The early developing capacity of human learners to seek out reliable informants, initiate pedagogical episodes, and monitor and redirect ongoing instruction is critical to understanding humans' remarkable capacity for cumulative culture.

Darwinism is a principle of biological continuity. This commentary argues against any claim of discontinuity between humans and other animals that must be based on absence of evidence. Instead, we offer additional examples of active teaching by chimpanzees.

As Kline envisages, there is an important relationship between cultural attraction and teaching. The very function of teaching is to make the content taught an attractor. Teaching, moreover, typically fulfills its function by exploiting a variety of factors of cultural attraction that help make its content learnable and teachable.

How far can teaching methods go to enhance learning? Optimal methods of teaching have been considered in research on supervised and unsupervised learning. Locally optimal methods are usually hybrids of teaching and self-directed approaches. The costs and benefits of specific methods have been shown to depend on the structure of the learning task, the learners, the teachers, and the environment.

Insufficient attention has been paid to the use of robots in classrooms. Robot “teachers” are being developed, but because Kline ignores such technological developments, it is not clear how they would fit within her framework. It is argued here that robots are not capable of teaching in any meaningful sense, and should be deployed only as educational tools.

We appreciate and endorse Kline's ethological taxonomy and its application. However, the definition of teaching she presents is problematic, as it replaces mentalistic intent with intention on the part of natural selection. We discuss problems with the strict adaptationist view and suggest instead that the five forms of teaching presented in the taxonomy may constitute exaptations rather than adaptations.

We address three issues: (1) There is a need for a comprehensive multidisciplinary understanding of teaching; (2) teaching is a natural cognitive ability for humans; and (3) there is a need to incorporate the mentalistic and cultural approaches to teaching. We suggest certain research studies that can help deepen our understanding of the cognition of teaching.

We argue that direct active teaching in humans exhibits at least two properties (open-endedness and content opacity) that make the recognition of teaching episodes without ostension untenable. Thus, while we welcome Kline's functional approach to the analysis of teaching, we think that she ignores important features of the socio-environmental niche in which human teaching likely evolved.

Kline does an admirable job of extending the functionalist framework developed by comparative researchers to help understand the function and form of human teaching. Functionalist approaches consider the adaptive value and underlying mechanisms of behaviour as separate but complementary questions, avoiding the conflation of ultimate and proximate explanations that has long hindered research on teaching and other forms of cooperation.

The collection of commentaries expands an already extensive field of research on teaching, and contributes new questions, techniques, and strengths to the evolutionary approach proposed in the target article. In my response, I show how reconciling multiple levels of explanation – mechanistic, ontogenetic, phylogenetic, and functional – enables researchers to build a more integrated, interdisciplinary approach to the study of teaching in humans and other animals.

In The Cognitive-Emotional Brain (Pessoa 2013), I describe the many ways that emotion and cognition interact and are integrated in the brain. The book summarizes five areas of research that support this integrative view and makes four arguments to organize each area. (1) Based on rodent and human data, I propose that the amygdala's functions go beyond emotion as traditionally conceived. Furthermore, the processing of emotion-laden information is capacity limited, thus not independent of attention and awareness. (2) Cognitive-emotional interactions in the human prefrontal cortex (PFC) assume diverse forms and are not limited to mutual suppression. Particularly, the lateral PFC is a focal point for cognitive-emotional interactions. (3) Interactions between motivation and cognition can be seen across a range of perceptual and cognitive tasks. Motivation shapes behavior in specific ways – for example, by reducing response conflict or via selective effects on working memory. Traditional accounts, by contrast, typically describe motivation as a global activation independent of particular control demands. (4) Perception and cognition are directly influenced by information with affective or motivational content in powerful ways. A dual competition model outlines a framework for such interactions at the perceptual and executive levels. A specific neural architecture is proposed that embeds emotional and motivational signals into perception and cognition through multiple channels. (5) A network perspective should supplant the strategy of understanding the brain in terms of individual regions. More broadly, in a network view of brain architecture, “emotion” and “cognition” may be used as labels of certain behaviors, but will not map cleanly into compartmentalized pieces of the brain.

We emphasize the importance of a neuroevolutionary perspective in moving beyond the cognition-emotion dichotomy. Cognitive behavior depends on cortical structures firmly rooted in the emotional brain from which they have evolved. As such, there cannot be cognition without emotion. Endocrine regulation of amygdala connectivity, a neural “switch” between impulsivity and deliberation, further underscores the phylogenetic impossibility of a cognition-emotion dichotomy.

I comment on Pessoa's (2013) idea that the interaction between emotion and cognition cannot be reduced to mutual interference. As an example that bolsters Pessoa's position, I discuss the effects of happy and sad moods on discourse and sentence comprehension. I distinguish between the effects of moods elicited without participants' knowledge (incidental) and moods elicited with participants' contribution (constructed).

The target article is a timely exposition on the impact of how emotion and cognition interact, a specifically important issue in surprise research. Psychologists debate whether disconfirmed expectations or sense-making processes determine surprise levels experienced for an event. We posit that, in surprise, cognition and emotion are intertwined, making it an interesting test case for the proposals in this article.

Pessoa's The Cognitive-Emotional Brain (2013) is an integrative approach to neuroscience that complements other developments in cognitive science, especially enactivism. Both accept complexity as essential to mind; both tightly integrate perception, cognition, and emotion, which enactivism unifies in its foundational concept of sense-making; and both emphasize that the spatial extension of mental processes is not reducible to specific brain regions and neuroanatomical connectivity. An enactive neuroscience is emerging.

Integration of cognition and emotion, discussed by Pessoa in The Cognitive-Emotional Brain (2013), is further illustrated by music. In music, I argue, this integration begins during mental control of the actions by which musical sounds are produced. Many emotional reactions to the music we hear are also strongly related to the actions by which musical sounds are produced. Studies involving music can further illuminate the integration of emotion and control of action throughout behavior.

We argue that although the “dual competition” model is useful when considering interactions between emotional and neutral stimuli, it fails to account for the influence of emotional arousal on perceptual or goal-directed behavior involving neutral stimuli. We present the “arousal-biased competition” framework as an alternative that accounts for both scenarios.

Pessoa (2013) makes a compelling case for conceiving of emotion and cognition as deeply integrated processes in the brain. We will begin our commentary by asking what implications this view of the brain has for an ontology of cognition – a theory of what cognition is and what cognitive processes exist. We will suggest that Pessoa's book, The Cognitive-Emotional Brain, provides strong support for an embodied theory of cognition. We end our commentary by offering some speculation about how Pessoa's arguments naturally extend to social cognition.

Pessoa's (2013) dual competition model outlines a framework for how cognition and emotion interact at the perceptual levels and provides evidence within the field of neuroscience to support this new perspective. Here, I discuss how behavioral work fares with this new model and how visual detection is influenced by information with affective or motivational content.

Contrary to Greene's dual-process theory of moral judgment (Greene 2013), this commentary suggests that the network view of the brain proposed by Pessoa, in which emotion and cognition may be used as labels in the context of certain behaviors, but will not map clearly into compartmentalized pieces of the brain, could represent a better explanation of the rationale behind people's moral behavior.