The explicit system for action selection integrates emotional information with the higher-order cognitive processes which culminate in the language system. Even the basic feels of emotion are what they are because they are integrated into the higher cognitive processes. The relation between emotion and intelligence would become increasingly important as the focus of brain science shifts to the integrative function of the prefrontal lobe.

Rolls defines emotion as innate reward and punishment. This could not explain our results showing that people learn faster in a negative mood. We argue that what people know about their world affects their emotional state. Negative emotion signals a failure to predict negative reward and hence prompts learning to resolve the ignorance. Thus what you don't know affects how you feel.

Rolls shares important data on hunger, thirst, sexuality, and learned behaviors, but is it pertinent to understanding the fundamental nature of emotionality? Important as such work is for understanding the motivated behaviors of animals, Rolls builds a constructivist theory of emotions and primary-process affective consciousness without considering past evidence on specific types of emotional tendencies and their diverse neural substrates.

Rolls's proposal that the amygdala is critical for the association of visual objects with reward is not consistent with recent ablation evidence. Stimulus-reward association learning is more likely to depend on basal forebrain efferents to the inferior temporal cortex, some of which pass through the amygdala. It is more likely that the amygdala is involved in rapid modulation of stimulus reward value.

Rolls attributes to consciousness the functions of reflection, planning, and error-correction. Neuropsychologically grounded cybernetic theory provides an analogous, broader conceptualization: consciousness constructs goals (and plans), alters the valence of goal-related phenomena, registers error-signals, and explores unexpected circumstances (reconfiguring goals and plans as necessary). Consciousness plays a fundamental unrecognized ontological role, as well, conferring the status of “discriminable object” on select aspects of otherwise indeterminate “being.”

Rolls demonstrates how reward/punishment systems are key mediators of cognitive appraisal, and this suggests a fundamental, causal role for emotion in thought and behaviour. However, this causal role for emotion seems to drop out of Rolls's model of consciousness, to be replaced by the old idea that emotion is essentially epiphenomenal. We suggest a modification to Rolls's model in which cognition and emotion activate each other reciprocally, both in appraisal and consciousness, thus allowing emotion to maintain its causal status where it matters most.

The “reward” systems described by Rolls are systems for drive-reinforced associations of contact and distant stimuli and not for emotional behaviours. The neural systems delineated may be associated with distinct categories of “affect,” namely “hedonic feelings,” “moods,” and “emotions.” Awareness of these affects requires external perceptual as well as internal feedback. Levels of feedback in evolution and development suggest sensory qualia may not require language.

This commentary argues for the inclusion of the neurophysiological arousal concept to help understanding the brain mechanisms of emotions and reward and the cognitive mechanisms involved.

Despite its explanatory value, the “higher order linguistic thought” model comes up short as an account of the felt aspect of motivational states.

Rolls's mechanistic account of emotion can help to bridge a rift within the field of evolutionary psychology. One side of the rift emphasizes the importance of innate psychological mechanisms that evolved to solve specific problems encountered in the ancestral environment. The other side emphasizes learning, development, and culture as open-ended evolutionary processes in their own right. Rolls shows how these two views can be reconciled, allowing a productive middle ground to be explored.

There are many advantages to defining emotions as states elicited by reinforcers, with the states having a set of different functions. This approach leads towards an understanding of the nature of emotion, of its evolutionary adaptive value, and of many principles of brain design. It also leads towards a foundation for many of the processes that underlie evolutionary psychology and behavioral ecology. It is shown that recent as well as previous evidence implicates the amygdala and orbitofrontal cortex in positive as well as negative emotions. The issue of why emotional states feel like something is part of the much larger problem of phenomenal consciousness. It is argued that thinking about one's own thoughts would have adaptive value by enabling first order linguistic thoughts to be corrected. It is suggested that reflecting on and correcting one's own thoughts and plans would feel like something, and that phenomenal consciousness may occur when this type of monitoring process is taking place.

The conceptual and investigative tools for the analysis of social behavior can be expanded by integrating biological theory, control systems theory, and Pavlovian conditioning. Biological theory has focused on the costs and benefits of social behavior from ecological and evolutionary perspectives. In contrast, control systems theory is concerned with how machines achieve a particular goal or purpose. The accurate operation of a system often requires feed-forward mechanisms that adjust system performance in anticipation of future inputs. Pavlovian conditioning is ideally suited to subserve this function in behavioral systems. Pavlovian mechanisms have been demonstrated in various aspects of sexual behavior, maternal lactation, and infant suckling. Pavlovian conditioning of agonistic behavior has been also reported, and Pavlovian processes may likewise be involved in social play and social grooming. Several further lines of evidence indicate that Pavlovian conditioning can increase the efficiency and effectiveness of social interactions, thereby improving their cost/benefit ratio. We extend Pavlovian concepts beyond the traditional domain of discrete secretory and other physiological reflexes to complex real-world behavioral interactions and apply abstract laboratory analyses of the mechanisms of associative learning to the daily challenges animals face as they interact with one another in their natural environments.

If biologists are going to incorporate learning into theories of animal behavior, why not go all the way and incorporate the enormous literatures on Pavlovian conditioning, plus those on operant and observational learning?

Domjan, Cusato & Villarreal's target article is reviewed in the context of historical difficulty for learning studies in discriminating between learned and unlearned components of behavior. The research surveyed in the target article meets the criterion of differentiating between some learned and the unlearned aspects of social behavior, with Pavlovian conditioning shown repeatedly as a route by which reproductive and aggressive behavior is modulated.

Some aspects of play may be explained by Pavlovian learning processes, but others are not so easily handled. Especially when there is a chance that specific actions can be misinterpreted; animals alter their behavior to reduce the likelihood that this will occur. The flexibility and fine-tuning of play make it an ideal candidate for comparative and evolutionary cognitive studies.

It is suggested that the control-system theory of Domjan et al. restates in engineering vocabulary the primary thesis of law-of-effect theories: namely, that classical-conditioning arrangements may involve CR-contingent reinforcement. The research cited by Domjan et al. is relevant to the idea that classical conditioning is an adaptive process, but is irrelevant to their control-system theory.

The proposed heavy dependence on Pavlovian conditioning to account for social behavior confounds phylogenically and ontogenically selected behavior patterns and ignores the extension of the principle of selection by consequences from biological to learning theory. Instead of acknowledging operant relations, Domjan et al. construct vaguely specified mechanisms based upon anticipatory cost-benefit considerations that are not supported by the Pavlovian conditioning literature.

A proper evaluation of the biological significance of Pavlovian conditioning requires consideration of performance mechanisms. Domjan et al.'s definition of net benefit is simplistic, and their model promotes convergence in behaviour, ignoring the possibility of innovation.

We applaud Domjan et al. for providing an elegant account of Pavlovian feed-forward mechanisms in social behavior that eschews the pitfall of purposivism. However, they seem to imply that they have provided a complete account without provision for operant conditioning. We argue that operant conditioning plays a central role in social behavior, giving examples from fish and infant behavior.

Domjan et al.'s model of how Pavlovian processes regulate social interaction can be extended to social learning, where an individual learns about the value of events, objects, or actions from information provided by another. The conditioned properties of a particular social partner, following from a history of interactions with that partner, can modulate the efficiency and specificity of social learning.