The Parieto-Frontal Integration Theory (P-FIT) of intelligence: converging neuroimaging evidence

"Is there a biology of intelligence which is characteristic of the normal human nervous system?" Here we review 37 modern neuroimaging studies in an attempt to address this question posed by Halstead (1947) as he and other icons of the last century endeavored to understand how brain and behavior are linked through the expression of intelligence and reason. Reviewing studies from functional (i.e., functional magnetic resonance imaging, positron emission tomography) and structural (i.e., magnetic resonance spectroscopy, diffusion tensor imaging, voxel-based morphometry) neuroimaging paradigms, we report a striking consensus suggesting that variations in a distributed network predict individual differences found on intelligence and reasoning tasks. We describe this network as the Parieto-Frontal Integration Theory (P-FIT). The P-FIT model includes, by Brodmann areas (BAs): the dorsolateral prefrontal cortex (BAs 6, 9, 10, 45, 46, 47), the inferior (BAs 39, 40) and superior (BA 7) parietal lobule, the anterior cingulate (BA 32), and regions within the temporal (BAs 21, 37) and occipital (BAs 18, 19) lobes. White matter regions (i.e., arcuate fasciculus) are also implicated. The P-FIT is examined in light of findings from human lesion studies, including missile wounds, frontal lobotomy/leukotomy, temporal lobectomy, and lesions resulting in damage to the language network (e.g., aphasia), as well as findings from imaging research identifying brain regions under significant genetic control. Overall, we conclude that modern neuroimaging techniques are beginning to articulate a biology of intelligence. We propose that the P-FIT provides a parsimonious account for many of the empirical observations, to date, which relate individual differences in intelligence test scores to variations in brain structure and function. Moreover, the model provides a framework for testing new hypotheses in future experimental designs.     

Gastric pathology and aphagia following lateral hypothalamic lesions in rats: effects of preoperative weight reduction.

The body weights of male albino rats were reduced gradually to 80% of normal body weight by resricting food intake (dieting), and then the rats were given lateral hypothalamic (LH) lesions. Compared with rats of normal body weight sustaining similar brain lesions, the dieted group displayed a shorter period of postoperative aphagia and less gastric pathology. In a second experiment, a group of rats was reduced to 80% of normal body weight by withholding all food (fasting) and then given LH lesions. Compared with dieted rats sustaining similar brain damage, the fasted group displayed a longer period of post operative aphagia and greater gastric pathology. Since the duration of aphagia could be shortened or lengthened by simple manipulations of preoperative body weight, the adequacy of sensorimotor or motivational hypotheses to account for aphagia is questioned. The results are more consistent with the suggestion that gastric abnormalities produced by LH lesions inhibit eating.     

Near-Death Experiences in patients with locked-in syndrome: Not always a blissful journey

Memories of Near-Death Experiences (NDEs) most often are recounted as emotionally positive events. At present, no satisfactory explanatory model exists to fully account for the rich phenomenology of NDEs following a severe acute brain injury. The particular population of patients with locked-in syndrome (LIS) provides a unique opportunity to study NDEs following infratentorial brain lesions. We here retrospectively characterized the content of NDEs in 8 patients with LIS caused by an acute brainstem lesion (i.e., “LIS NDEs”) and 23 NDE experiencers after coma with supratentorial lesions (i.e., “classical NDEs”). Compared to “classical NDEs”, “LIS NDEs” less frequently experienced a feeling of peacefulness or well-being. It could be hypothesized that NDEs containing less positive emotions might have a specific neuroanatomical substrate related to impaired pontine/paralimbic connectivity or alternatively might be related to the emotional distress caused by the presence of conscious awareness in a paralyzed body.     

A SAS/IML program using the Kalman filter for estimating state space models

To help disseminate the knowledge and software implementation of a state space model (SSM), this article provides a SAS/IML (SAS Institute, 2010) program for estimating the parameters of general linear Gaussian SSMs using the Kalman filter algorithm. In order to use this program, the user should have SAS installed on a computer and have a valid license for SAS/IML. Since the code is completely open, it is expected that this program can be used not only by applied researchers, but also by quantitative methodologists who are interested in improving their methods and promoting SSM as a research instrument.     

Online measures of basic language skills in children with early focal brain lesions

Twenty children with early focal lesions were compared with 150 age-matched control subjects on 11 online measures of the basic skills underlying language processing, a digit span task, and 6 standardized measures. Although most of the children with brain injury scored within the normal range on the majority of the tasks, they also had a disproportionately high number of outlier scores on the reaction time tests. This evidence for a moderate impairment of the basic skills underlying language processing contrasts with other evidence suggesting that these children acquire normal control of the functional use of language. Furthermore, these children scored within the normal range on a measure of general cognitive ability, suggesting that there is no particular sparing of linguistic functions at the expense of general cognitive functions. Using the MPD procedure (Valdés-Pérez & Pericliev, 1997), we found that the controls and the five clinical groups could be best distinguished with two measures of online processing (word repetition and visual number naming) and one standardized test subcomponent (the CELF Oral Directions subtest). The 12 children with left hemisphere lesions scored significantly lower than the 8 other children on the CELF-RS measure. Within the group of children with cerebral infarct, the nature of the processing disability could be linked fairly well to site of lesion. Otherwise, there was little relation between site or size of lesion and the pattern of deficit. These results support a model in which damage to the complex functional circuits subserving language leads to only minor deficits in process efficiency, because of the plasticity of developmental processes.     

Handling from weaning to adulthood does not prevent hyperdefensiveness induced by septal, medial accumbens, or medial hypothalamic lesions

Weanling male hooded rats were handled or not handled for 10 min each day, 5 days/week, for 6 weeks. At the end of this time all animals received one of the following: a septal lesion, a medial accumbens lesion, a medial hypothalamic lesion, or a sham lesion. The animals were tested for defensiveness toward the experimenter on Days 3, 7, and 14 postoperatively. All behavioral testing was done blind. Each of the three lesions increased defensiveness toward the experimenter. However, with each lesion, there was no difference in the defensiveness scores of preoperatively handled and nonhandled animals at any of the postoperative test sessions while those of animals with medial hypothalamic lesions did not. These results are consistent with observations that hyperdefensiveness occurs in human subjects following tumors in each of these brain areas in spite of their familiarity with the situation in which they are observed. They contrast with observations that mouse killing induced by septal lesions is readily prevented by preoperative exposure to a mouse.     

Early lexical development in children with focal brain injury

Early lexical development in 27 children with focal brain injury was studied cross-sectionally and longitudinally. Data were obtained from children between 12 and 35 months of age who acquired their lesion prenatally or within the first 6 months of life. Results for the group as a whole provide clear evidence for delays in lexical comprehension and production, and for a larger number of comprehension/production dissociations than would be expected by chance. In addition, a significant number of children were observed having unusual difficulty mastering predication and/or using an atypically high proportion of closed class words (suggesting reliance on holistic/formulaic speech). Analyses by lesion type revealed no effect of lesion size. Analyses according to side of lesion revealed that children with right-hemisphere damage produced a higher proportion of closed class words, suggesting heavy reliance on well-practiced but under-analyzed speech formulae. Children with left-hemisphere damage were slightly better in comprehension than children with right-hemisphere damage. In addition, left posterior lesions were associated with greater delays in expressive language, and delays were more protracted in children with left posterior damage. No differential effects of left posterior damage were found for lexical comprehension.     

Mapping cognition to the brain through neural interactions

Brain imaging methods, such as positron emission tomography (PET) and functional magnetic resonance imaging (fMRI), provide a unique opportunity to study the neurobiology of human memory. As these methods can measure most of the brain, it is possible to examine the operations of large-scale neural systems and their relation to cognition. Two neuroimaging studies, one concerning working memory and the other episodic memory retrieval, serve as examples of application of two analytic methods that are optimised for the quantification of neural systems, structural equation modelling, and partial least squares. Structural equation modelling was used to explore shifting prefrontal and limbic interactions from the right to the left hemisphere in a delayed match-to-sample task for faces. A feature of the functional network for short delays was strong right hemisphere interactions between hippocampus, inferior prefrontal, and anterior cingulate cortices. At longer delays, these same three areas were strongly linked, but in the left hemisphere, which was interpreted as reflecting change in task strategy from perceptual to elaborate encoding with increasing delay. The primary manipulation in the memory retrieval study was different levels of retrieval success. The partial least squares method was used to determine whether the image-wide pattern of covariances of Brodmann areas 10 and 45/47 in right prefrontal cortex (RPFC) and the left hippocampus (LGH) could be mapped on to retrieval levels. Area 10 and LGH showed an opposite pattern of functional connectivity with a large expanse of bilateral limbic cortices that was equivalent for all levels of retrieval as well as the baseline task. However, only during high retrieval was area 45/47 included in this pattern. The results suggest that activity in portions of the RPFC can reflect either memory retrieval mode or retrieval success depending on other brain regions to which it is functionally linked, and imply that regional activity must be evaluated within the neural context in which it occurs. The general hypothesis that learning and memory are emergent properties of large-scale neural network interactions is discussed, emphasising that a region can play a different role across many functions and that role is governed by its interactions with anatomically related regions.     

The opercular-subopercular syndrome: four cases with review of the literature

We present four cases of the 'opercular syndrome' of volitional paresis of the facial, lingual, and laryngeal muscles (bilateral facio-glosso-pharyngo-masticatory paresis). Case histories and CT brain images are presented, along with a review of the literature concerning this long-recognized but little-known syndrome. The neuroanatomic basis of the syndrome classically involves bilateral lesions of the frontal operculum. We propose, on the basis of our cases and others, that the identical syndrome can arise from lesions of the corticobulbar tracts, not involving the cortical operculum. Our cases included one with bilateral subcortical lesions, one with a unilateral left opercular lesion and a possible, non-visualized right hemisphere lesion, one with unilateral cortical and unilateral subcortical pathology, and one with bilateral cortical lesions. These lesion localizations suggest that any combination of cortical or subcortical lesions of the operculum or its connections on both sides of the brain can produce a syndrome indistinguishable from the classical opercular syndrome. We propose the new term 'opercular-subopercular syndrome' to encompass cases with predominantly or partially subcortical lesions.     

Cerebellar lesions abolish an avoidance response in rabbit

Unilateral lesions of the cerebellar dentate-interpositus (DI) nuclei abolished or impaired the reacquisition of a nictitating membrane (NM) conditioned avoidance response (CAR) in rabbits. Animals that sustained damage to the DI nuclei and surrounding areas displayed little or no reacquisition of the NM-CAR ipsilateral to the lesion, but displayed excellent acquisition of the response when training was shifted to the contralateral eye. These results agree with previous investigations of the effects of cerebellar lesions on the classically conditioned NM and leg-flexion responses. The DI nuclei appear to be essential for the retention of conditioned responses in a variety of associative paradigms.     

Word-category violations in patients with Broca's aphasia: an ERP study

An event-related brain potential experiment was carried out to investigate on-line syntactic processing in patients with Broca's aphasia. Subjects were visually presented with sentences that were either syntactically correct or contained violations of word-category. Three groups of subjects were tested: Broca patients (N = 11), non-aphasic patients with a right hemisphere (RH) lesion (N = 9), and healthy aged-matched controls (N = 15). Both control groups appeared sensitive to the violations of word-category as shown by clear P600/SPS effects. The Broca patients displayed only a very reduced and delayed P600/SPS effect. The results are discussed in the context of a lexicalist parsing model. It is concluded that Broca patients are hindered to detect on-line violations of word-category, if word class information is incomplete or delayed available.     

Prevalence,and intellectual outcome of unilateral focal cortical brain damage as a function of age,sex and aetiology

Neurologists and neuropsychologists are aware that aging men are more at risk than women for brain damage, principally because of the well known male-predominant risk for cardiovascular disease and related cerebrovascular accidents. However, a disproportion in prevalence of brain damage between the sexes in childhood may be less suspected. Furthermore, sex-specific risk for other aetiologies of brain damage may be little known, whether in the pediatric or adult populations. Proposals of a sex difference in cognitive recovery from brain damage have also been controversial. Six hundred and thirty five "consecutive" cases with cortical focal lesions including cases of all ages and both sexes were reviewed. Aetiology of the lesion was determined for each case as was postlesion IQ. Risk was highly male prevalent in all age groups, with a predominance of cardiovascular aetiology explaining much of the adult male prevalence. However, several other aetiological categories were significantly male prevalent in juveniles (mitotic, traumatic, dysplasic) and adults (mitotic, traumatic). There was no sex difference in outcome (i.e., postlesion IQ) of these cortical brain lesions for the cohort as a whole, after statistical removal of the influence of lesion extent, aetiology and presence of epilepsy. Mechanisms potentially responsible for sex differences in prevalence, aetiology of brain damage, and recovery, are reviewed and discussed.     

PsychoPro 2.0: Using multidimensional scaling to examine the perceptual categorization of race

Recently, researchers examining cognitive mechanisms involved in the detection of racial markers have been using psychophysics to examine perceptual thresholds delineating race using a program called PsychoPro (MacLin, MacLin, Peterson, Chowdhry, & Joshi, 2009). PsychoPro allows researchers to collect data using facial stimuli morphed along a racial continuum with psychophysical methodology. PsychoPro 2.0 was updated so researchers could collect paired-comparison data that can then be analyzed using multidimensional scaling (MDS). MDS allows researchers to examine the perceptions of racial boundaries along perceptual dimensions not bounded by the stimulus continuum, by having the observer make paired comparisons to racial stimuli based on their perceived similarity. Results indicate that ratings were made on the basis of at least six different racial dimensions that suggest the emergence of a third race not represented by the original morph starting images. Details of the software, the new paired-comparison feature and its relevance in psychophysical studies in general and racial categorization in particular are presented.     

Mapping Cognition to the Brain Through Neural Interactions

Brain imaging methods, such as positron emission tomography (PET) and functional magnetic resonance imaging (fMRI), provide a unique opportunity to study the neurobiology of human memory. As these methods can measure most of the brain, it is possible to examine the operations of large-scale neural systems and their relation to cognition. Two neuroimaging studies, one concerning working memory and the other episodic memory retrieval, serve as examples of application of two analytic methods that are optimised for the quantification of neural systems, structural equation modelling, and partial least squares. Structural equation modelling was used to explore shifting prefrontal and limbic interactions from the right to the left hemisphere in a delayed match-to-sample task for faces. A feature of the functional network for short delays was strong right hemisphere interactions between hippocampus, inferior prefrontal, and anterior cingulate cortices. At longer delays, these same three areas were strongly linked, but in the left hemisphere, which was interpreted as reflecting change in task strategy from perceptual to elaborate encoding with increasing delay. The primary manipulation in the memory retrieval study was different levels of retrieval success. The partial least squares method was used to determine whether the image-wide pattern of covariances of Brodmann areas 10 and 45/47 in right prefrontal cortex (RPFC) and the left hippocampus (LGH) could be mapped on to retrieval levels. Area 10 and LGH showed an opposite pattern of functional connectivity with a large expanse of bilateral limbic cortices that was equivalent for all levels of retrieval as well as the baseline task. However, only during high retrieval was area 45/47 included in this pattern. The results suggest that activity in portions of the RPFC can reflect either memory retrieval mode or retrieval success depending on other brain regions to which it is functionally linked, and imply that regional activity must be evaluated within the neural context in which it occurs. The general hypothesis that learning and memory are emergent properties of large-scale neural network interactions is discussed, emphasising that a region can play a different role across many functions and that role is governed by its interactions with anatomically related regions.     

Using a computer simulation of three slot machines to investigate a gambler’s preference among varying densities of near-miss alternatives

The present article describes a software program in Visual Basic .NET designed to simulate three slot machines on a computer screen. This software is described in detail regarding utility, downloading, and usage; and data are presented illustrating the software’s potential for researchers interested in gambling behavior. A simulation of multiple slot machines such as this enables researchers to evaluate players’ preferences across various machines. In the highlighted experiment, 18 recreational slot machine players played the software for extra course credit and a chance at cash prizes. All participants played a version of the simulation in which every 5th response on average was a win, whereas the remaining trials were a loss. However, on those loss trials, a varying distribution of almost wins or near misses (i.e., two winning symbols on the payoff line and the final winning symbol directly above or below the payoff line) were presented in percentages of 15, 30, or 45. While no preferences across the three options could be predicted on the basis of reinforcement history alone, deviations from equal choices across the games were noted and appeared to be the result of the presentations of near-miss losing trials. Implications for a greater understanding of pathological gambling are presented.     

The Occipital Cortex in the Blind

ABSTRACT— Studying the brains of blind individuals provides a unique opportunity to investigate how the brain changes and adapts in response to afferent (input) and efferent (output) demands. We discuss evidence suggesting that regions of the brain normally associated with the processing of visual information undergo remarkable dynamic change in response to blindness. These neuroplastic changes implicate not only processing carried out by the remaining senses but also higher cognitive functions such as language and memory. A strong emphasis is placed on evidence obtained from advanced neuroimaging techniques that allow researchers to identify areas of human brain activity, as well as from lesion approaches (both reversible and irreversible) to address the functional relevance and role of these activated areas. A possible mechanism and conceptual framework for these physiological and behavioral changes is proposed.     

Neocortex and feeding behavior in the rat.

Consummatory behavior and weight-regulation capacity were measured in 12 normal rats and in 43 rats that survived complete (C), sequential unilateral (U), anterolateral (A), or posterior (P) neocortical ablations. Groups C and A displayed aphagia and adipsia followed by a sequence of recovery stages gualitatively identical to, but shorter than, recovery typically seen following lateral hypothalamic lesions. After recovery, Group C displayed long-term effects of finickiness and pradial drinking. These effects as well as a measure of recovery of body-weight-regulation capacity were significantly intercorrelated with lesion size, and body-weight set point remained significantly lower than normal. Group U was relatively unaffected by the first unilateral ablation and showed, relative the second ablation but displayed the long-term effects. Group P, though significantly affected by the lesion, did not display the pattern or intensity of effects described for the other bilaterally ablated groups.     

Prevalence and situational causes of stereotyped behaviors in blind infants and preschoolers

Parents of 85 blind children aged from 10 months to the 6th year of life were asked regarding the frequency, duration and typical situations of the occurrence of various stereotypic behavors in their children. The Bielefeld Parents' Questionnaire for Blind and Sighted Infants and Preschoolers was used as the instrument of measurement. All of the children displayed at least one stereotypic behavior; most displayed several stereotypic behaviors according to the parents' reports. Eye poking and body rocking dominated within the prevalence hierarchy. Four typical situations could be identified in which stereotypic behaviors were shown: monotony, arousal, demand, and during feeding or eating. The results suggested that repetitive hand and finger movements, stereotypic manipulation of objects, and making a face(s) mainly occur within arousal situations whereas eye poking, whimpering, and sucking thumbs or fingers especially are linked to monotony.This research was carried out within a research program on early intervention and family care for blind infants and preschoolers. This program is part of the Special Research Unit on Prevention and Intervention in Childhood and Adolescence (Sonderforschungsbereich No. 227) at the University of Bielefeld funded by the German Research Association (DFG). The authors would like to thank Gail Weingart and Jonathan Harrow (University of Bielefeld) for translating this paper from German.