Of mice and men, nature and nurture, and a few red herrings.

Burkart et al.'s proposal is based on three false premises: (1) theories of the mind are either domain-specific/modular (DSM) or domain-general (DG); (2) DSM systems are considered inflexible, built by nature; and (3) animal minds are deemed as purely DSM. Clearing up these conceptual confusions is a necessary first step in understanding how general intelligence evolved.

Serotonin selectively influences moral judgment and behavior through effects on harm aversion.

Aversive emotional reactions to real or imagined social harms infuse moral judgment and motivate prosocial behavior. Here, we show that the neurotransmitter serotonin directly alters both moral judgment and behavior through increasing subjects' aversion to personally harming others. We enhanced serotonin in healthy volunteers with citalopram (a selective serotonin reuptake inhibitor) and contrasted its effects with both a pharmacological control treatment and a placebo on tests of moral judgment and behavior. We measured the drugs' effects on moral judgment in a set of moral 'dilemmas' pitting utilitarian outcomes (e.g., saving five lives) against highly aversive harmful actions (e.g., killing an innocent person). Enhancing serotonin made subjects more likely to judge harmful actions as forbidden, but only in cases where harms were emotionally salient. This harm-avoidant bias after citalopram was also evident in behavior during the ultimatum game, in which subjects decide to accept or reject fair or unfair monetary offers from another player. Rejecting unfair offers enforces a fairness norm but also harms the other player financially. Enhancing serotonin made subjects less likely to reject unfair offers. Furthermore, the prosocial effects of citalopram varied as a function of trait empathy. Individuals high in trait empathy showed stronger effects of citalopram on moral judgment and behavior than individuals low in trait empathy. Together, these findings provide unique evidence that serotonin could promote prosocial behavior by enhancing harm aversion, a prosocial sentiment that directly affects both moral judgment and moral behavior.

Syntax-induced pattern deafness.

Perceptual systems often force systematically biased interpretations upon sensory input. These interpretations are obligatory, inaccessible to conscious control, and prevent observers from perceiving alternative percepts. Here we report a similarly impenetrable phenomenon in the domain of language, where the syntactic system prevents listeners from detecting a simple perceptual pattern. Healthy human adults listened to three-word sequences conforming to patterns readily learned even by honeybees, rats, and sleeping human neonates. Specifically, sequences either started or ended with two words from the same syntactic category (e.g., noun-noun-verb or verb-verb-noun). Although participants readily processed the categories and learned repetition patterns over nonsyntactic categories (e.g., animal-animal-clothes), they failed to learn the repetition pattern over syntactic categories, even when explicitly instructed to look for it. Further experiments revealed that participants successfully learned the repetition patterns only when they were consistent with syntactically possible structures, irrespective of whether these structures were attested in English or in other languages unknown to the participants. When the repetition patterns did not match such syntactically possible structures, participants failed to learn them. Our results suggest that when human adults hear a string of nouns and verbs, their syntactic system obligatorily attempts an interpretation (e.g., in terms of subjects, objects, and predicates). As a result, subjects fail to perceive the simpler pattern of repetitions--a form of syntax-induced pattern deafness that is reminiscent of how other perceptual systems force specific interpretations upon sensory input.

Five-month-old infants' identification of the sources of vocalizations.

Humans speak, monkeys grunt, and ducks quack. How do we come to know which vocalizations animals produce? Here we explore this question by asking whether young infants expect humans, but not other animals, to produce speech, and further, whether infants have similarly restricted expectations about the sources of vocalizations produced by other species. Five-month-old infants matched speech, but not human nonspeech vocalizations, specifically to humans, looking longer at static human faces when human speech was played than when either rhesus monkey or duck calls were played. They also matched monkey calls to monkey faces, looking longer at static rhesus monkey faces when rhesus monkey calls were played than when either human speech or duck calls were played. However, infants failed to match duck vocalizations to duck faces, even though infants likely have more experience with ducks than monkeys. Results show that by 5 months of age, human infants generate expectations about the sources of some vocalizations, mapping human faces to speech and rhesus faces to rhesus calls. Infants' matching capacity does not appear to be based on a simple associative mechanism or restricted to their specific experiences. We discuss these findings in terms of how infants may achieve such competence, as well as its specificity and relevance to acquiring language.

The evolutionary origins of human patience: temporal preferences in chimpanzees, bonobos, and human adults.

To make adaptive choices, individuals must sometimes exhibit patience, forgoing immediate benefits to acquire more valuable future rewards [1-3]. Although humans account for future consequences when making temporal decisions [4], many animal species wait only a few seconds for delayed benefits [5-10]. Current research thus suggests a phylogenetic gap between patient humans and impulsive, present-oriented animals [9, 11], a distinction with implications for our understanding of economic decision making [12] and the origins of human cooperation [13]. On the basis of a series of experimental results, we reject this conclusion. First, bonobos (Pan paniscus) and chimpanzees (Pan troglodytes) exhibit a degree of patience not seen in other animals tested thus far. Second, humans are less willing to wait for food rewards than are chimpanzees. Third, humans are more willing to wait for monetary rewards than for food, and show the highest degree of patience only in response to decisions about money involving low opportunity costs. These findings suggest that core components of the capacity for future-oriented decisions evolved before the human lineage diverged from apes. Moreover, the different levels of patience that humans exhibit might be driven by fundamental differences in the mechanisms representing biological versus abstract rewards.

The apes' edge: positional learning in chimpanzees and humans.

A wide variety of organisms produce actions and signals in particular temporal sequences, including the motor actions recruited during tool-mediated foraging, the arrangement of notes in the songs of birds, whales and gibbons, and the patterning of words in human speech. To accurately reproduce such events, the elements that comprise such sequences must be memorized. Both memory and artificial language learning studies have revealed at least two mechanisms for memorizing sequences, one tracking co-occurrence statistics among items in sequences (i.e., transitional probabilities) and the other one tracking the positions of items in sequences, in particular those of items in sequence-edges. The latter mechanism seems to dominate the encoding of sequences after limited exposure, and to be recruited by a wide array of grammatical phenomena. To assess whether humans differ from other species in their reliance on one mechanism over the other after limited exposure, we presented chimpanzees (Pan troglodytes) and human adults with brief exposure to six items, auditory sequences. Each sequence consisted of three distinct sound types (X, A, B), arranged according to two simple temporal rules: the A item always preceded the B item, and the sequence-edges were always occupied by the X item. In line with previous results with human adults, both species primarily encoded positional information from the sequences; that is, they kept track of the items that occurred in the sequence-edges. In contrast, the sensitivity to co-occurrence statistics was much weaker. Our results suggest that a mechanism to spontaneously encode positional information from sequences is present in both chimpanzees and humans and may represent the default in the absence of training and with brief exposure. As many grammatical regularities exhibit properties of this mechanism, it may be recruited by language and constrain the form that certain grammatical regularities take.

Word segmentation with universal prosodic cues.

When listening to speech from one's native language, words seem to be well separated from one another, like beads on a string. When listening to a foreign language, in contrast, words seem almost impossible to extract, as if there was only one bead on the same string. This contrast reveals that there are language-specific cues to segmentation. The puzzle, however, is that infants must be endowed with a language-independent mechanism for segmentation, as they ultimately solve the segmentation problem for any native language. Here, we approach the acquisition problem by asking whether there are language-independent cues to segmentation that might be available to even adult learners who have already acquired a native language. We show that adult learners recognize words in connected speech when only prosodic cues to word-boundaries are given from languages unfamiliar to the participants. In both artificial and natural speech, adult English speakers, with no prior exposure to the test languages, readily recognized words in natural languages with critically different prosodic patterns, including French, Turkish and Hungarian. We suggest that, even though languages differ in their sound structures, they carry universal prosodic characteristics. Further, these language-invariant prosodic cues provide a universally accessible mechanism for finding words in connected speech. These cues may enable infants to start acquiring words in any language even before they are fine-tuned to the sound structure of their native language.

The tuning of human neonates' preference for speech.

Human neonates prefer listening to speech compared to many nonspeech sounds, suggesting that humans are born with a bias for speech. However, neonates' preference may derive from properties of speech that are not unique but instead are shared with the vocalizations of other species. To test this, thirty neonates and sixteen 3-month-olds were presented with nonsense speech and rhesus monkey vocalizations. Neonates showed no preference for speech over rhesus vocalizations but showed a preference for both these sounds over synthetic sounds. In contrast, 3-month-olds preferred speech to rhesus vocalizations. Neonates' initial biases minimally include speech and monkey vocalizations. These listening preferences are sharpened over 3 months, yielding a species-specific preference for speech, paralleling findings on infant face perception.

Action comprehension in non-human primates: motor simulation or inferential reasoning?

Some argue that action comprehension is intimately connected with the observer's own motor capacities, whereas others argue that action comprehension depends on non-motor inferential mechanisms. We address this debate by reviewing comparative studies that license four conclusions: monkeys and apes extract the meaning of an action (i) by going beyond the surface properties of actions, attributing goals and intentions to the agent; (ii) by using environmental information to infer when actions are rational; (iii) by making predictions about an agent's goal, and the most probable action to obtain the goal given environmental constraints; (iv) in situations in which they are physiologically incapable of producing the actions. Motor theories are, thus, insufficient to account for primate action comprehension in the absence of inferential mechanisms.

Evidence of an evolutionary precursor to human language affixation in a non-human primate.

Human language, and grammatical competence in particular, relies on a set of computational operations that, in its entirety, is not observed in other animals. Such uniqueness leaves open the possibility that components of our linguistic competence are shared with other animals, having evolved for non-linguistic functions. Here, we explore this problem from a comparative perspective, asking whether cotton-top tamarin monkeys (Saguinus oedipus) can spontaneously (no training) acquire an affixation rule that shares important properties with our inflectional morphology (e.g. the rule that adds -ed to create the past tense, as in the transformation of walk into walk-ed). Using playback experiments, we show that tamarins discriminate between bisyllabic items that start with a specific 'prefix' syllable and those that end with the same syllable as a 'suffix'. These results suggest that some of the computational mechanisms subserving affixation in a diversity of languages are shared with other animals, relying on basic perceptual or memory primitives that evolved for non-linguistic functions.

Will travel for food: spatial discounting in two new world monkeys.

Nonhuman animals steeply discount the future, showing a preference for small, immediate over large, delayed rewards. Currently unclear is whether discounting functions depend on context. Here, we examine the effects of spatial context on discounting in cotton-top tamarins (Saguinus oedipus) and common marmosets (Callithrix jacchus), species known to differ in temporal discounting. We presented subjects with a choice between small, nearby rewards and large, distant rewards. Tamarins traveled farther for the large reward than marmosets, attending to the ratio of reward differences rather than their absolute values. This species difference contrasts with performance on a temporal task in which marmosets waited longer than tamarins for the large reward. These comparative data indicate that context influences choice behavior, with the strongest effect seen in marmosets who discounted more steeply over space than over time. These findings parallel details of each species' feeding ecology. Tamarins range over large distances and feed primarily on insects, which requires using quick, impulsive action. Marmosets range over shorter distances than tamarins and feed primarily on tree exudates, a clumped resource that requires patience to wait for sap to exude. These results show that discounting functions are context specific, shaped by a history of ecological pressures.

Human cerebral response to animal affective vocalizations.

It is presently unknown whether our response to affective vocalizations is specific to those generated by humans or more universal, triggered by emotionally matched vocalizations generated by other species. Here, we used functional magnetic resonance imaging in normal participants to measure cerebral activity during auditory stimulation with affectively valenced animal vocalizations, some familiar (cats) and others not (rhesus monkeys). Positively versus negatively valenced vocalizations from cats and monkeys elicited different cerebral responses despite the participants' inability to differentiate the valence of these animal vocalizations by overt behavioural responses. Moreover, the comparison with human non-speech affective vocalizations revealed a common response to the valence in orbitofrontal cortex, a key component on the limbic system. These findings suggest that the neural mechanisms involved in processing human affective vocalizations may be recruited by heterospecific affective vocalizations at an unconscious level, supporting claims of shared emotional systems across species.

Free-ranging rhesus monkeys spontaneously individuate and enumerate small numbers of non-solid portions.

Fundamental questions in cognitive science concern the origins and nature of the units that compose visual experience. Here, we investigate the capacity to individuate and store information about non-solid portions, asking in particular whether free-ranging rhesus monkeys (Macaca mulatta) quantify portions of a non-solid substance presented in discrete pouring actions. When presented with portions of carrot pieces poured from a cup into opaque boxes, rhesus picked the box with the greatest number of portions for comparisons of 1 vs. 2, 2 vs. 3, and 3 vs. 4, but not for comparisons of 4 vs. 5 and 3 vs. 6. Additional experiments indicate that rhesus based their decisions on both the number of portions and the total amount of food. These results show that the capacity to individuate non-solid portions is not unique to humans, and does not depend on structures of natural language. Further, the fact that rhesus' ability to represent non-solid portions is constrained by the same 4-item limit typically ascribed to the system of parallel individuation that operates over solid objects suggests that the visual system recruits common working memory processes for retaining information about solid objects and non-solid portions. We discuss our results with respect to theories of visual processing, as well as to the role that the human language faculty may have played in both the evolution and development of quantification.

The evolution of the music faculty: a comparative perspective.

We propose a theoretical framework for exploring the evolution of the music faculty from a comparative perspective. This framework addresses questions of phylogeny, adaptive function, innate biases and perceptual mechanisms. We argue that comparative studies can make two unique contributions to investigations of the origins of music. First, musical exposure can be controlled and manipulated to an extent not possible in humans. Second, any features of music perception found in nonhuman animals must not be part of an adaptation for music, and must rather be side effects of more general features of perception or cognition. We review studies that use animal research to target specific aspects of music perception (such as octave generalization), as well as studies that investigate more general and shared systems of the mind/brain that may be relevant to music (such as rhythm perception and emotional encoding). Finally, we suggest several directions for future work, following the lead of comparative studies on the language faculty.

Rhesus monkeys correctly read the goal-relevant gestures of a human agent.

When humans point, they reveal to others their underlying intent to communicate about some distant goal. A controversy has recently emerged based on a broad set of comparative and phylogenetically relevant data. In particular, whereas chimpanzees (Pan troglodytes) have difficulty in using human-generated communicative gestures and actions such as pointing and placing symbolic markers to find hidden rewards, domesticated dogs (Canis familiaris) and silver foxes (Urocyon cinereoargenteus) readily use such gestures and markers. These comparative data have led to the hypothesis that the capacity to infer communicative intent in dogs and foxes has evolved as a result of human domestication. Though this hypothesis has met with challenges, due in part to studies of non-domesticated, non-primate animals, there remains the fundamental question of why our closest living relatives, the chimpanzees, together with other non-human primates, generally fail to make inferences about a target goal of an agent's communicative intent. Here, we add an important wrinkle to this phylogenetic pattern by showing that free-ranging rhesus monkeys (Macaca mulatta) draw correct inferences about the goals of a human agent, using a suite of communicative gestures to locate previously concealed food. Though domestication and human enculturation may play a significant role in tuning up the capacity to infer intentions from communicative gestures, these factors are not necessary.

Nonhuman primates prefer slow tempos but dislike music overall.

Human adults generally find fast tempos more arousing than slow tempos, with tempo frequently manipulated in music to alter tension and emotion. We used a previously published method [McDermott, J., & Hauser, M. (2004). Are consonant intervals music to their ears? Spontaneous acoustic preferences in a nonhuman primate. Cognition, 94(2), B11-B21] to test cotton-top tamarins and common marmosets, two new-World primates, for their spontaneous responses to stimuli that varied systematically with respect to tempo. Across several experiments, we found that both tamarins and marmosets preferred slow tempos to fast. It is possible that the observed preferences were due to arousal, and that this effect is homologous to the human response to tempo. In other respects, however, these two monkey species showed striking differences compared to humans. Specifically, when presented with a choice between slow tempo musical stimuli, including lullabies, and silence, tamarins and marmosets preferred silence whereas humans, when similarly tested, preferred music. Thus despite the possibility of homologous mechanisms for tempo perception in human and nonhuman primates, there appear to be motivational ties to music that are uniquely human.

A paradox in the evolution of primate vocal learning.

The importance of auditory feedback in the development of spoken language in humans is striking. Paradoxically, although auditory-feedback-dependent vocal plasticity has been shown in a variety of taxonomic groups, there is little evidence that our nearest relatives--non-human primates--require auditory feedback for the development of species-typical vocal signals. Because of the apparent lack of developmental plasticity in the vocal production system, neuroscientists have largely ignored the neural mechanisms of non-human primate vocal production and perception. Recently, the absence of evidence for vocal plasticity from developmental studies has been contrasted with evidence for vocal plasticity in adults. We argue that this new evidence makes non-human primate vocal behavior an attractive model system for neurobiological analysis.

Vervet monkeys and humans show brain asymmetries for processing conspecific vocalizations, but with opposite patterns of laterality.

A robust finding in the human neurosciences is the observation of a left hemisphere specialization for processing spoken language. Previous studies suggest that this auditory specialization and brain asymmetry derive from a primate ancestor. Most of these studies focus on the genus Macaca and all demonstrate a left hemisphere bias. Due to the narrow taxonomic scope, however, we lack a sense of the distribution of this asymmetry among primates. Further, although the left hemisphere bias appears mediated by conspecific calls, other possibilities exist including familiarity, emotional relevance and more general acoustic properties of the signal. To broaden the taxonomic scope and test the specificity of the apparent hemisphere bias, we conducted an experiment on vervets (Cercopithecus aethiops)-a different genus of old world monkeys and implemented the relevant acoustic controls. Using the same head orienting procedure tested with macaques, results show a strong left ear/right hemisphere bias for conspecific vocalizations (both familiar and unfamiliar), but no asymmetry for other primate vocalizations or non-biological sounds. These results suggest that although auditory asymmetries for processing species-specific vocalizations are a common feature of the primate brain, the direction of this asymmetry may be relatively plastic. This finding raises significant questions for how ontogenetic and evolutionary forces have impacted on primate brain evolution.