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Parallel, interdependent channels for location and orientation in sensorimotor transformations for reaching and grasping.

1. Subjects were presented with a cylinder, whose orientation with respect to the vertical and location in space varied from trial to trial. They grasped a similar cylinder in their hands and were instructed to align the grasped cylinder with the target cylinder. In some experiments the task was performed from memory, and subjects attempted to reproduce both location and orientation of the cylinder. In others, they attempted to reproduce only its orientation, either from memory or while the cylinder was in view. 2. Multivariate linear regression analysis was used to determine persistent and variable errors in performance. This analysis related the subjects' performance (reproduced orientation) to target parameters (location and orientation). 3. We have interpreted the experimental results starting from the assumption that there are two parallel neural processes underlying reaching and grasping: one relating proximal arm motion to target location and the other relating distal hand motion to target orientation. 4. Variable errors did not vary with task conditions, even when subjects were asked to dissociate target orientation from target location by matching target orientation with the arm at the side, irrespective of the location of the target. This finding suggests that the neural transformations involving target location and target orientation are performed in parallel. 5. Persistent errors did vary with task condition. The subjects made the largest errors in matching target orientation when the target was in view, but they were asked to match its orientation at a location that differed from that of the target. These errors depended mostly on the elevation of the target and on its slant (inclination relative to the vertical). They were related to the posture of the arm in a manner that suggested that matching orientation is influenced by both extrinsic (spatial) and intrinsic (arm posture) parameters. 6. The fact that persistent errors depend on target location and on arm posture as well as on target orientation implies that the neural transformation from target orientation to hand orientation is not independent of the transformation dealing with target location.

Attention↗

Uniform discrimination of pattern orientation by honeybees.

To explore how honeybees, Apis cerana, discriminate the orientation of patterns, we trained workers to discriminate between a black stripe of a certain orientation on a white disc and a pure white disc. We tested trained bees for their ability to discriminate between the trained orientation and deviations from it. This was done either in a dual choice situation where the bees had to choose between the trained orientation and one deviation from it at a time, or in a multiple choice situation where bees had to choose simultaneously between the trained orientation and 11 successive deviations from it. In the dual choice situation, bees did not discriminate behaviourally between the trained orientation and deviations up to 25 degrees, whereas in a multiple choice situation, they discriminated between the trained orientation and a deviation of 15 degrees or more. Thus, orientation can be analysed more precisely in multiple choice experiments. The response of the bees was independent of the orientation of the trained orientation; the 12 different trained orientations all yielded identical results. This finding, considered together with a model that we present for orientation discrimination, suggests that at least three orientation-sensitive channels (a neuron or a set of neurons that respond maximally to a particular orientation) participate in the analysis of pattern orientation. (c) 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Evaluation of the energetic contribution of interhelical Coulombic interactions for coiled coil helix orientation specificity.

Coiled coils are formed by two or more alpha-helices that align in a parallel or an antiparallel relative orientation. The factors that determine a preference for a given relative helix orientation are incompletely understood. The helix orientation preference for the designed coiled coil, Acid-a1-Base-a1, was measured previously. This model system therefore provides a means for the experimental determination of the energetic contribution of a variety of interactions to helix orientation specificity. The antiparallel preference for Acid-a1-Base-a1 is imparted by a single buried polar interaction. Interhelical Coulombic interactions between residues at the e and g positions have been proposed to influence helix orientation preference. In the Acid-a1-Base-a1 heterodimer, potentially attractive Coulombic interactions are expected in both orientations. To determine the energetic consequences of Coulombic interactions for helix orientation preference, we have positioned a single charged residue in each peptide such that exclusively favorable interhelical Coulombic interactions can occur only in the parallel orientation. In contrast, two potentially repulsive interactions are expected in the antiparallel orientation. Because the buried polar interaction can occur only in the antiparallel orientation, interhelical Coulombic interactions favor the parallel orientation and the potential to form a buried polar interaction favors the antiparallel orientation. We find no clear preference for an antiparallel orientation in the resulting heterodimer, Acid-Ke-Base-Eg, suggesting that interhelical Coulombic interactions and a buried polar interaction are of approximately equal importance for helix orientation specificity. Stability measurements indicate that maintenance of all favorable electrostatic interactions and/or avoidance of two potentially repulsive interactions contributes approximately 2.1 kcal/mol to helix orientation preference.

Amino Acid Sequence↗

Pattern adaptation and cross-orientation interactions in the primary visual cortex.

The responsiveness of neurons in the primary visual cortex (V1) is substantially reduced after a few seconds of visual stimulation with an effective pattern. This phenomenon, called pattern adaptation, is uniquely cortical and is the likely substrate of a variety of perceptual after-effects. While adaptation to a given pattern reduces the responses of V1 neurons to all subsequently viewed test patterns, this reduction shows some specificity, being strongest when the adapting and test patterns are identical. This specificity may indicate that adaptation affects the interaction between groups of neurons that are jointly activated by the adapting stimulus. We investigated this possibility by studying the effects of adaptation to visual patterns containing one or both of two orientations--the preferred orientation for a cell, and the orientation orthogonal to it. Because neurons in the primary visual cortex are sharply tuned for orientation, stimulation with orthogonal orientations excites two largely distinct populations of neurons. With intracellular recordings of the membrane potential of cat V1 neurons, we found that adaptation to the orthogonal orientation alone does not evoke the hyperpolarization that is typical of adaptation to the preferred orientation. With extracellular recordings of the firing rate of macaque V1 neurons, we found that the responses were not reduced by adaptation to the orthogonal orientation alone nearly as much as by adaptation to the preferred orientation. In the macaque we also studied the effects of adaptation to plaids containing both the preferred and the orthogonal orientations. We found that adaptation to these stimuli could modify the interactions between orientations. It increased the amount of cross-orientation suppression displayed by some cells, even turning some cells that showed cross-orientation facilitation when adapted to a blank stimulus into cells that show cross-orientation suppression. This result suggests that pattern adaptation can affect the interaction between the groups of neurons tuned to the orthogonal orientations, either by increasing their mutual inhibition or by decreasing their mutual excitation.

Animals↗

Tryptophans in membrane proteins: indole ring orientations and functional implications in the gramicidin channel.

Orientational constraints generated from solid-state NMR of uniformly aligned gramicidin A in hydrated lipid bilayers have been used to determine the indole ring orientations for the four tryptophans of the gramicidin A monomer with respect to the bilayer normal and the channel axis. 15N epsilon 1 labeled tryptophan has been incorporated into gramicidin at positions 9, 11, 13, and 15. The chemical shift tensor orientation has been oriented with respect to the N-H bond via doubly labeled sample in which the 15N epsilon 1-1H has been exchanged for 2H. By observation of the dipolar coupled 15N chemical shift powder pattern of the amino acid, sigma cc has been shown to be perpendicular to the plane of the ring and that sigma aa makes an angle of 25 degrees with respect to the N-H bond. The indole ring orientations were obtained from a consideration of both the chemical shift and the 15N-1H dipolar interaction. These four rings have very similar orientations with respect to the bilayer normal as given by the range of angles between the bilayer and ring normals (64-67 degrees). Furthermore, the N-H bond orientations with respect to the bilayer normal varies by only 10 degrees among the four sites. This orientational analysis has been based on an assumption that large amplitude librational motions in the hydrated bilayer samples are not averaging the nuclear spin interactions. This assumption was verified by analyzing the 2H quadrupole spectra of d5-Trp11-labeled gramicidin A in oriented preparations. The orientations predicted for the five C-H bonds in the indole ring from the 15N data agreed (root-mean-square deviation of 3.7 degrees) with the observed orientations from quadrupole splittings of the C-D bonds in the ring. From the orientation of the indole rings with respect to the bilayer normal and a polypeptide backbone conformation, the four typtophans of the gramicidin monomer are oriented with respect to the backbone of the channel conformation. The similarity among the indole orientations with respect to the bilayer normal is therefore consistent with the electrophysiological results that the individual replacement of the indole rings with phenyl rings results in a incremental decrease in the conductance of the channels formed. The indole orientations with respect to the backbone as defined by the side-chain torsion angles is not uniquely determined but yields a discrete set of possible values.(ABSTRACT TRUNCATED AT 400 WORDS)

Amino Acid Sequence↗

Factors associated with students' orientations to nursing.

Factors associated with students' orientations to nursing This paper presents the results of a study focusing on the factors associated with orientations to nursing. Students' orientations to nursing have not as yet been a focus of nursing research. In some other professions, however, professional orientation has been associated with learning motivation and study performance, and has been seen as a predictor of work satisfaction. In this study, students' orientations to nursing were defined in terms of caring, nursing expertise and life orientation. The hypothesis of whether students' pre-educational experiences of nursing, gender, choice of nursing specialty, problems with nursing studies and intention to stay in nursing were associated with different orientations was tested. The extent to which students were orientated to caring, nursing expertise and their own life was also examined. The orientation to nursing measurement tool, which has been developed on the basis of a qualitative study, was used to collect the data. Nurse teachers collected the data from nursing students (n=184) who were studying in three different nursing programmes in Finland. Non-parametric assessments (Mann-Whitney U-test and Kruskal-Wallis test) of the differences between the students' orientations were carried out. A majority of the students were highly life-orientated, and two-thirds had average nursing expertise or caring orientation scores. The results supported the study hypothesis of an association between students' orientations and their gender, choice of nursing speciality, problems with nursing studies and intention to stay in nursing. However, the hypothesis of an association between students' pre-educational nursing experiences and orientation to nursing was not supported. The contradictions between students' orientation to nursing and the philosophy of nursing underlying the study programme may be a source of motivational problems and dissatisfaction with nursing education. Therefore, nurse educators are challenged to discuss curriculum matters and student supervision in order to promote flexibility in planning personal study programmes.

Adolescent↗

The reproduction of vertical and oblique orientations in the visual, haptic, and somato-vestibular systems.

This study investigates whether the vertical orientation may be predominantly used as an amodal reference norm by the visual, haptic, and somato-vestibular perceptual systems to define oblique orientations. We examined this question by asking the same sighted adult subjects to reproduce, in the frontal (roll) plane, the vertical (0 degree) and six oblique orientations in three tasks involving different perceptual systems. In the visual task, the subjects adjusted a moveable rod so that it reproduced the orientation of a visual rod seen previously in a dark room. In the haptic task, the blindfolded sighted subjects scanned an oriented rod with one hand and reproduced its orientation, with the same hand, on a moveable response rod. In the somato-vestibular task, the blind-folded sighted subjects, sitting in a rotating chair, adjusted this chair in order to reproduce the tested orientation of their own body. The results showed that similar oblique effects (unsigned angular error difference between six oblique orientations and vertical orientation) were observed across the three tasks. However, there were no positive correlations between the visual, haptic, and somato-vestibular oblique effects. Moreover, in some oblique orientations, there was a tendency to overestimate the angle between the oblique orientation and the vertical orientation. This effect varied according to the orientation value and the modality. Taken together, these findings suggest that although vertical orientation is used as a reference norm in the visual, haptic, and somato-vestibular systems to define oblique orientations, specific processing mechanisms seem to be at work in each perceptual system.

Adult↗

Cellular analogs of visual cortical epigenesis. I. Plasticity of orientation selectivity.

A differential pairing procedure was applied in vivo to individual neurons in the primary visual cortex of anesthetized paralyzed cats, in order to produce changes in their relative orientation preference. While we recorded from a single cell, its visual response to a light bar was driven iontophoretically to a "high" level when stimulating with an initially nonpreferred orientation (S+), and alternately reduced to a "low" level when stimulating with the preferred orientation (S). This associative procedure was devised to test the possible role of neuronal coactivity in controlling the plasticity of orientation selectivity. Among 87 cells tested, 35 (40%) showed significant long-lasting changes, either in the relative orientation preference for the two "paired" stimuli S+ and S-, in the global orientation tuning profile, or in both. Measurements of relative orientation preference demonstrated significant effects in 27 cells (31%), all in favor of the positively reinforced orientation (S+). Modifications of orientation selectivity (studied over the entire orientation spectrum in 45 of the conditioned cells) usually consisted (21 out of 25 modified cells) of a competitive reorganization of the orientation tuning curve: the preferred orientation shifted toward S+, and a loss of relative visual responsiveness was observed for orientations close to the negatively reinforced orientation (S-). The largest changes were found in deprived kittens at the peak of the critical period, although the probability of inducing a significant change studied during the first year of postnatal life was independent of age. These functional modifications demonstrated at the cellular level are analogous to those induced by a global manipulation of the visual environment, when only a restricted spectrum of orientations is experienced during the critical period. Our results support the hypothesis that covariance levels between pre- and postsynaptic activity determine the sign and the amplitude of the modification of efficacy of cortical synapses.

Animals↗

Goal orientations of young male ice hockey players and their parents.

In this study, the researchers investigated the relationship between parent and player dispositional goal orientations associated with playing youth hockey. The authors used the Task and Ego Orientation in Sport Questionnaire (J. L. Duda & J. Whitehead, 1998) to measure task and ego orientation in 123 boys (10-13 years old) and 1 of their parents. Sons rated their own goal orientations for hockey and their perceptions of their parent's goal orientations. Parents rated their goal orientations for their son and their perceptions of their son's goal orientations. Mothers and fathers did not differ in their goal orientations for their son. Travel-team and nontravel-team players did not differ. For ego orientation, the son's self-ratings correlated significantly with the parent's goals for the son, but not for task orientation. Sons reported being significantly more ego-oriented than their parents desired. Sons perceived that their parents had goal orientations similar to their own. The data from this study are congruent with the assertion that parents socialize their children's goal orientations and that ego orientation may be more salient and easily communicated than task orientation.

Adolescent↗

Response variability and orientation discrimination of single cells in striate cortex of cat.

The response of single cells in the striate cortex of cat to a moving light bar of variable orientation was measured by a method providing data on the mean response as well as the standard deviation (SD) at the different stimulus orientations. At the optimal stimulus orientation the SD was about 1/3 of the mean response. Marked differences in this respect were found between simple and complex cells, i.e., the SD for the simple cells was about 1/2 of the mean response and about 1/4 for the complex cells. The variation coefficient (Vc = SD/mean) was minimal at the optimal orientation and increased relatively in the same manner for simple and complex cells as the stimulus orientation was varied away from optimal orientation. The Vc varied with the mean response at optimal orientation in a nonlinear manner. A function is proposed which fits this relationship and which is equally applicable for both simple and complex cells. The mean orientation discrimination (MOD) was defined as that change in orientation angle away from the optimal which produced a response statistically different--on the 1% level--from the response to the optimal orientation. There were differences in MOD between the two sides of the orientation tuning curve: the mean of the smaller of the two values was 13.5 deg and of the larger 19.7 deg. No significant difference in MOD was found between simple and complex cells despite the fact that the halfwidth of the tuning curves for the two cell types was 19.5 deg and 31.6 deg, respectively. The preciseness in localization of the most sensitive part within the receptive field of single cells was calculated from the variability in time of occurrence of the smallest interspike interval. The degree of preciseness was found to be of the order of 1/4 of the receptive field diameter in both simple and complex cells. When nonoptimal stimulus orientations were presented, the preciseness significantly decreased in complex cells whereas it remained unchanged in simple cells. It is suggested that the same type of intracortical wiring produces orientation selectivity in simple and complex cells, and that the differences in tuning width are mainly due to a larger extension of inhibitory fields in the simple cells. Considering the cortical visual cells as elementary units in a network built for orientation detection and discrimination, the tuning width seems of minor importance for that function.

Animals↗

Fine orientation discrimination and shape constancy in young infants.

The present study aimed to investigate simultaneously fine orientation discrimination and shape constancy in young infants. The design employed two variants of the habituation paradigm. Infants in one group were habituated to a single orientation (5 or 15 degrees) of a single stimulus presented repeatedly, and they were then tested with the complementary orientation (15 or 5 degrees). Infants in a second group were habituated to several orientations (5, 10, and 15 degrees) of the same stimulus, and they were then tested with a familiar orientation of the stimulus, with two novel orientations of the same stimulus, and with a new stimulus. Between-groups comparison showed that infants habituated more efficiently to re-presentations of a single orientation than to multiple orientations of the same stimulus, providing evidence of fine orientation discrimination; posthabituation comparison within the single-orientation group confirmed that infants discriminated small orientation changes. Posthabituation comparison within the multiple-orientation group showed that infants generalized over novel orientation changes of the familiar stimulus though they discriminated change to a novel stimulus. Cumulatively, the results of this study demonstrate that under one set of conditions young infants show sensitivity to relatively fine variations in pattern orientation, but that under a different set of conditions young infants give evidence of shape constancy with the same patterns.

Discrimination Learning↗

Double orientation tuning in the cat visual cortex units.

Orientation tuning of 271 neurons of the cat visual cortex (area 17) was studied with a light bar flashing in the receptive field. Under different conditions, 27-57% of units were found to have double-orientation tuning: they demonstrated the main preferred orientation and an additional preferred orientation. The statistical reliability and reproducibility of additional preferred orientation were shown. The quality of orientation tuning in the second maximum did not differ statistically from the first one. The angle between preferred orientation and additional preferred orientation was either 90 degrees (29% of cases) or an acute one (60.1 +/- 3.1 degrees, 71% of cases). The ratio of discharge frequency in responses to additional preferred orientation and preferred orientation was equal to 0.74 +/- 0.05. Neurons with double-orientation tuning clearly preferred 67 degrees and 157 degrees, while monomodal units preferred 0 degrees and 90 degrees. Probability of the double-tuning increased under bar lengths of near 3 degrees and near 10 degrees and with increase of stimulus/background contrast. At the same time some neurons displayed double-orientation tuning only with relatively low stimulus/background contrast. The proportion of units with double-orientation tuning was lowered by about 1.5-times under general Nembutal narcotization as compared with local anesthesia of the animal. In about one-third of units simultaneous stimulation by two flashing lines crossing in the receptive field center under an angle specific for the cell, evoked a response from 1.5 to four times larger than to the preferred orientation.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Are neurons in cat posteromedial lateral suprasylvian visual cortex orientation sensitive? Tests with bars and gratings.

There is controversy in the literature concerning whether or not neurons in the cat's posteromedial lateral suprasylvian (PMLS) visual cortex are orientation selective. Previous studies that have tested cells with simple bar stimuli have found that few, if any, PMLS cells are orientation selective. Conversely, studies that have used repetitive stimuli such as gratings have found that most or all PMLS cells are orientation selective. It is not known whether this difference in results is due to the stimuli used or the laboratories using them. The present experiments were designed to answer this question by testing individual PMLS neurons for orientation sensitivity with both bar and grating stimuli. Using quantitative response measures, we found that most PMLS neurons respond well enough to stationary flashed stimuli to use such stimuli to test for orientation sensitivity. On the basis of these tests, we found that about 85% of the cells with well-defined receptive fields are orientation sensitive to flashed gratings, and a similar percentage are orientation sensitive to flashed bars. About 80% of the cells were orientation sensitive to both types of stimuli. The preferred orientations typically were similar for the two tests, and they were orthogonal to the preferred direction of movement. The strength of the orientation sensitivity (measured as the ratio of discharge to the preferred and nonpreferred orientations) was similar to both types of stimuli. However, the width of the orientation tuning curves was systematically broader to bars than to gratings. Several hypotheses are considered as to why previous studies using bars failed to find evidence for orientation sensitivity. In addition, a mechanism for the difference in orientation tuning to bars and gratings is suggested.

Animals↗

Iso-orientation domains in cat visual cortex are arranged in pinwheel-like patterns.

The mammalian cortex is organized in a columnar fashion: neurons lying below each other from the pia to the white matter usually share many functional properties. Across the cortical surface, cells with similar response properties are also clustered together, forming elongated bands or patches. Some response properties, such as orientation preference in the visual cortex, change gradually across the cortical surface forming 'orientation maps'. To determine the precise layout of iso-orientation domains, knowledge of responses not only to one but to many stimulus orientations is essential. Therefore, the exact depiction of orientation maps has been hampered by technical difficulties and remained controversial for almost thirty years. Here we use in vivo optical imaging based on intrinsic signals to gather information on the responses of a piece of cortex to gratings in many different orientations. This complete set of responses then provides detailed information on the structure of the orientation map in a large patch of cortex from area 18 of the cat. We find that cortical regions that respond best to one orientation form highly ordered patches rather than elongated bands. These iso-orientation patches are organized around 'orientation centres', producing pinwheel-like patterns in which the orientation preference of cells is changing continuously across the cortex. We have also analysed our data for fast changes in orientation preference and find that these 'fractures' are limited to the orientation centres. The pinwheels and orientation centres are such a prominent organizational feature that it should be important to understand their development as well as their function in the processing of visual information.

Animals↗

A model for the formation of orientation columns.

A mathematical model is proposed to describe the formation of orientation columns in mammalian visual cortex. The model is similar in concept to that proposed for ocular dominance column formation (Swindale 1980), the essential difference being that orientation is a vector rather than a scalar variable. It is assumed that initially orientation selectivity is weak and randomly distributed, and that selectivity develops in such a way that the orientation preferences of neurons less than about 200 microns apart tend to change in a similar direction, whereas the preferences of cells further apart tend to develop in opposite directions. No hypotheses are made about the anatomical or physiological basis of these interactions, and it is not necessary to assume that they are the result of environmental stimulation, as with existing models for the development of orientation selectivity (see, for example, von der Malsburg, 1973). The model reproduces the experimental data on orientation columns: roughly linear sequences of orientation change are produced, and these alternate unpredictably between clockwise and anticlockwise directions of change. Continuous sequences may span several 180 degrees cycles of rotation. The sequences are generally smooth, but abrupt discontinuities of up to 90 degrees also occur. The iso-orientation domains for large orientation ranges (60-90 degrees) are periodically spaced branching stripes that resemble those demonstrated in animals by the 2-deoxyglucose technique. The domains for narrower orientation ranges are periodically spaced but are more irregular in shape, though sometimes thin and elongated. The model makes a number of predictions that can be tested experimentally. Of particular interest are the discontinuities in the orientation sequences: these should be distributed with a spacing roughly equal to, or half, that of the iso-orientation domains. Each should be surrounded by one or two complete sets of iso-orientation domains, and each may be associated with regions where cells are not orientation selective. These regions may be more extensive in younger animals, when the columns are at an intermediate stage of formation, and less numerous where the columns run parallel and unbranched over large areas.

Animals↗

Relationship between lateral inhibitory connections and the topography of the orientation map in cat visual cortex.

The functional and structural topography of lateral inhibitory connections was investigated in visual cortical area 18 using a combination of optical imaging and anatomical tracing techniques in the same tissue. Orientation maps were obtained by recording intrinsic signals in regions of 8.4-19 mm2. To reveal the inhibitory connections provided by large basket cells, biocytin was iontophoretically injected at identified orientation sites guided by the pattern of surface blood vessels. The axonal and dendritic fields of two retrogradely labelled large basket cells were reconstructed in layer III. Their axonal fields extended up to 1360 microns from the parent somata. In addition to single basket cells, the population of labelled basket cell axons was also studied. For this analysis anterogradely labelled basket axons running horizontally over 460-1280 microns from the core of an injection site in layer III were taken into account. The distribution of large basket cell terminals according to orientation preferences of their target regions was quantitatively assessed. Using the same spatial resolution as the orientation map, a frequency distribution of basket cell terminals dependent on orientation specificity could be derived. For individual basket cells, the results showed that, on average, 43% of the terminals provided input to sites showing similar orientation preferences (+/- 30 degrees) to those of the parent somata. About 35% of the terminals were directed to sites representing oblique-orientation [+/- (30-60) degrees], and 22% of them terminated at cross-orientation sites [+/- (60-90) degrees]. Furthermore, the possible impact of large basket cells on target cells at different distances and orientation preferences was estimated by comparing the occurrence of orientation preferences with the occurrence of basket terminals on the distance scale. It was found that a basket cell could elicit iso-orientation inhibition with a high impact between 100-400 and 800-1200 microns, strong cross-orientation inhibition at approximately 400-800 microns, and oblique-orientation inhibition between 300-500 and 700-900 microns from the parent soma. The non-isotropic topography of large basket axons suggests a complex function for this cell class, possibly including inhibition related to orientation and direction selectivity depending on the location of the target cells and possible target selectivity.

Animals↗

Orientational dynamics of T2 DNA during agarose gel electrophoresis: influence of gel concentration and electric field strength.

The understanding, on a molecular level, of the mechanisms responsible for the improved separation in DNA gel electrophoresis when using modulated electric fields requires detailed information about conformational distribution and dynamics in the DNA/gel system. The orientational order due to electrophoretic migration ("electrophoretic orientation") is an interesting piece of information in this context that can be obtained through linear dichroism spectroscopy [M. Jonsson, B. Akerman, and B. Nordén, (1988) Biopolymers 27, 381-414]. The technique permits measurement of the orientation factor S of DNA (S = 1 corresponds to perfect orientation) within an electrophoretic zone in the gel during the electrophoresis. It is reported that the degree of orientation of T2 DNA [170 kilo base pairs (kpb)] is considerable (S = 0.17 in 1% agarose at 10 V/cm) compared to relatively modest orientations of short fragments found earlier (for 23-kbp DNA, S = 0.03 in 1% agarose at 10 V/cm), showing that large DNA coils are substantially deformed during the migration. Growth and relaxation dynamics of the orientational order of the T2 DNA are also reported, as functions of gel concentration (0.3-2%), electric field strength (0-40 V/cm), and pulse characteristics. The rise profile of the DNA orientation, when applying a constant field, is a nonmonotonic function that displays a pronounced overshoot, followed by a minor undershoot, before it reaches steady-state orientation (after 12 s in 1% agarose, 9 V/cm). The orientational relaxation in absence of field shows a multiexponential decay in a time region of some 10 s, when most of the DNA anisotropy has disappeared. A surprising phenomenon is a memory over minutes of the DNA/gel system to previous pulses: with two consecutive rectangular pulses (of the same polarity), the orientational overshoot and undershoot as a response to the second pulse are significantly reduced compared to the first pulse. The time required to recover 90% of their amplitudes is typically 1200 s (1% agarose, 9 V/cm), which may be compared to the time required to relax 90% of the DNA orientation, which is only 6 s. The major part of the over- and undershoot recovery is thus a reorganization of a system in which DNA is already randomly oriented. The different response amplitudes and relaxation times, including the amplitude and recovery time of the overshoot, of the orientational order of DNA in the electrophoretic gel have been studied as functions of gel concentration and field strength. The results are discussed against relevant theories of polymer dynamics.

Chemical Phenomena↗

Neural motion after-effects in the cat's striate cortex: orientation selectivity.

Single striate cortical neurones were recorded from adult cats, lightly anaesthetized with N2O/O2/halothane. The receptive fields for the dominant eye were subjected to direction-specific adaptation by a square-wave grating of optimal spatial frequency and velocity, drifting continuously in each neurone's preferred direction. Recovery of the neural motion after-effect induced by prior adaptation was assessed with the same grating pattern which now moved alternately in the preferred and opposite directions. In controls the same tests for recovery followed a period of exposure to a uniform field of identical luminance to the adapting grating. Three sets of measurements were made to establish whether the adaptation was orientation- as well as direction-specific. In the first, test grating orientation was maintained constant and optimal for each neurone whilst adapting orientation was systematically varied. In the second, test orientation was varied whilst maintaining adapting orientation constant. In the third set, adapting and test orientations were initially fixed at each neurone's optimum; they were next set, non-optimally to one side of the optimum. Results from the latter configuration were compared with similar tests in which the test grating remained at that non-optimal orientation whilst the orientation of the adapting grating was now altered to a new point on the other flank of each neurone's orientation tuning curve that was matched for strength of adaptation. Thus the degree of adaptation was identical in each case, but zero orientation difference between adapting and test gratings in one case was contrasted with a substantial orientation difference in the other. The results from all three sets of data were unequivocal: in simple neurones, and in standard and intermediate classes of complex neurones, but not in special complex neurones, the sequential effects of adapting gratings on the responses and sensitivity to subsequently presented test gratings were maximal when their orientations were matched and optimal for each neurone, less marked when orientations were matched but non-optimal. In conclusion, adaptation induced by pattern motion was orientation- as well as direction-specific only in standard (length summating) and intermediate complex neurones, and in simple cells; in special complex neurones it was not.

Adaptation, Ocular↗