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Biomedical subjects

D Fitzpatrick

Publications and source records attributed to D Fitzpatrick.

At least 37 records · Page 2Linked to original sources

Cognitive evoked potentials to anticipated oesophageal stimulus in humans: quantitative assessment of the cognitive aspects of visceral perception.

Evoked potential studies provide an objective measure of the neural pathways involved with perception. The effects of cognitive factors, such as anticipation or awareness, on evoked potentials are not known. The aim was to compare the evoked potential response to oesophageal stimulation with the cortical activity associated with anticipation of the same stimulus. In 12 healthy men (23.5 +/- 4 years), oesophageal electrical stimulation (15 mA, 0.2 Hz, 0.2 msec) was applied, and the evoked potentials recorded using scalp electrodes. A computerized model of randomly skipped stimuli (4:1 ratio) was used to separately record the evoked potentials associated with stimulation and those associated with an anticipated stimulus. The electrical stimulus represented the nontarget stimulus and the skipped impulse the target (anticipatory) stimulus. This anticipatory evoked potential was also compared to auditory P300 evoked potentials. Reproducible evoked potentials and auditory P300 responses were elicited in all subjects. Anticipatory evoked potentials (peak latency 282.1 +/- 7.9 msec, amplitude 8.2 +/- 0.7 microV, P < 0.05 vs auditory P300 evoked potential) were obtained with the skipped stimulus. This anticipatory evoked potential was located frontocentrally, while the auditory P300 potential was located in the centro-parietal cortex. The anticipatory evoked potential associated with expectation of an oesophageal stimulus, although of similar latency to that of the auditory P300 evoked response, originates from a different cortical location. The recording of cognitive evoked potentials to an expected oesophageal stimulus depends on attention to, and awareness of, the actual stimulus. Anticipatory evoked potentials to GI stimuli may provide an objective electrophysiological tool for the assessment of the cognitive factors associated with visceral perception.

Adult↗

Apoptosis in vascular endothelial cells caused by serum deprivation, oxidative stress and transforming growth factor-beta.

Vascular endothelial cell apoptosis has previously been shown to play a role in the pathogenesis of hypertension-induced vessel deletion and damage. In the present in vitro study we analyse several possible relevant causative factors of vascular endothelial cell apoptosis, namely, serum deprivation and nutrient depletion, oxidative stress in the forms of hypoxia, hyperoxia or free radical damage, and altered levels of transforming growth factor-beta1 (TGF-beta1) protein. An established cell line, bovine aortic endothelial cells (BAEC), was maintained in complete growth medium (RPMI-1640 plus 15% fetal calf serum and antibiotics, abbreviated as RPMI) in 25cm2 flasks or in 12-well plates on glass coverslips. Confluent but actively-growing cultures were treated with either hypoxia (PO2 of RPMI = 50mmHg), serum-free media (SFM), SFM plus hypoxia, hyperoxia (PO2 of RPMI = 450mmHg), hydrogen peroxide (H2O2, 1mM) in SFM, or TGF-beta1 protein (10ng/mL) in SFM. Appropriate control cultures were used. BAEC were collected 48h or 72h after all treatments except for TGF-beta1 and H2O2 treatments that were collected at 16-18h. Cell death was assessed using morphological characteristics or in situ end labeling (ISEL), cell proliferation assessed using proliferating cell nuclear antigen (PCNA), and TGF-beta1 expression assessed using transcript levels or immunohistochemistry. All treatments significantly increased levels of apoptosis over control cultures (P<0.05), and decreased levels of cell proliferation. Treatment with TGF-beta1 protein or SFM plus hypoxia induced greatest levels of apoptosis. TGF-beta1 protein and transcript levels were decreased in treated cultures, results suggesting that a paracrine source of TGF-beta1 protein would be needed as a cause of endothelial cell apoptosis in viva. Future therapies against inappropriate vessel deletion in disease states may use the known gene-driven nature of apoptosis to modify this sort of cell death in endothelial cells.

Animals↗

Unequal representation of cardinal and oblique contours in ferret visual cortex.

We have measured the amount of cortical space activated by differently oriented gratings in 25 adult ferrets by optical imaging of intrinsic signal. On average, 7% more area of the exposed visual cortex was preferentially activated by vertical and horizontal contours than by contours at oblique angles. This anisotropy may reflect the real-world prevalence of contours in the cardinal axes and could explain the greater sensitivity of many animals to vertical and horizontal stimuli.

Animals↗

Cortical evoked responses following esophageal balloon distension and electrical stimulation in healthy volunteers.

Recording of evoked potential responses represents an objective and quantifiable method to study visceral afferent sensory pathways in humans. We examined the evoked responses to mechanical distension (balloon) and electrical stimulation of the proximal and distal esophagus. A standard manometric catheter with a latex balloon and an additional electrode attached to its body was placed in the lower esophagus in 15 healthy young volunteers. Repeated nonpainful balloon distension stimuli above the individual sensation threshold (0.17 Hz, 12-20 ml) or short electrical impulses (0.2 Hz, 12-16 mA) were delivered in an alternate fashion at 23 and 33 cm from the nares. Evoked potential responses (EP) were recorded through 22 scalp surface electrodes using the standard 10/20 International EEG system of electrode placement. Balloon distension produced a reproducible triphasic response at both sites. Peak latencies of three negative EP peaks were 92+/-17, 229+/-40, and 339+/-36 msec with proximal stimulation versus 154+/-24, 275+/-24, and 384+/-30 msec obtained with distal stimulation (P < 0.001). Electrical stimulation produced a triphasic response with significantly shorter peak latencies at both sites when compared to mechanical stimulation (P < 0.001). Peak latencies were 74+/-12, 137+/-11, and 245+/-27 msec proximal versus 83+/-12, 148+/-32, and 247+/-51 msec with distal stimulation (P < 0.01). The calculated conduction velocities for both modes of stimulation (balloon: 1.73+/-0.9 m/sec vs electrical: 10.1+/-3.4 m/sec) are compatible with conduction through C fibers and Adelta fibers, respectively. Both modes of stimulation produce characteristic brain responses that are conveyed through different types of afferent fibers. The respective contributions of both types of fibers to esophageal function and symptomatology can be specifically addressed using this approach in both normal and pathologic conditions.

Adult↗

Fear of seizures: an investigation and treatment.

Despite the wealth of epidemiological evidence showing the comorbidity of epilepsy with various forms of psychopathology, there has been little systematic investigation of its relationship with affective disorders. This study aims to explore the phenomenon of seizure fear. Survey data have suggested that patients commonly fear death and/or brain damage as a result of their seizures, but documented cases of seizure phobia are rare and infrequently address issues of treatment. The study describes the exploration and successful cognitive behavioural treatment of a case of seizure phobia in a 26 year old woman with a 9 year history of epilepsy following a subarachnoid haemorrhage. The findings represent an important development in the much-neglected study of seizure fear. The case highlights some of the factors which may contribute to the genesis and maintenance of epilepsy-related psychopathologies and consequently has important implications for theoretical modelling in this area. Furthermore, it provides encouraging evidence to support the use of cognitive-behavioural techniques in treatment of these disorders.

Adaptation, Psychological↗

Assessment of the visceral afferent and autonomic pathways in response to esophageal stimulation in control subjects and in patients with diabetes.

OBJECTIVE: To examine the effects of esophageal stimulation on vagal afferent and efferent pathways in volunteers without diabetes and patients with diabetes. DESIGN: Prospective physiological study. PARTICIPANTS: Fourteen control subjects without diabetes and 6 patients with diabetes. INTERVENTIONS: Electrical and mechanical stimulation of the esophagus. OUTCOME MEASURES: Cortical evoked potentials and the power spectra of heart rate variability. RESULTS: For the control subjects, there was a significant decrease in the ratio of the low frequency to high frequency (LF:HF) power (i.e., increased vagal efferent modulation) during stimulation. Reproducible cortical evoked potentials were obtained from all control subjects. In the 6 patients with diabetes, who had viscerosensory and autonomic neuropathy, the cortical evoked potentials showed an erratic non-reproducible response to electrical esophageal stimulation; however, the LF:HF ratio decreased in these patients during stimulation, suggesting an intact subcortical reflex circuit. CONCLUSIONS: Vago-afferent fibres can be studied using minimally invasive techniques, and the power spectral analysis of heart rate variability permits study of autonomic vago-efferent pathways.

Adolescent↗

Orientation selectivity and the arrangement of horizontal connections in tree shrew striate cortex.

Horizontal connections, formed primarily by the axon collaterals of pyramidal neurons in layer 2/3 of visual cortex, extend for millimeters parallel to the cortical surface and form patchy terminations. Previous studies have provided evidence that the patches formed by horizontal connections exhibit modular specificity, preferentially linking columns of neurons with similar response characteristics, such as preferred orientation. The issue of how these connections are distributed with respect to the topographic map of visual space, however, has not been resolved. Here we combine optical imaging of intrinsic signals with small extracellular injections of biocytin to assess quantitatively the specificity of horizontal connections with respect to both the map of orientation preference and the map of visual space in tree shrew V1. Our results indicate that horizontal connections outside a radius of 500 microm from the injection site exhibit not only modular specificity, but also specificity for axis of projection. Labeled axons extend for longer distances, and give off more terminal boutons, along an axis in the map of visual space that corresponds to the preferred orientation of the injection site. Inside of 500 microm, the pattern of connections is much less specific, with boutons found along every axis, contacting sites with a wide range of preferred orientations. The system of long-range horizontal connections can be summarized as preferentially linking neurons with co-oriented, co-axially aligned receptive fields. These observations suggest specific ways that horizontal circuits contribute to the response properties of layer 2/3 neurons and to mechanisms of visual perception.

Animals↗

Estimation of habituation and signal-to-noise ratio of cortical evoked potentials to oesophageal electrical and mechanical stimulation.

Electrical and mechanical stimulation of the oesophagus has been recently proposed to examine the physiological effects of autonomic stimulation in humans. Cortical evoked potentials (EPs) to oesophageal stimulation provide an assessment of afferent fibres and central processing. However, habituation takes place during averaging of cortical EPs and reduces the signal-to-noise ratio (SNR) as the number of stimuli increases. The SNR of cortical EPs to oesophageal stimulation is computed for 15 normal subjects. Habituation is characterised by the Euclidean distance between the EEG response to single stimuli and the averaged EP, to serve as an objective measure of similarity between the averaged EP and the single-stimulus EEG. With electrical stimulation, the SNR is highest (0.41 +/- 0.21) for 1-12 stimuli and then significantly decreases to 0.2 +/- 0.08 for 13-24 stimuli (p < 0.001). With balloon distension (BD), the SNR is highest (0.22 +/- 0.16) for 1-12 stimuli and lowest (0.12 +/- 0.14) for 13-24 stimuli, but these SNRs are not significantly different from each other. Both electrical and mechanical stimulation of the oesophagus produce rapidly adapting EPs. The SNR of the EPs is higher with electrical stimulation than with BD. The EPs response to BD has a higher variability and is more noisy. Consequently, these results suggest that the overall cortical EP response to electrical stimulation of the oesophagus is more reproducible than that due to balloon distension.

Adult↗

The magnitude of the central response to esophageal electrical stimulation is intensity dependent.

BACKGROUND & AIMS: Cerebral evoked potential (EP) responses to visceral stimulation represent a powerful method to assess visceral afferent pathways. The aim of this study was to establish basic stimulation parameters (dose-response relationship in EP amplitude and topographic brain organization) during electrical esophageal stimulation. METHODS: Electrical esophageal stimulation was performed in repeated series of 24 stimuli in 15 healthy subjects (25 years) by steps of 5 mA, ranging from 0.5 mA (sham) to 25 mA. EPs were obtained using scalp electrodes positioned according to the 10/20 International electroencephalographic system. Topographic EP maps were created using interpolation techniques. RESULTS: No cerebral responses were recorded with sham stimulation. A significant intensity-dependent increase of the major EP peaks (N1-P2) was observed between 5 and 25 mA (P < 0.05). A significant shortening of the mean peak latency of the first peak (N1) occurred with increasing stimulus intensity (P < 0.0001). Topographic brain maps localized the early EP peaks centrally, whereas later peaks were spread symmetrically over the centroparietal region. CONCLUSIONS: The clear dose-response relationship in the brain response with increasing stimulus intensities probably reflects increased recruitment of afferent fibers. Early peaks originate from deep central brain structures, whereas later peaks are localized exclusively in cortical regions.

Adult↗

Seizure control after stimulation of the vagus nerve: clinical outcome measures.

BACKGROUND: Currently, decreases in seizure frequency are the accepted efficacy outcome measure of therapeutic interventions in the management of patients with epilepsy. In a longitudinal randomized controlled trial of 10 subjects with intractable complex partial seizures who received left vagal nerve stimulation (VNS) to control seizures, it was found that the total number of consecutive seizure-free days is a significant efficacy outcome measure. Unlike measures in which percentage decreases in seizure frequency are calculated, measures of consecutive seizure days indicate improvement in the amount of time for which patients may function at a higher level in activities of daily living. METHODS: Fourteen day blocks of consecutive seizure-free days and 14 day blocks of consecutive days in which subjects had seizures were tabulated. RESULTS: A Pearson correlation coefficient showed that prior to VNS subjects had few, if any, seizure free blocks of time and after VNS they had more blocks of time seizure free r = -1.00 and r = -0.99. The blocks of seizure-free days increased tenfold (mean 0.85 to mean 8.00) from 1991-1995 while mean seizure frequency in those blocks in which subjects had seizures only decreased from (mean 20.14 to mean 17.59) for the same time period. Correlations between total number of seizures after 24 months of VNS and after 50 months of VNS were r = 0.85 showing a consistency in the effect of VNS. CONCLUSIONS: Monitoring the number of consecutive seizure-free days is a significant clinical outcome measure of VNS.

Adult↗

Chronic stimulation of the left vagus nerve: cognitive motor effects.

BACKGROUND: Early studies of cognitive motor control have shown deficits in complex reaction time tests of epileptic subjects. The purpose of this efficacy study was to determine whether chronic (28 months) stimulation of the left vagus nerve (VNS) to control seizures increased these deficits in 6 epileptic subjects with intractable complex partial seizures. METHODS: Subjects were assessed for simple reaction time, Test A, and subsequent Tests B and C which involved more complex cognitive strategies. Tests were done pre-operatively (SI) and at intervals, 6-8 weeks (S2-S3), and at 6 month intervals (S4-S6) over a 28 month period. Data were collected and collated on an Apple II E computer (Apple, Cupertino CA. U.S.A.) and on electronic switch pad. Data were analyzed using a repeated measures analysis of covariance technique with 2 within subject factors, day, and time of day. RESULTS: 2/11 cognitive measures showed a statistically significant difference. Error rate associated with Test A (simple reaction time) significantly decreased for the factor of day (repeated visits) p = .01. For Test C, error rates decreased in the afternoon (p = .03). This test involved the subjects ability to respond quickly to one signal while simultaneously ignoring a second signal. Data analysis of the covariate showed that the effects of VNS are weak in comparison to baseline differences and the frequency of nerve stimulation negatively predicts the number of wrong errors. High frequency stimulation results showed fewer errors than low frequency stimulation T = -2.31, p = .03. CONCLUSION: Chronic stimulation of the left vagus nerve to control seizure activity does not impair cognitive motor control.

Adult↗

Chronic stimulation of the left vagus nerve in epilepsy: balance effects.

BACKGROUND: Stimulation of the left vagus nerve (VNS) has been shown to control seizures in double blinded crossover studies in man. Animal studies have reported vagal afferent induced depression of nociceptive and motor reflexes which may be caused by an effect on the descending reticular system controlling spinal cord function. Anticonvulsant drug therapy may cause postural instability. The effects of VNS are assessed not only from the perspective of seizure control but also from the view of potential harm to other bodily systems. Long term (2 1/4 years) effects of VNS were compared to postural stability analyses. METHODS: 8 subjects, 2 were females, mean age 34.5 +/- 8.23 SD years, with intractable complex partial seizures, taking 3 anticonvulsant drugs were assessed for postural stability in quiet standing and while moving forwards, backwards and sideways with eyes open (EO) and eyes closed (EC). Data were collected and collated using an AMTI Biomechanics immovable forceplate, Newton M.A. U.S.A. The study design was longitudinal with pre-operative baseline data collected prior to neurostimulation and at intervals post operatively. RESULTS: 4/8 balance measures showed significant changes from pre-operative values and after 2 1/4 years of stimulation. Area of sway (EO) in quiet standing p = .022 and total sway (EC) in the moving state p = .019 and total sway (EC) in quiet standing showed an increase in sway p = .003. Area of sway (EC) p = .004 tended to decrease. Regression analysis for frequency of stimulation showed an increase in sway with higher frequencies T = 1.99, P = .05. CONCLUSION: Chronic VNS does not augment postural instability.

Adult↗

Modulation of neurocardiac function by oesophageal stimulation in humans.

1. The heart and the oesophagus have similar sensory pathways, and sensations originating from the oesophagus are often difficult to differentiate from those of cardiac origin. We hypothesized that oesophageal sensory stimuli could alter neurocardiac function through autonomic reflexes elicited by these oesophageal stimuli. In the present study, we examined the neurocardiac response to oesophageal stimulation and the effects of electrical and mechanical oesophageal stimulation on the power spectrum of beat-to-beat heart rate variability in male volunteers. 2. In 14 healthy volunteers, beat-to-beat heart rate variability was compared at rest and during oesophageal stimulation, using either electrical (200 microns, 16 mA, 0.2 Hz) or mechanical (0.5 s, 14 ml, 0.2 Hz) stimuli. The power spectrum of beat-to-beat heart rate variability was obtained and its low- and high-frequency components were determined. 3. Distal oesophageal stimulation decreased heart rate slightly (both electrical and mechanical) (P < 0.005), and markedly altered heart rate variability (P < 0.001). Both electrical and mechanical oesophageal stimulation increased the absolute and normalized area of the high-frequency band within the power spectrum (P < 0.001), while simultaneously decreasing the low-frequency power (P < 0.005). 4. In humans, oesophageal stimulation, whether electrical or mechanical, appears to amplify respiratory-driven cardiac vagoafferent modulation while decreasing sympathetic modulation. The technique provides access to vagoafferent fibres and thus may yield useful information on the autonomic effects of visceral or oesophageal sensory stimulation.

Adolescent↗

The cerebral response to electrical stimuli in the oesophagus is altered by increasing stimulus frequencies.

Recording of cerebral evoked responses (EP) allows the assessment of visceral afferent pathways and gut-brain communication, but the optimal stimulation parameters remain to be established. The present study determined the optimal stimulation frequency of electrical stimulation of the oesophagus to elicit EP responses. In 13 healthy male volunteers (24.1 +/- 5.9 years), a 5 mm stainless-steel electrode was placed in the distal oesophagus for electrical stimulation (ES). EP were recorded from 21 scalp electrodes placed according to the 10/20 International system. ES (15 mA, 200 microseconds) were delivered in repeated series of 24 stimuli. Stimulus frequency was randomly altered in different series using a pseudologarithmic range (0.1, 0.2, 0.3, 0.5, and 1 Hz). Two series of stimuli were applied using each stimulation frequency. Two-dimensional topographic brain maps were created using interpolation techniques at each stimulation frequency. With increasing stimulus frequency, a significant and progressive decrease of EP amplitudes was observed between frequencies of 0.1 Hz and 1.0 Hz (P1/N2: 7.6 +/- 1.2 vs 1.4 +/- 0.3* microV, N2/P2: 17.2 +/- 1.7 vs 4.6 +/- 0.4* microV, P2/N3: 6.9 +/- 0.7 vs 4.2 +/- 0.5* microV; * = P < 0.05). In addition, there was a significant shortening of the mean peak latency of the intercalated P2 peak (P < 0.0005), with a similar trend for the P3 peak (P < 0.06), with increasing stimulus frequency from 0.1-1.0 Hz. Topographic brain maps localized the maximal early peaks (N1,P1.N2) in the paracentral cortical region (C3, Cz, C4), whereas the later peaks (P2 to P3) were symmetrically spread over the centroparietal and temporal regions (Cz, Pz, T5, T4). There was no difference in the cortical location of maximal EP amplitudes with increasing stimulus frequency. In conclusion, there is a clear relationship between stimulus frequency and amplitude of EP, suggesting rapid attenuation of the cerebral autonomic neural responses with increased electrical stimulation frequency. The effect of increased frequency on peak latencies suggests an alteration of stimulus processing in the thalamocortical region due to an altered perception of stimuli. Early EP peaks originate from basal structures of primarily the dominant hemisphere, while later peaks are localized in centroparietal cortical regions.

Adult↗

Neurocardiac response to esophageal electric stimulation in humans: effects of varying stimulation frequencies.

The purpose of this study was to determine if the cardioautonomic responses to esophageal electric stimulation were mediated entirely through modulation of respiratory frequency or a direct vagal effect. We performed electric stimulation of the esophagus in 13 healthy male controls (24 +/- 6 yr) using a manometric catheter to which a stainless steel electrode was attached. Stimulation frequencies ranged from 0.1 to 1 Hz and were applied in random fashion. We computed the power spectra of the heart rate variability and respiratory frequency as measures of autonomic function. Electric stimulation of the esophagus produced significant increases in the high-frequency power of the heart rate autospectrum at all stimulation frequencies (maximal at 0.2 Hz). However, regardless of the frequency of esophageal stimulation, the respiratory rate was not changed from baseline. These studies indicate that enhancement of cardiac vagal modulation observed in response to esophageal electric stimulation is not primarily due to changes in respiratory frequency, but rather occurs through a direct, vagally mediated action through sensory neural pathways involving vagal esophageal afferents.

Adult↗

Cerebral evoked responses to gastrointestinal stimulation in humans.

Recent advances have permitted recording of evoked potentials (EPs) in response to electrical and mechanical stimulation of the gastrointestinal (GI) organs via methods used primarily in clinical neurophysiology. Current research involving stimulation of the esophagus, rectum, and colon, and recording the corresponding responses on the scalp, is being practiced in only a few laboratories. This review examines the engineering aspects of recording EPs, such as characteristics of the stimuli, placement of stimulus electrodes in the GI tract, and enhancement of evoked potential signals. We also discuss the physiological concepts involved in the generation of EPs, and how these compare with somatosensory evoked responses. Current experimental techniques employed by various investigators and results reported from their laboratories are compared. We believe that cerebral EPs to GI stimulation could be useful in studying a number of pathophysiological conditions such as gastroesophageal reflux disease, diffuse esophageal spasm, chronic inflammatory bowel disorders, chronic abdominal pain, and irritable bowel syndrome, among others. We hope that the present review will generate interest in the use of EPs arising out of GI stimulation, aiding in understanding their physiological implications in healthy subjects and in GI disorders.

Cerebral Cortex↗

Identification of ligand binding determinants in the somatostatin receptor subtypes 1 and 2.

The somatostatin (SRIF) receptors (SSTRs) 1 and 2 bind SRIF and SRIF 28 with high affinity, although a number of synthetic hexapeptide and octapeptide analogs of SRIF bind selectively to SSTR2. Extracellular loop three and its adjoining trans-membrane-spanning regions contain elements essential for the binding of such analogs to murine SSTR2. In particular, a stretch of amino acids from residues 294-297 (FDFV) in murine SSTR2 in trans-membrane domain seven can determine affinity for the SSTR2-selective analogs. Within this region, Phe294 has previously been predicted to be essential for the binding of octapeptides (Kaupmann, K., Bruns, C., Raulf, F., Weber, H., Mattes, H., and Lubbert, H. (1995) EMBO J. 14, 727-735) based on the observation that SSTR1 can bind the octapeptide SMS-201-995 with reasonable affinity after a Ser-to-Phe conversion in the analogous region of this receptor (SSTR1S305F). We find that SSTR1S305F has low affinity for a number of SSTR2-selective hexapeptides, suggesting that these analogs have different binding requirements than SMS-201-995. A correlation is seen between the ability of SSTR1S305F to bind hexapeptide analogs and the presence of a phenylalanine, but not tyrosine, at position two in these small cyclic molecules. Thus, a single hydroxyl group in hexapeptides can play a critical role in determining receptor binding to these receptor mutants. We also find that the second extracellular loop of SSTR1 is important for the selectivity of certain SRIF agonists for binding to SSTR1. Taken together, our data indicate that there are multiple elements in the somatostatin receptors that can determine the binding affinity and selectivity of peptide analogs.

Amino Acid Sequence↗

A systematic map of direction preference in primary visual cortex.

Neurons in the primary visual cortex respond selectively to the orientation of edges and their direction of motion. Orientation preference is mapped in a systematic fashion across the cortical surface, such that neurons in adjacent columns have similar but slightly shifted preferred orientations. Microelectrode studies have suggested that direction preference is also arranged in a systematic fashion, but exactly how this response property is mapped remains unclear. Here we show by optical imaging of intrinsic signals in ferret cortical area 17 that there is a mosaic-like map of direction preference. This map consists of numerous regions within which direction preference changes in a slow, continuous fashion. These regions are separated by winding boundaries (fractures) across which direction preference shifts abruptly, often by 180 degrees. Comparison of direction and orientation preference maps shows that these fractures subdivide iso-orientation domains into regions selective for opposite directions of motion.

Animals↗