Executive functions and the frontal lobes: a conceptual view

Several problems in understanding executive functions and their relationships to the frontal lobes are discussed. Data are then presented from several of our studies to support the following statements: (1) the examination of patients with focal frontal lobe lesions is a necessary first step in defining the relation of executive functions to the frontal lobes; (2) there is no unitary executive function. Rather, distinct processes related to the frontal lobes can be differentiated which converge on a general concept of control functions; (3) a simple control-automatic distinction is inadequate to explain the complexity of control-automatic processes; (4) the distinction between complex and simple tasks cannot explain the differences in functions between the frontal lobes and other brain regions; and (5) the most important role of the frontal lobes may be for affective responsiveness, social and personality development, and self-awareness and unconsciousness. Received: 31 March 1999 / Accepted: 23 July 1999     

Cognitive control, hierarchy, and the rostro-caudal organization of the frontal lobes

Cognitive control supports flexible behavior by selecting actions that are consistent with our goals and appropriate for our environment. The prefrontal cortex (PFC) has an established role in cognitive control, and research on the functional organization of PFC promises to contribute to our understanding of the architecture of control. A recently popular hypothesis is that the rostro-caudal axis of PFC supports a control hierarchy whereby posterior-to-anterior PFC mediates progressively abstract, higher-order control. This review discusses evidence for a rostro-caudal gradient of function in PFC and the theories proposed to account for these results, including domain generality in working memory, relational complexity, the temporal organization of behavior and abstract representational hierarchy. Distinctions among these frameworks are considered as a basis for future research.     

Information search following damage to the frontal lobes

Five adult patients with focal brain damage in different areas of the frontal lobes and five matched controls were administered a 20 Questions test. Other tests of planning, problem-solving, and decision-making were also administered to frontal lobe patients. The results indicate that damage to the frontal lobe impairs the ability to impose and utilize hierarchical organization in mental representation.     

Adaptive versus veridical decision making and the frontal lobes

Adaptive decision making and veridical decision making are based on different mechanisms. Veridical decision making is based on the identification of the correct response, which is intrinsic to the external situation and is actor-independent. Adaptive decision making is actor-centered and is guided by the actor's priorities. The prefrontal cortex is particularly critical for adaptive decision making and less so for veridical decision making. However, most experimental procedures used in cognitive psychology and neuropsychology focus on veridical decision making and ignore adaptive decision making. Innovative experimental procedures are required to characterize the contribution of the prefrontal cortex to adaptive decision making. We have designed a prototype for such procedures, the Cognitive Bias Task, and present the novel findings generated by this task.     

Aging effects on memory encoding in the frontal lobes

Functional magnetic resonance imaging (fMRI) was used to compare frontal-lobe activation in younger and older adults during encoding of words into memory. Participants made semantic or nonsemantic judgments about words. Younger adults exhibited greater activation for semantic relative to nonsemantic judgments in several regions, with the largest activation in the left inferior frontal gyrus. Older adults exhibited greater activation for semantic judgments in the same regions. but the extent of activation was reduced in left prefrontal regions. In older adults, there was a significant association between behavioral tests of declarative and working memory and extent of frontal activation. These results suggest that age-associated decreases in memory ability may be due to decreased frontal-lobe contributions to the initial encoding of experience.     

Inhibition and the control of behavior. From Gall to Freud via Phineas Gage and the frontal lobes.

In his On the Functions of the Brain and Each of Its Parts, Franz Joseph Gall proposed that significant behaviors resulted from certain independent, irreducible, and fundamental faculties or propensities. There seems to be nothing in his system for explaining how two or more faculties interact, especially how one comes to predominate over another, other than the implicit notion that the stronger will be manifested. This peculiarity of Gall's system is consistent with the predominately "excitatory" physiology of his time in which there was no place for an independent inhibitory process. Despite the experimental demonstration of inhibitory phenonema, the sensory-motor physiological theories in the last three quarters of the 19th century were similarly deficient. The lack is seen most clearly in the experimental and clinical literature on frontal lobe function, especially in relation to the kinds of changes seen in the Gage case. The deficiencies of sensory-motor physiology are also to be found in Freud's model of the mental apparatus and in his concept of inhibition via a lateral cathexis. In the paper an attempt is made to trace the historical sequence of these ideas and to make their experimental and theorectical basis explicit.     

The role of the frontal lobes in the regulation of cognitive development.

Between the ages of 1.5 and 5 years, and again between the ages of 5 and 10 years, a sequence of changes takes place in children's behavior which indicates a fundamental reorganization of their attentional, executive, and self-reflexive processes. In the present article, these changes are summarized, and evidence is adduced to support the claims (1) that these changes are frontally mediated and (2) that the underlying mechanism that generates them is similar to the one that generates the changes in EEG coherence during the same time period. The psychological model that has been hypothesized to explain the cycles of cognitive development (Case, 1992) is then compared to the physiological model that has been proposed to explain cycles of EEG development (Thatcher, 1992). It is shown that the two models are complementary, both in the underlying developmental sequence that they postulate and in the recursive dynamic they propose for producing movement through this sequence. A number of implications and predictions are derived, which follow from the proposition that the two sets of changes are different manifestations of a common underlying process.     

The role of internal inhibition and the frontal lobes in child development

The theory that children show a predominance of excitation over internal inhibition and that this is due to inadequate development of the frontal lobes is explored and extended. Experimental results show that children resemble leucotomised monkeys in their difficulty in overcoming an initial preference in discrimination tasks.     

A selective and critical review of neuropsychological deficits and the frontal lobes

Presumptions about the functions of the frontal lobes, and the sensitivity and specificity of certain tests to measure frontal lobe functions, are having a substantial influence on both clinical and research conclusions. In this paper the authors examine the details of the studies that have contributed to these presumptions, and find that the evidence to support these conclusions is weak. A detailed evaluation of the evidence relating to the Wisconsin Card Sorting Test and the Thurstone Word Fluency Test is also presented. Finally, the development of the belief that frontal lobe functions can be specifically measured is reviewed. The authors of this paper conclude that the bewildering array of deficits attributed to frontal lesions still seems to prevail.     

Strategy application disorder: the role of the frontal lobes in human multitasking

Strategy application disorder is a term used to describe a pattern of deficits, usually associated with frontal lobe dysfunction, where people show disorganisation, absentmindedness and problems with planning and decision making in everyday life despite normal performance on traditional neuropsychological tests. It is argued that the prototypical situation which presents problems for these cases are those which require multitasking, and although good cases are rare in the literature, those that do exist show a characteristic neuropsychological pattern. Moreover, this pattern is confirmed in recent group studies of multitasking and of the relationship between multitasking tests (such as the Six Element Test), failures in everyday life and other neuropsychological measures. At present the evidence suggests that the potential frontal brain regions most implicated in multitasking are the anterior cingulate; B.A. 10 and immediately adjacent areas; and the right dorsolateral prefrontal cortex, with each making a unique contribution to different aspects of performance. Furthermore, recent studies show striking dissociations between performances on multitasking tests and two of the most commonly administered measures of executive function: the verbal fluency test and the Wisconsin Card Sorting Test, which sets a minimum level for a fractionation of the executive syndrome in humans. Received: 31 March 1999 / Accepted: 23 July 1999     

Distinguishing intentions from desires: Contributions of the frontal and parietal lobes

The ability to represent desires and intentions as two distinct mental states was investigated in patients with parietal (N = 8) and frontal (N = 6) lesions and in age-matched controls (N = 7). A task was used where the satisfaction of the desire and the fulfilment of the intention did not co-vary and were manipulated in a 2 × 2 set. In two experiments we show that lesions to the frontal lobe may impair the ability to deal with desires when their outcome is not congruent with that of the intention, and that parietal damage – especially if it encompasses the left temporo-parietal junction – may cause severe difficulties in the processing of both desires and intentions. The implications of the results for the neuropsychological and the developmental literature are discussed.     

Neurolinguistic analysis of verbal utterances in patients with focal lesions of frontal lobes

Verbal utterances of 105 patients have been analyzed. Six groups of subjects were examined including patients with left dorsolateral frontal lesions, left orbitofrontal lesions, right frontal lobe lesions, posterior aphasics, posterior brain-damaged patients without aphasia, and a matched control group. Results indicate that the left frontal lobe is involved in organization of linguistic information. Its dorsolateral part seems to be concerned with forming the sequential pattern of an utterance, and the orbital part with the directed development of a narrative. It is conceivable that the right frontal lobe is important for the global, nonlinguistic organization of information to be uttered.     

Stare in the crowd: frontal face guides overt attention independently of its gaze direction

Whether or not a stare in the midst of many faces can guide visual attention is a controversial issue. Two experiments are reported that investigate the hypothesis that visual attention is guided toward a frontal face in the search for a stare among faces with varied head angles. The participants were required to search for a face with a direct gaze in a context where the target could be at any of various head angles and the target's head angle was unpredictable in one trial. The search performance was better for a frontal-face target than for deviated-face targets. Furthermore, eye-movement analyses revealed that a frontal-face stimulus tended to be initially fixated prior to deviated-face stimuli, and many of the initially fixated frontal-face stimuli had an averted gaze. The findings suggest that a frontal face guides overt attention independently of its gaze direction in the search for a stare in a crowd. The validity of prioritising a frontal face in order to find a direct gaze among faces and the characteristics of a human-face detection system are discussed.     

Executive control and the experience of regret

The experience of regret rests on a counterfactual analysis of events. Previous research indicates that regret emerges at around 6 years of age, marginally later than the age at which children begin to answer counterfactual questions correctly. We hypothesized that the late emergence of regret relative to early counterfactual thinking is a result of the executive demands of simultaneously holding in mind and comparing dual representations of reality (counterfactual and actual). To test this hypothesis, we administered two regret tasks along with four tests of executive function (two working memory tasks, a switch task, and an inhibition task) to a sample of 104 4- to 7-year-olds. Results indicated that switching, but not working memory or inhibition, was a significant predictor of whether or not children experienced regret. This finding corroborates and extends previous research showing that the development of counterfactual thinking in children is related to their developing executive competence.     

Human gaze control during real-world scene perception

In human vision, acuity and color sensitivity are best at the point of fixation, and the visual-cognitive system exploits this fact by actively controlling gaze to direct fixation towards important and informative scene regions in real time as needed. How gaze control operates over complex real-world scenes has recently become of central concern in several core cognitive science disciplines including cognitive psychology, visual neuroscience, and machine vision. This article reviews current approaches and empirical findings in human gaze control during real-world scene perception.