Commentaries surrounding Skinner (1984) were re-examined and applied to Hull et al. Hull et al. were found to address many of these concerns by paying attention to neuroscience, by providing some discussion of the origins of behavior, and by forwarding a deterministic account that may prove as revolutionary as that of Copernicus and Darwin.

Dichotomous definitions of culture and language do not generate productive questions. Instead, more progress can be made by identifying components of each that other animals might plausibly possess. The evolutionary, ecological approach advocated by Rendell and Whitehead holds great promise for helping us to understand the conditions under which natural selection can favor similar capacities in differently organized brains.

We suggest that consciousness (C) should be addressed as a multilevel concept. We can provisionally identify at least three, rather than two, levels: Gray's system should relate at least to the lowest of these three levels. Although it is unlikely to be possible to develop a behavioural test for C, it is possible to speculate as to the evolutionary advantages offered by C and how C evolved through succeeding levels. Disturbances in the relationships between the levels of C could underlie mental illness, especially schizophrenia.

Selectionist models date back to Empedocles in Ancient Greece. The novelty of Darwinian selection is that it is able to produce adaptively valuable things without being sensitive to adaptive value. Darwin achieved this result by a restriction of environmental feedback to the replicative process. Immune system selection definitely does not respect this restriction, and it is doubtful whether operant learning does.

In O'Regan & Noë's (O&N's) account for the phenomenal experience of seeing, awareness is equated to what is within the current focus of attention. They find no place for a distinction between phenomenal and access awareness. In doing so, they essentially present a dualistic solution to the mind-brain problem, and ignore that we do have phenomenal experience of what is outside the focus of attention.

Sensory motor contingencies belong to a functionalistic framework. Functionalism does not explain why and how objective functional relations produce phenomenal experience. O'Regan & Noë (O&N) as well as other functionalists do not propose a new ontology that could support the first person subjective phenomenal side of experience.

Cetacean cultural transmission is associated with lengthened postmenopausal life histories and relatively stable matrilineal social structures. Although Homo erectus was not marine adapted, broadly comparable selection pressures, life history profiles, and social structures can be inferred.

Gray's target article presents a model of consciousness that includes several ideas similar to those developed over the past century to explain how sensorimotor information is interpreted by the nervous system. This commentary discusses these ideas and introduces some additional hypotheses, also derived from sensorimotor investigations, that might help improve Gray's model.

The idea that similar selective processes operate in gene-based evolution, immunology, and operant psychology provides an intuitively appealing metaphor. This idea also isolates questions that operant psychologists should ask and makes some empirical predictions. However, the idea currently lacks the detail needed to precisely separate it from some plausible alternatives. This sort of thinking is the kind that operant psychologists should do if operant theorizing is to survive the competition among ideas.

Cetaceans are likely candidates for social learning and culture. Meager experimental evidence suggests that some cetaceans possess the requisite cognitive skills for social learning. Equally sparse ethnographic data provide clues about possible outcomes of social learning. Although the available evidence is consistent with the notion of culture in cetaceans, caution is warranted due to the many gaps in the data.

Behavioral theorists such as B. F. Skinner have argued that the thesis of selection by consequences applies to behavior just as much as to morphology. This commentary specifically examines certain respects in which the thesis of “selection by consequences” applies to the development of ontogenic operant behavior.

Gray's account of a brain mechanism for generating the contents of consciousness is incomplete. Adaptive advantages of conscious functioning need to be sought within the first-person affective sensation motivating flexibly goal-directed actions, as in Humphrey's sensory feedback theory.

Sensorimotor contingencies are certainly of great importance for perception but they are no substitute for the internal representation of perceived information. I argue that internal, non-iconic representations of perceptions must, and do, exist and that sensorimotor contingencies are an integral part of them. Further, I argue that mastery of the sensory apparatus or environment is not a prerequisite for perception and that perception is possible in the absence of any control over the perceptual process.

We disagree with two of Rendell and Whitehead's assertions. Culture may be an ancestral characteristic of terrestrial cetacean ancestors; not derived via marine variability, modern cetacean mobility, or any living cetacean social structure. Furthermore, evidence for vocal behavior as culture, social stability, and cognitive ability, is richer in birds than Rendell and Whitehead portray and comparable to that of cetaceans and primates.

What makes conscious experiences necessary for in- formation processing or behaviour (no one knows)? Would it be easier first to divide consciousness into different levels (probably not)? Is consciousness tied to information processing or brain states (no one knows)? Would the target article's comparator be improved by adding a continuously adjusting feedback (probably not)?

Instead of using only one notion of selection I argue for a broader typology of different types of selection. Three such types are differentiated, namely simple one-step selection, iterated one-step selection, and multi-step selection. It is argued that this more general and more inclusive typology might face more effectively the possible challenges of a general account of selection.

I argue that O'Regan & Noë's (O&N's) theory is in a no better position than any other theory to solve the “hard problem” of consciousness. Getting rid of the explanatory gap by exchanging sensorimotor contingencies for neural representations is an illusion.

Theories that make action central to perception are plausible, though largely untried, for space perception. However, explaining object recognition, and high-level perception generally, will require reference to representations of the world in some form. Nonetheless, action is central to cognition, and explaining high-level perception will be aided by integrating an understanding of action with other aspects of perception.

The use of anecdotes is not a viable research strategy to study animal culture. Social learning processes can often be documented with careful quantitative analyses of observational data. Unfortunately, suggestions that killer whales engage in teaching are entirely based on subjective interpretations of qualitative observations. Thus, “evidence” of teaching in killer whales cannot be used to argue for the occurrence of culture in marine mammals.

I argue that population-level selection does not necessarily have to be invoked to explain the polymorphism at the MHC locus. I argue that the authors' attempt to model operant conditioning in Darwinian terms faces a serious problem. Depending on how many operant responses we take to comprise a sequence, different conclusions about whether or not evolution is occurring in an operant lineage will be reached.