Neuroanatomical substrates of age-related cognitive decline

There are many reports of relations between age and cognitive variables and of relations between age and variables representing different aspects of brain structure and a few reports of relations between brain structure variables and cognitive variables. These findings have sometimes led to inferences that the age-related brain changes cause the age-related cognitive changes. Although this conclusion may well be true, it is widely recognized that simple correlations are not sufficient to warrant causal conclusions, and other types of correlational information, such as mediation and correlations between longitudinal brain changes and longitudinal cognitive changes, also have limitations with respect to causal inferences. These issues are discussed, and the existing results on relations of regional volume, white matter hyperintensities, and diffusion tensor imaging measures of white matter integrity to age and to measures of cognitive functioning are reviewed. It is concluded that at the current time the evidence that these aspects of brain structure are neuroanatomical substrates of age-related cognitive decline is weak. The final section contains several suggestions concerning measurement and methodology that may lead to stronger conclusions in the future.     

Musicians processing music: Measurement of brain potentials with EEG

Abstract

In the present study, the EEG was recorded from the scalp of musicians while mentally active in their field. Analytic, creative and memory processes of the brain were observable using a special electrophysiological method called DC-potential recording. Music students listened to a sequence of four notes and subsequently were either to reverse the sequence (task 1 = analytic) or to compose a new continuation (task 2 = creative). In task 3, the initial segment of a well-known melody was presented and had to be continued (memory task). All tasks had to be solved mentally (imagery). In tasks 1 and 2, either tonal or atonal sequences were presented.

While processing, the results show that the analytic task elicited the highest brain activity. The analytic task involved mainly parieto-temporal areas of both hemispheres, the left hemisphere showing a tendency for domination. The memory task produced predominant activity over the right hemisphere. The creative task caused the lowest brain activation and elicited an unexpected lateralisation to the left, though we expected creativity to be a right hemispheric holistic-synthetic phenomenon.

Comparing listening with processing of the perceived music, we found a significant shift from an insignificant right hemispheric to an insignificant left hemispheric predominance (except with the memory task). This indicates that musicians do not lateralise to the left hemisphere per se when listening to music. Whether one finds a left hemispheric lateralisation in listening tasks or a right hemispheric one probably depends on the amount of simultaneous analytic-sequential processing the musician undertakes.     

Modelling age-related changes in information processing

Researchers in cognitive ageing seldom take advantage of explicit quantitative models of information processing to account for age differences in cognition. Where quantitative models have been used, these models typically remain silent about the details of information processing. The lack of explicit cognitive models has consequences for the interpretation of a number of empirical results. Using a specific class of models called random walk models, I review evidence showing that the empirical relations taken as support for global age-related changes are consistent with a number of possible age effects on information processing. In addition, I demonstrate that these models can be used to account for age differences within the context of individual experiments and such modelling has important implications for the interpretation of age differences in performance.     

Age-related changes in auditory temporal processing

Two tone bursts separated by a silent interval and imbedded in a background white noise were presented to elderly subjects (M age = 71.3 years) and young subjects (M age = 22.2 years). Subjects were required to judge when the two tone bursts fused perceptually by adjusting the duration of tone-one. The bursts were separated by six discrete interstimulus intervals of 4, 8, 16, 24, 32, or 40 ms. The tone-two burst was held constant at 100 ms. Fusion point was defined as that critical tone-one duration at which the two tones fused (were perceived as one). Elderly subjects reached fusion point at longer critical tone-one durations than young subjects at each interval tested. The function relating tone-one duration and interval conformed to an exponential curve and is discussed with respect to an exponential decay model of the inhibitory interactions of neural systems responding to onsets and offsets of sensory events.     

Language and auditory processing in autism

Autism is characterized by varying degrees of disorders in language, communication and imagination. What are the prospects for making sense of this heterogeneous condition? Advances in identifying phenotypes in relation to subgroups within autism, based on disproportionate language impairment, have been recently reported by Tager-Flusberg and Joseph. The symptom severity of these subgroups requires investigation for underlying deficits, such as in auditory processing. Other recent reports support the view that a deficit in auditory processing might be a key factor in autism.     

Schema-based processing in auditory scene analysis

What is the involvement of what we know in what we perceive? In this article, the contribution of melodic schema-based processes to the perceptual organization of tone sequences is examined. Two unfamiliar six-tone melodies, one of which was interleaved with distractor tones, were presented successively to listeners who were required to decide whether the melodies were identical or different. In one condition, the comparison melody was presented after the mixed sequence: a target melody interleaved with distractor tones. In another condition, it was presented beforehand, so that the listeners had precise knowledge about the melody to be extracted from the mixture. In the latter condition, recognition performance was better and a bias toward same responses was reduced, as compared with the former condition. A third condition, in which the comparison melody presented beforehand was transposed up in frequency, revealed that whereas the performance improvement was explained in part by absolute pitch or frequency priming, relative pitch representation (interval and/or contour structure) may also have played a role. Differences in performance as a function of mean frequency separation between target and distractor sequences, when listeners did or did not have prior knowledge about the target melody, argue for a functional distinction between primitive and schema-based processes in auditory scene analysis.     

On the relative role of inhibition in age-related working memory decline

In this cross-sectional study, the mediating role of inhibition relative to speed in age-related working memory decline on different tasks was investigated. It was hypothesized that the role of inhibition is marginal or absent in a relatively "passive" or simple working memory task, but present in a relatively "active" or complex task in which the processing, rather than the storage capacity of working memory, is addressed. This hypothesis was tested with a structural equation model that was fitted on a subsample (N = 213) of the Maastricht Aging Study (MAAS). The first trial of the Verbal Learning Test (VLT) served as the passive test and the Self-Paced Auditory Serial Addition Task (SPASAT) as the active test. Results showed that the role of inhibition was absent in predicting VLT performance, whereas it was comparable to the role of speed in predicting SPASAT performance. These findings suggest a relative, task-dependent role of inhibition in explaining age-related memory decline.     

Pre-attentive auditory processing of lexicality

The effects of lexicality on auditory change detection based on auditory sensory memory representations were investigated by presenting oddball sequences of repeatedly presented stimuli, while participants ignored the auditory stimuli. In a cross-linguistic study of Hungarian and German participants, stimulus sequences were composed of words that were language-familiar, lexical, meaningful in Hungarian but language-unfamiliar, not lexical, meaningless in German, and words with the opposite characteristics. The roles of frequently presented stimuli (Standards) and infrequently presented one (Deviants) were fully crossed. Language-familiar and language-unfamiliar Deviants elicited the Mismatch Negativity component of the event-related brain potential. We found differences in processes of change detection depending on whether the Standard was language-familiar, or not. Whereas, the lexicality of the Deviant had no effect on the processes of change detection. Also, language-familiar Standards processed differently than language-unfamiliar ones. We suggest that pre-attentive (default) tuning to meaningful words sets up language-specific preparatory processes that affect change detection in speech sequences.     

Stimulus-dependent flexibility in non-human auditory pitch processing

Songbirds and humans share many parallels in vocal learning and auditory sequence processing. However, the two groups differ notably in their abilities to recognize acoustic sequences shifted in absolute pitch (pitch height). Whereas humans maintain accurate recognition of words or melodies over large pitch height changes, songbirds are comparatively much poorer at recognizing pitch-shifted tone sequences. This apparent disparity may reflect fundamental differences in the neural mechanisms underlying the representation of sound in songbirds. Alternatively, because non-human studies have used sine-tone stimuli almost exclusively, tolerance to pitch height changes in the context of natural signals may be underestimated. Here, we show that European starlings, a species of songbird, can maintain accurate recognition of the songs of other starlings when the pitch of those songs is shifted by as much as ±40%. We observed accurate recognition even for songs pitch-shifted well outside the range of frequencies used during training, and even though much smaller pitch shifts in conspecific songs are easily detected. With similar training using human piano melodies, recognition of the pitch-shifted melodies is very limited. These results demonstrate that non-human pitch processing is more flexible than previously thought and that the flexibility in pitch processing strategy is stimulus dependent.     

The specificity of age-related decline in interpretation of emotion cues from prosody

Older adults are not as good as younger adults at decoding prosodic emotions. We sought to determine the specificity of this finding. Performance of older and younger adults was compared on a prosodic emotion task, a "pure" prosodic emotion task, a linguistic prosody task, and a "pure" linguistic prosody task. Older adults were less accurate at interpreting prosodic emotion cues and nonemotional contours, concurrent semantic processing worsened interpretation, and performance was further degraded when identifying negative emotions and questions. Older adults display a pervasive problem interpreting prosodic cues, but further study is required to clarify the stage at which performance declines.     

A demonstration of auditory top-down processing

A simple demonstration of auditory top-down processing is described in which one speech embedded in several others becomes much clearer when participants read a text of the target speech at the same time as they hear it.     

Simultaneous auditory and tactile information processing

Subjects were required to perform perceptual tasks when stimuli were presented simultaneously in the auditory and tactile modalities and when they were presented in one of the modalities alone. The results indicated that when the demands on cognitive processes are small, auditory and tactile stimuli presented simultaneously can be processed as well as when stimuli are presented in only one modality. In a task which required a large amount of cognitive processing, it became difficult for subjects to maintain high levels of performance in both modalities and the distribution of attention became an important determinant of performance. The data were consistent with a theory that cognitive, but not perceptual, processing is disrupted when subjects have difficulty performing two perceptual tasks simultaneously.     

Environmentally induced analgesia: age-related decline in a neurally mediated, nonopioid system

In order to examine the function of an endogenous system of pain inhibition during aging, rats (3, 14, and 24 months old) were exposed to 90 s of hind-paw shock. To investigate the pharmacology and anatomy involved in the production of hind-paw shock induced analgesia, the effects of naltrexone (7 mg/kg), scopolamine (5 mg/kg), and adrenalectomy were examined. Results revealed that there was an age-related reduction in the degree of analgesia produced by hind-paw shock. Naltrexone and adrenalectomy did not alter the analgesia elicited by hind-paw shock. Scopolamine reduced the analgesia produced by hind-paw shock, and the effectiveness of scopolamine blockage declined with age. The results of a second experiment demonstrated that the effect of scopolamine was specific to the analgesia induced by hind-paw shock because scopolamine was ineffective in modifying the analgesia produced by a different stressor (cold water). These results suggest that the decline in hind-paw shock induced analgesia is the result of an alteration in the function of the cholinergic system.     

Task-dependent costs in processing two simultaneous auditory stimuli

A listener presented with two speech signals must at times sacrifice the processing of one signal in order to understand the other. This study was designed to distinguish costs related to interference from a second signal (selective attention) from costs related to performing two tasks simultaneously (divided attention). Listeners presented with two processed speech-in-noise stimuli, one to each ear, either (1) identified keywords in both or (2) identified keywords in one and detected the presence of speech in the other. Listeners either knew which ear to report in advance (single task) or were cued afterward (partial-report dual task). When the dual task required two identification judgments, performance suffered relative to the single-task condition (as measured by percent correct judgments). Two different tasks (identification for one stimulus and detection for the other) resulted in much smaller reductions in performance when the cue came afterward. We concluded that the degree to which listeners can simultaneously process dichotic speech stimuli seems to depend not only on the amount of interference between the two stimuli, but also on whether there is competition for limited processing resources. We suggest several specific hypotheses as to the structural mechanisms that could constitute these limited resources.     

Executive functioning as a potential mediator of age-related cognitive decline in normal adults

Critical requirements for the hypothesis that executive functioning is a potential mediator of age-related effects on cognitive functioning are that variables assumed to reflect executive functioning represent a distinct construct and that age-related effects on other types of cognitive functioning are reduced when measures of executive functioning are statistically controlled. These issues were investigated in a study involving 261 adults between 18 and 84 years of age. Although age-related effects on various cognitive abilities were substantially reduced after statistical control of the variance in measures hypothesized to represent executive functioning, there was only weak evidence for the existence of distinct constructs corresponding to executive functioning or to aspects of executive control concerned with inhibition, updating, or time sharing.     

Feature and conjunction processing in the auditory modality

In five experiments, participants made speeded target/nontarget classification responses to singly presented auditory stimuli. Stimuli were defined via vocal identity and location in Experiments 1 and 2 and frequency and location in the remaining experiments. Performance was examined in two conditions inspired by visual search: In the feature condition, responses were based on the detection of unique stimulus features; in the conjunction condition, unique combinations of features were critical. Experiment 1 showed a conjunction benefit, since classifications were faster in the conjunction condition than in the feature condition. Potential confounds were eliminated in Experiments 2 and 3, which resulted in the observation of conjunction costs. In Experiments 4 and 5, we examined, respectively, whether the cost could be explained in terms of differences in interstimulus similarity and target template complexity across the main conditions. Both accounts were refuted. It seems that when the identification of particular feature combinations is necessary, conjunction processing in audition becomes an effortful process.     

Within- and between-dimensional processing in the auditory modality

Participants made speeded target-nontarget responses to singly presented auditory stimuli in 2 tasks. In within-dimension conditions, participants listened for either of 2 target features taken from the same dimension; in between-dimensions conditions, the target features were taken from different dimensions. Judgments were based on the presence or absence of either target feature. Speech sounds, defined relative to sound identity and locale, were used in Experiment 1, whereas tones, comprising pitch and locale components, were used in Experiments 2 and 3. In all cases, participants performed better when the target features were taken from the same dimension than when they were taken from different dimensions. Data suggest that the auditory and visual systems exhibit the same higher level processing constraints.