The three introductory questions posed by Hobson et al. point toward further investigations of cellular, circuit, and systems mechanisms involved in cognitive function that include the effect of CNS-state related modulatory systems on these mechanisms.

[Hobson et al.]

Page's manifesto makes a case for localist representations in neural networks, one of the advantages being ease of interpretation. However, even localist networks can be hard to interpret, especially when at some hidden layer of the network distributed representations are employed, as is often the case. Hidden Markov models can be used to provide useful interpretable representations.

The hypothesis of two separate reasoning systems, one subserving individual goals and the other our genes, is theoretically implausible and not supported by the data. As an alternative, I propose a single system for analytical reasoning backed up by simple mechanisms for the selection of relevant information. This system can generate normative behavior as well as systematic deviations from it.

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 comprehension problem in Broca's patients does not stem from an inability to represent argument traces. There can be good comprehension in the presence of (object) traces and impaired comprehension can result in constructions where there are no (object) argument traces. This leads to an alternative understanding of Broca's comprehension, one that places the locus of the impairment in an inability to construct syntactic representation on time.

Although Norris, McQueen & Cutler have provided convincing evidence that there is no need for contributions from the lexicon to phonetic processing, their simplification of the communication between levels comes at a cost to the processes themselves. Although their arrangement may ultimately prove correct, its validity is not due to a successful application of Occam's razor.

Stanovich & West (S&W) are wrong to think that all “reject-the-norm” theorists simply wish to reduce the normative/descriptive gap. They have misunderstood Issac Levi's reasons for rejecting Tversky and Kahneman's normative assumptions in the “base-rate” experiments. In their discussion of the taxicab experiment, (S&W) erroneously claim that subjects' responses indicate whether they have reasoned in accordance with Bayesian principles or not.

Page's “localist” code, a population code with occasional, maximally firing elements, does not seem to us usefully or testably different from sparse population coding. Some of the evidence adduced by Page for his proposal is not actually evidence for it, and coding by maximal firing is challenged by lower firing observed in neuronal responses to natural stimuli.

The notion of simulation in dreaming of threat recognition and avoidance faces difficulties deriving from (1) some typical characteristics of dream artifacts (some “surreal,” some not) and (2) metaphysical issues involving the need for some representation in the theory of a perspective subject making use of the artifact.

[Hobson et al.; Revonsuo]

This commentary focuses on the larger implications of Grodzinsky's hypothesis. Although Grodzinsky argues persuasively that the syntactic comprehension deficits in Broca's aphasia involve mainly an inability to comprehend sentences requiring a transformational movement of phrasal constituents, his larger claim for a distinct and dedicated “language organ” in the left hemisphere is much less tenable.

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.

Models are not adequately evaluated simply by whether they capture the data, after the fact. Other criteria are needed. One criterion is parsimony; but utility and generality are at least as important. Even with respect to parsimony, however, the case against feedback is not as straightforward as Norris et al. present it. We use feedback consistency effects to illustrate these points.

Stanovich & West (S&W) distinguish between evolutionary rationality and normative rationality, and System 1 and System 2 mental processes. They hold that the main function of System 2 has to do with normative and not evolutionary rationality. We ask how System 2 could then be an adaptation, especially given S&W's own work on individual differences.

In this manuscript Hobson et al. propose a model exploring qualitative differences between the three states of consciousness, waking, NREM sleep, and REM sleep, in terms of state-related brain activity. The model consists of three factors, each of which varies along a continuum, creating a three-dimensional space: activation (A), information flow (I), and mode of information processing (M). Hobson has described these factors previously (1990; 1992a). Two of the dimensions, activation and modulation, deal directly with subcortical influences upon cortical structures – the reticular activation system, with regard to the activation dimension and the locus coeruleus and the pontine raphe neuclei, with regard to the modulation dimension. The focus of this review is a further exploration of the interaction between dreaming and the cortical and subcortical structures relevant to REM sleep eye movements.

[Hobson et al. ]

An elaboration on some loose ends in Grodzinsky's analysis shows that data from the field of aphasia contribute to the formulation of theoretical linguistic principles, and provides extra arguments in favor of Grodzinsky's claim that linguistic theory is the best tool for the investigation of aphasia. This illustrates and emphasizes the importance of communication between researchers in the field of (Broca's) aphasia and of theoretical linguistics.

The central thesis of our target article is that feedback is never necessary in spoken word recognition. In this response we begin by clarifying some terminological issues that have led to a number of misunderstandings. We provide some new arguments that the feedforward model Merge is indeed more parsimonious than the interactive alternatives, and that it provides a more convincing account of the data than alternative models. Finally, we extend the arguments to deal with new issues raised by the commentators such as infant speech perception and neural architecture.

The commentators have raised some interesting issues but none question the viability of a localist approach to connectionist modelling. Once localist models are properly defined they can be seen to exhibit many properties relevant to the modelling of both psychological and brain function. They can be used to implement exemplar models, prototype models and models of sequence memory and they form a foundation upon which symbolic models can be constructed. Localist models are insensitive to interference and have learning rules that are biologically plausible. They have more explanatory value than their distributed counterparts and they relate transparently to a number of classic mathematical models of behaviour.

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?

A general conceptual framework for large-scale neocortical dynamics based on data from many laboratories is applied to a variety of experimental designs, spatial scales, and brain states. Partly distinct, but interacting local processes (e.g., neural networks) arise from functional segregation. Global processes arise from functional integration and can facilitate (top down) synchronous activity in remote cell groups that function simultaneously at several different spatial scales. Simultaneous local processes may help drive (bottom up) macroscopic global dynamics observed with electroencephalography (EEG) or magnetoencephalography (MEG).

A local/global dynamic theory that is consistent with EEG data and the proposed conceptual framework is outlined. This theory is neutral about properties of neural networks embedded in macroscopic fields, but its global component makes several qualitative and semiquantitative predictions about EEG measures of traveling and standing wave phenomena. A more general “metatheory” suggests what large-scale quantitative theories of neocortical dynamics may be like when more accurate treatment of local and nonlinear effects is achieved.

The theory describes the dynamics of excitatory and inhibitory synaptic action fields. EEG and MEG provide large-scale estimates of modulation of these synaptic fields around background levels. Brain states are determined by neuromodulatory control parameters. Purely local states are dominated by local feedback gains and rise and decay times of postsynaptic potentials. Dominant local frequencies vary with brain region. Other states are purely global, with moderate to high coherence over large distances. Multiple global mode frequencies arise from a combination of delays in corticocortical axons and neocortical boundary conditions. Global frequencies are identical in all cortical regions, but most states involve dynamic interactions between local networks and the global system. EEG frequencies may involve a “matching” of local resonant frequencies with one or more of the many, closely spaced global frequencies.

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.