from both cups, positive cues only from the baited cup, or negative cues from the empty cup. As in previous work, subjects readily used both positive and negative visual cues to secure reward. However, auditory outcomes were both similar to and different from those of earlier studies. Specifically, all subjects came to exploit positive auditory cues, but none responded to negative versions. The animals were also clearly different in visual versus auditory performance. Results indicate that a significant proportion of capuchins may be able to use positive auditory cues, with experience and learning likely playing a critical role. These findings raise the possibility that experience may be significant in visually based performance in this task as well, and highlight that coming to grips with evident differences between visual versus auditory processing may be important for understanding primate cognition more generally.

A Serial Reaction Time (SRT) task with symmetrical joystick responding for nonhuman primates.

The serial reaction time (SRT) task is a simple procedure in which participants produce differentiated responses to each of a series of stimuli presented at varying locations. Learning about stimulus order is revealed through decreased latencies for structured versus randomized sequences. Although widely used with humans and well suited to nonhumans, this paradigm is little used in comparative research. In the present article, we describe an SRT procedure that uses colored circles as stimuli, a circular layout of locations, and symmetrical joystick deflections as responses. In two experiments, we showed that four rhesus macaques (Macaca mulatta) learned to track sequences up to eight items long, with three animals showing faster responding to repeating sequences than to randomized versions. After extended training, these participants also showed evidence of faster responding at all positions within repeating sequences. This method minimizes response effort, equates effort and travel distance across stimulus locations, and is applicable to any joystick-capable species.

A chimpanzee recognizes synthetic speech with significantly reduced acoustic cues to phonetic content.

A long-standing debate concerns whether humans are specialized for speech perception, which some researchers argue is demonstrated by the ability to understand synthetic speech with significantly reduced acoustic cues to phonetic content. We tested a chimpanzee (Pan troglodytes) that recognizes 128 spoken words, asking whether she could understand such speech. Three experiments presented 48 individual words, with the animal selecting a corresponding visuographic symbol from among four alternatives. Experiment 1 tested spectrally reduced, noise-vocoded (NV) synthesis, originally developed to simulate input received by human cochlear-implant users. Experiment 2 tested "impossibly unspeechlike" sine-wave (SW) synthesis, which reduces speech to just three moving tones. Although receiving only intermittent and noncontingent reward, the chimpanzee performed well above chance level, including when hearing synthetic versions for the first time. Recognition of SW words was least accurate but improved in experiment 3 when natural words in the same session were rewarded. The chimpanzee was more accurate with NV than SW versions, as were 32 human participants hearing these items. The chimpanzee's ability to spontaneously recognize acoustically reduced synthetic words suggests that experience rather than specialization is critical for speech-perception capabilities that some have suggested are uniquely human.

Two organizing principles of vocal production: Implications for nonhuman and human primates.

Vocal communication in nonhuman primates receives considerable research attention, with many investigators arguing for similarities between this calling and speech in humans. Data from development and neural organization show a central role of affect in monkey and ape sounds, however, suggesting that their calls are homologous to spontaneous human emotional vocalizations while having little relation to spoken language. Based on this evidence, we propose two principles that can be useful in evaluating the many and disparate empirical findings that bear on the nature of vocal production in nonhuman and human primates. One principle distinguishes production-first from reception-first vocal development, referring to the markedly different role of auditory-motor experience in each case. The second highlights a phenomenon dubbed dual neural pathways, specifically that when a species with an existing vocal system evolves a new functionally distinct vocalization capability, it occurs through emergence of a second parallel neural pathway rather than through expansion of the extant circuitry. With these principles as a backdrop, we review evidence of acoustic modification of calling associated with background noise, conditioning effects, audience composition, and vocal convergence and divergence in nonhuman primates. Although each kind of evidence has been interpreted to show flexible cognitively mediated control over vocal production, we suggest that most are more consistent with affectively grounded mechanisms. The lone exception is production of simple, novel sounds in great apes, which is argued to reveal at least some degree of volitional vocal control. If also present in early hominins, the cortically based circuitry surmised to be associated with these rudimentary capabilities likely also provided the substrate for later emergence of the neural pathway allowing volitional production in modern humans.

The evolution of laughter in great apes and humans.

It has long been claimed that human emotional expressions, such as laughter, have evolved from nonhuman displays. The aim of the current study was to test this prediction by conducting acoustic and phylogenetic analyses based on the acoustics of tickle-induced vocalizations of orangutans, gorillas, chimpanzees, bonobos and humans. Results revealed both important similarities and differences among the various species' vocalizations, with the phylogenetic tree reconstructed based on these acoustic data matching the well-established genetic relationships of great apes and humans. These outcomes provide evidence of a common phylogenetic origin of tickle-induced vocalizations in these taxa, which can therefore be termed "laughter" across all five species. Results are consistent with the claims of phylogenetic continuity of emotional expressions. Together with observations made on the use of laughter in great apes and humans, findings of this study further indicate that there were two main periods of selection-driven evolutionary change in laughter within the Hominidae, to a smaller degree, among the great apes and, most distinctively, after the separation of hominins from the last common ancestor with chimpanzees and bonobos.

I Like the Sound of Your Voice: Affective Learning about Vocal Signals.

This paper provides the first demonstration that the content of what a talker says is sufficient to imbue the acoustics of his voice with affective meaning. In two studies, participants listened to male talkers utter positive, negative, or neutral words. Next, participants completed a sequential evaluative priming task where a neutral word spoken by one of the same talkers was presented before each target word to be evaluated. We predicted, and found, that voices served as evaluative primes that influenced the speed with which participants evaluated the target words. These two experiments demonstrate that the human voice can take on affective meaning merely based on the positive or negative value of the words uttered by that voice. Implications for affective processing, the pragmatics of communication, and person-perception are discussed.

Reconstructing the evolution of laughter in great apes and humans.

Human emotional expressions, such as laughter, are argued to have their origins in ancestral nonhuman primate displays. To test this hypothesis, the current work examined the acoustics of tickle-induced vocalizations from infant and juvenile orangutans, gorillas, chimpanzees, and bonobos, as well as tickle-induced laughter produced by human infants. Resulting acoustic data were then coded as character states and submitted to quantitative phylogenetic analysis. Acoustic outcomes revealed both important similarities and differences among the five species. Furthermore, phylogenetic trees reconstructed from the acoustic data matched the well-established trees based on comparative genetics. Taken together, the results provide strong evidence that tickling-induced laughter is homologous in great apes and humans and support the more general postulation of phylogenetic continuity from nonhuman displays to human emotional expressions. Findings also show that distinctively human laughter characteristics such as predominantly regular, stable voicing and consistently egressive airflow are nonetheless traceable to characteristics of shared ancestors with great apes.

Asymmetries in the individual distinctiveness and maternal recognition of infant contact calls and distress screams in baboons.

A key component of nonhuman primate vocal communication is the production and recognition of clear cues to social identity that function in the management of these species' individualistic social relationships. However, it remains unclear how ubiquitous such identity cues are across call types and age-sex classes and what the underlying vocal production mechanisms responsible might be. This study focused on two structurally distinct call types produced by infant baboons in contexts that place a similar functional premium on communicating clear cues to caller identity: (1) contact calls produced when physically separated from, and attempting to relocate, mothers and (2) distress screams produced when aggressively attacked by other group members. Acoustic analyses and field experiments were conducted to examine individual differentiation in single vocalizations of each type and to test mothers' ability to recognize infant calls. Both call types showed statistically significant individual differentiation, but the magnitude of the differentiation was substantially higher in contact calls. Mothers readily discriminated own-offspring contact calls from those of familiar but unrelated infants, but did not do so when it came to distress screams. Several possible explanations for these asymmetries in call differentiation and recognition are considered.

GSU Praat Tools: scripts for modifying and analyzing sounds using Praat acoustics software.

The Praat acoustics program (Boersma, 2001) is powerful freeware that is widely used bybehavioral scientists working with digital sound. This article describes GSU Praat Tools, a script package that helps simplify and automate such work. The routines use Praat's scripting language to create new menus and commands within the existing interface, and can operate either on individual files or in batch mode. The new functions facilitate selecting, displaying, editing, filtering, and otherwise modifying sounds, quantifying acoustic features, and saving results in text-based data files. The package includes an installation script and user's manual, and is available free from psyvoso.googlepages.com/softwaredownload.

An acoustic analysis of laughter produced by congenitally deaf and normally hearing college students.

Relatively few empirical data are available concerning the role of auditory experience in nonverbal human vocal behavior, such as laughter production. This study compared the acoustic properties of laughter in 19 congenitally, bilaterally, and profoundly deaf college students and in 23 normally hearing control participants. Analyses focused on degree of voicing, mouth position, air-flow direction, temporal features, relative amplitude, fundamental frequency, and formant frequencies. Results showed that laughter produced by the deaf participants was fundamentally similar to that produced by the normally hearing individuals, which in turn was consistent with previously reported findings. Finding comparable acoustic properties in the sounds produced by deaf and hearing vocalizers confirms the presumption that laughter is importantly grounded in human biology, and that auditory experience with this vocalization is not necessary for it to emerge in species-typical form. Some differences were found between the laughter of deaf and hearing groups; the most important being that the deaf participants produced lower-amplitude and longer-duration laughs. These discrepancies are likely due to a combination of the physiological and social factors that routinely affect profoundly deaf individuals, including low overall rates of vocal fold use and pressure from the hearing world to suppress spontaneous vocalizations.

Listeners judge talker sex more efficiently from male than from female vowels.

Speech routinely provides cues as to the sex of the talker, in voiced sounds, these cues mainly reflect dimorphism in vocal anatomy. This dimorphism is not symmetrical, however, since during adolescent development, males specifically diverge from a previously shared trajectory with females. We therefore predicted that listeners would show a corresponding perceptual advantage for male sounds in talker-sex discrimination, a hypothesis tested using very brief, one- to eight-cycle vowel segments. The expected performance asymmetry was observed in threshold-like tests of multiple different vowels in Experiments 1-3, and a signal detection design in Experiment 4 helped rule out possible response bias effects. In confirming our counterintuitive prediction, the present study illustrates that a biological and evolutionary perspective can be helpful in understanding indexical cuing in speech.

Nonlinear acoustics in the pant hoots of common chimpanzees (Pan troglodytes): vocalizing at the edge.

Common chimpanzee (Pan troglodytes) "pant hoots" are multi-call events that build from quiet, consistently harmonic introductory sounds to loud, screamlike "climax" calls with acoustic irregularities known as "nonlinear phenomena" (NLP). Two possible functions of NLP in climax calls are to increase direct auditory impact on listeners and to signal physical condition. These possibilities were addressed by comparing climax calls from 12 wild chimpanzee males with "screams" and pant hoot "introduction" calls from the same individuals. Climax calls that included NLP were found to have higher fundamental frequencies (F0s) than introduction or climax calls that were purely harmonic. NLP onsets within climax calls were also specifically associated with local F0 maxima, suggesting vocalizers are vibrating their vocal folds at the upper limits of stability. Furthermore, climax calls showed far fewer NLP than did screams recorded from the same individuals, while showing equivalent or higher F0 values. Overall, the results are consistent with the hypothesis that the relative prevalence of NLP is a signal of physical condition, with callers "vocalizing at the edge" of regular, stable production while producing few NLP. The results are discussed in light of the initial hypotheses.

Acoustics and behavioral contexts of "gecker" vocalizations in young rhesus macaques (Macaca mulatta).

Loud, pulsed "gecker" vocalizations are commonly produced by young rhesus macaques in distressful circumstances. The acoustics, usage, and responses associated with these calls were examined using audio recordings and observational data from captive, socially living rhesus up to 24 months old. One-hundred-eleven gecker bouts were recorded from ten individuals (six males, four females), with most geckers produced during the first 6 months of age. A gecker call consisted of a bout of up to 28 pulses of spectrally structured noise with a single prominent frequency peak. Nine contexts of calling were identified, but little evidence of context-specific acoustic variation was found. While geckering often triggered responses by the vocalizer's mother, the most common outcome was the absence of any reaction. Females geckered longer and at higher rates than did males, while also showing acoustic evidence of greater vocal effort. Mothers nonetheless responded more often and more positively to males. Overall, results show that gecker acoustics vary somewhat with vocalizer sex, age, and likely arousal level, but do not reflect detailed aspects of behavioral context. Circumstances of production suggest that geckers function primarily to draw the attention of mothers, who in turn are selective in responding.

The relative roles of vowels and consonants in discriminating talker identity versus word meaning.

Three experiments tested the hypothesis that vowels play a disproportionate role in hearing talker identity, while consonants are more important in perceiving word meaning. In each study, listeners heard 128 stimuli consisting of two different words. Stimuli were balanced for same/different meaning, same/different talker, and male/female talker. The first word in each was intact, while the second was either intact (Experiment 1), or had vowels ("Consonants-Only") or consonants wels-Only") replaced by silence (Experiments 2, 3). Different listeners performed a same/ different judgment of either talker identity (Talker) or word meaning (Meaning). Baseline testing in Experiment 1 showed above-chance performance in both, with greater accuracy for Meaning. In Experiment 2, Talker identity was more accurately judged from Vowels-Only stimuli, with modestly better overall Meaning performance with Consonants-Only stimuli. However, performance with vowel-initial Vowels-Only stimuli in particular was most accurate of all. Editing Vowels-Only stimuli further in Experiment 3 had no effect on Talker discrimination, while dramatically reducing accuracy in the Meaning condition, including both vowel-initial and consonant-initial Vowels-Only stimuli. Overall, results confirmed a priori predictions, but are largely inconsistent with recent tests of vowels and consonants in sentence comprehension. These discrepancies and possible implications for the evolutionary origins of speech are discussed.

Characterizing noise in nonhuman vocalizations: Acoustic analysis and human perception of barks by coyotes and dogs.

Measuring noise as a component of mammalian vocalizations is of interest because of its potential relevance to the communicative function. However, methods for characterizing and quantifying noise are less well established than methods applicable to harmonically structured aspects of signals. Using barks of coyotes and domestic dogs, we compared six acoustic measures and studied how they are related to human perception of noisiness. Measures of harmonic-to-noise-ratio (HNR), percent voicing, and shimmer were found to be the best predictors of perceptual rating by human listeners. Both acoustics and perception indicated that noisiness was similar across coyote and dog barks, but within each species there was significant variation among the individual vocalizers. The advantages and disadvantages of the various measures are discussed.

Nonlinear acoustics in pant hoots of common chimpanzees (Pan troglodytes): frequency jumps, subharmonics, biphonation, and deterministic chaos.

The pant hoot calls produced by common chimpanzees (Pan troglodytes) are multi-call vocalizations that have figured prominently in investigations of acoustic communication in this species. Although pant hoots are predominantly harmonically structured, they can exhibit an acoustic complexity that has recently been linked to nonlinearity in the vocal-fold dynamics underlying typical mammalian sound production. We examined the occurrence of these sorts of nonlinear phenomena in pant hoot vocalizations, contrasting quieter and lower-pitched "introduction" components with loud and high-pitched "climax" calls in the same bouts. Spectrographic evidence revealed four kinds of nonlinear phenomena, including discrete frequency jumps, subharmonics, biphonation, and deterministic chaos. While these events were virtually never observed during the introduction, they occurred in more than half of the climax calls. Biphonation was by far the most common phenomenon, followed by subharmonics, chaos, and frequency jumps. Individual callers varied in the degree to which their climax calls exhibited nonlinear phenomena, but were consistent in showing more biphonation than other forms. These outcomes show that nonlinear phenomena are routinely present in chimpanzee pant hoots, and help lay the foundation for investigating the function of such events.