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Adaptation of movement endpoints to perturbations of visual feedback.

We investigated the extent to which humans can quickly adapt their goal-directed arm movements to perturbed feedback. We predicted that the magnitude of adaptation to a changed relationship between vision and kinesthesia would depend on the type of perturbation, being largest when the perturbation can be generalized within egocentric frames of reference. To test this prediction we asked subjects to align a real 5-cm cube so that they could feel, but not see, with a simulation that they saw via a mirror. Subjects made successive movements between target locations in a sequence of adaptation and test phases. During adaptation phases, subjects received continuous visual feedback about the position of the real cube. The feedback was either veridical or perturbed. The perturbations were consistent with either a uniform translation, a scaling or a rotation. The latter two were relative to a central position between all the targets. During test phases, subjects received no visual feedback. We compared test movement endpoints after perturbed feedback with ones after veridical feedback. We found about 40% adaptation to translation, 20% to scaling and 10% to rotation. This difference in magnitude is consistent with the ease with which the transformation can be generalized within egocentric frames of reference. Changing the task so that it required different arm postures did not change the magnitude of adaptation, so postural configuration of the arm does not appear to be critical. Nevertheless, transfer to the unexposed arm was incomplete for translations and rotations, though it was complete for scaling, suggesting that at least part of the adaptation is posture based. We conclude that the adaptation to different kinds of perturbations not only differs in extent but also involves different (egocentric) mechanisms.

Adaptation, Physiological↗

Functional adaptation of reactive saccades in humans: a PET study.

It is known that the saccadic system shows adaptive changes when the command sent to the extraocular muscles is inappropriate. Despite an abundance of supportive psychophysical investigations, the neurophysiological substrate of this process is still debated. The present study addresses this issue using H2(15)O positron emission tomography (PET). We contrasted three conditions in which healthy human subjects were required to perform saccadic eye movements toward peripheral visual targets. Two conditions involved a modification of the target location during the course of the initial saccade, when there is suppression of visual perception. In the RAND condition, intra-saccadic target displacement was random from trial-to-trial, precluding any systematic modification of the primary saccade amplitude. In the ADAPT condition, intra-saccadic target displacement was uniform, causing adaptive modification of the primary saccade amplitude. In the third condition (stationary, STAT), the target remained at the same location during the entire trial. Difference images reflecting regional cerebral-blood-flow changes attributable to the process of saccadic adaptation (ADAPT minus RAND; ADAPT minus STAT) showed a selective activation in the oculomotor cerebellar vermis (OCV; lobules VI and VII). This finding is consistent with neurophysiological studies in monkeys. Additional analyses indicated that the cerebellar activation was not related to kinematic factors, and that the absence of significant activation within the frontal eye fields (FEF) or the superior colliculus (SC) did not represent a false negative inference. Besides the contribution of the OCV to saccadic adaptation, we also observed, in the RAND condition, that the saccade amplitude was significantly larger when the previous trial involved a forward jump than when the previous trial involved a backward jump. This observation indicates that saccade accuracy is constantly monitored on a trial-to-trial basis. Behavioral measurements and PET observations (RAND minus STAT) suggest that this single-trial control of saccade amplitude may be functionally distinct from the process of saccadic adaptation.

Adaptation, Physiological↗

Functions of the nucleus of the optic tract (NOT). I. Adaptation of the gain of the horizontal vestibulo-ocular reflex.

We studied the role of the nucleus of the optic tract (NOT) in adapting the gain of the angular vestibulo-ocular reflex (aVOR) in rhesus and cynomolgus monkeys using lesions and temporary inactivation with muscimol. The aVOR gain was adaptively reduced by forced sinusoidal rotation (0.25 Hz, 60 degrees/s) in a self-stationary visual surround, i.e., a visual surround that moved with the subject, or by wearing x0.5 reducing lenses during natural head movements. The aVOR gains dropped by 20-30% after 2 h and by about 30% after 4 h. Muscimol injections caused a loss of adaptation of contraversive-eye velocities induced by the aVOR, and their gains promptly returned to or above preadapted levels. The gains of the adapted ipsiversive and vertical eye velocities produced by the aVOR were unaffected by muscimol injections. Lesions of NOT significantly reduced or abolished the animals' ability to adapt the gain of contraversive aVOR-induced eye velocities, and the monkeys were unable to suppress these contraversive-eye velocities in a self-stationary surround. The lesions did not affect ipsiversive aVOR-induced eye velocities, and the animals were still able to suppress them. Lesions of NOT also affected the unadapted or "default" aVOR gains. After unilateral NOT lesions, gains of ipsiversive aVOR-induced eye velocity were reduced, while gains of contraversive aVOR-induced eye velocity were either unaffected or slightly increased. Consistent with this, muscimol injections into the NOT of unadapted monkeys slightly reduced the gains of ipsiversive and increased the gains of contraversive-eye velocities by about 8-10%. We conclude that each NOT processes ipsiversive retinal-slip information about visual surround movement relative to the head induced by the aVOR. In the presence of visual surround movement, the retinal-slip signal is suppressed, leading to adaptive changes in the gain of aVOR-induced contraversive horizontal eye velocities. NOT also has a role in controlling and maintaining the current state of the aVOR gains. Thus, it plays a unique role in producing and supporting adaptation of the gain of the horizontal aVOR that is likely to be important for stabilizing gaze during head movement. Pathways through the inferior olive are presumably important for this adaptation.

Adaptation, Physiological↗

A mathematical model of adaptive behavior in quadruped locomotion.

Locomotion involves repetitive movements and is often executed unconsciously and automatically. In order to achieve smooth locomotion, the coordination of the rhythms of all physical parts is important. Neurophysiological studies have related that basic rhythms are produced in the spinal network called, the central pattern generator (CPG), where some neural oscillators interact to self-organize coordinated rhythms. We present a model of the adaptation of locomotion patterns to a variable environment, and attempt to elucidate how the dynamics of locomotion pattern generation are adjusted by the environmental changes. Recent experimental results indicate that decerebrate cats have the ability to learn new gait patterns in a changed environment. In those experiments, a decerebrate cat was set on a treadmill consisting of three moving belts. This treadmill provides a periodic perturbation to each limb through variation of the speed of each belt. When the belt for the left forelimb is quickened, the decerebrate cat initially loses interlimb coordination and stability, but gradually recovers them and finally walks with a new gait. Based on the above biological facts, we propose a CPG model whose rhythmic pattern adapts to periodic perturbation from the variable environment. First, we design the oscillator interactions to generate a desired rhythmic pattern. In our model, oscillator interactions are regarded as the forces that generate the desired motion pattern. If the desired pattern has already been realized, then the interactions are equal to zero. However, this rhythmic pattern is not reproducible when there is an environmental change. Also, if we do not adjust the rhythmic dynamics, the oscillator interactions will not be zero. Therefore, in our adaptation rule, we adjust the memorized rhythmic pattern so as to minimize the oscillator interactions. This rule can describe the adaptive behavior of decerebrate cats well. Finally, we propose a mathematical framework of an adaptation in rhythmic motion. Our framework consists of three types of dynamics: environmental, rhythmic motion, and adaptation dynamics. We conclude that the time scale of adaptation dynamics should be much larger than that of rhythmic motion dynamics, and the repetition of rhythmic motions in a stable environment is important for the convergence of adaptation.

Adaptation, Physiological↗

Local adaptation in the monocarpic perennial Carlina vulgaris at different spatial scales across Europe.

Spatial variation in environmental conditions can lead to local adaptation of plant populations, particularly if gene flow among populations is low. Many studies have investigated adaptation to contrasting environmental conditions, but little is known about the spatial scale of adaptive evolution. We studied population differentiation and local adaptation at two spatial scales in the monocarpic grassland perennial Carlina vulgaris. We reciprocally transplanted seedlings among five European regions (northwestern Czech Republic, central Germany, Luxembourg, southern Sweden and northwestern Switzerland) and among populations of different sizes within three of the regions. We recorded survival, growth and reproduction over three growing periods. At the regional scale, several performance traits and the individual fitness of C. vulgaris were highest if the plants were grown in their home region and they decreased with increasing transplant distance. The effects are likely due to climatic differences that increased with the geographical distance between regions. At the local scale, there were significant interactions between the effects of the population of origin and the transplant site, but these were not due to an enhanced performance of plants at their home site and they were not related to the geographical or environmental distance between the site of origin and the transplant site. The size of the population of origin did not influence the strength of local adaptation. The results of our study suggest that C. vulgaris consists of regionally adapted genotypes, and that distance is a good predictor of the extent of adaptive differentiation at large scales ( > 200 km) but not at small scales. We conclude that patterns of local adaptation should be taken into account for the efficient preservation of genetic resources, when assessing the status of a plant species and during conservation planning.

Adaptation, Biological↗

Parallel genotypic adaptation: when evolution repeats itself.

Until recently, parallel genotypic adaptation was considered unlikely because phenotypic differences were thought to be controlled by many genes. There is increasing evidence, however, that phenotypic variation sometimes has a simple genetic basis and that parallel adaptation at the genotypic level may be more frequent than previously believed. Here, we review evidence for parallel genotypic adaptation derived from a survey of the experimental evolution, phylogenetic, and quantitative genetic literature. The most convincing evidence of parallel genotypic adaptation comes from artificial selection experiments involving microbial populations. In some experiments, up to half of the nucleotide substitutions found in independent lineages under uniform selection are the same. Phylogenetic studies provide a means for studying parallel genotypic adaptation in non-experimental systems, but conclusive evidence may be difficult to obtain because homoplasy can arise for other reasons. Nonetheless, phylogenetic approaches have provided evidence of parallel genotypic adaptation across all taxonomic levels, not just microbes. Quantitative genetic approaches also suggest parallel genotypic evolution across both closely and distantly related taxa, but it is important to note that this approach cannot distinguish between parallel changes at homologous loci versus convergent changes at closely linked non-homologous loci. The finding that parallel genotypic adaptation appears to be frequent and occurs at all taxonomic levels has important implications for phylogenetic and evolutionary studies. With respect to phylogenetic analyses, parallel genotypic changes, if common, may result in faulty estimates of phylogenetic relationships. From an evolutionary perspective, the occurrence of parallel genotypic adaptation provides increasing support for determinism in evolution and may provide a partial explanation for how species with low levels of gene flow are held together.

Adaptation, Biological↗

Adenylate cyclase activity in fish gills in relation to salt adaptation.

The influence of salt adaptation on specific adenylate cyclase activity (measured by conversion of [alpha-32p]-ATP into [alpha-32p]-cAMP) was investigated in gill plasma membranes of rainbow trout (Salmo gairdneri) adapted to various salinities (deionized water, DW; fresh water, FW; 3/4 sea water, 3/4 SW; sea water, SW) and in sea water adapted-mullet (Mugil sp.). Basal activity declined by a factor of 2 in trout with increasing external salinity (pmoles cAMP/mg protein/10 min: 530 in DW, 440 in FW, 340 in 3/4 SW; 250 in SW) and was very low in SW adapted-mullet: 35. The Km for ATP was similar (0.5 mM) in both FW adapted- and SW adapted- trout in either the absence (basal activity) or in the presence of stimulating agents (isoproterenol; NaF) while the Vm varied. Analysis of stimulation ratios with respect to basal levels of the enzyme showed that hormones (glucagon, VIP) and pharmacological substances (isoproterenol, NaF) display a greater potency in high salt than in low salt adapted- fish gills. In contrast, salt adaptation did not have any effect on the regulation of adenylate cyclase by PGE1. These results are interpreted in relation to the general process of osmoregulation.

Adaptation, Physiological↗

Post-receptor chromatic mechanisms revealed by flickering vs fused adaptation.

For both normals and dichromats, adaptation to certain pairs of alternating monochromatic caused reduced sensitivity for detection of some test wavelengths when compared to sensitivity losses caused by adaptation to the fused (supra-CFF) sums of the same lights. Since the two adaptation conditions caused equivalent photopigment depletion, and since sensitivities to some wavelengths did not differ between conditions, the effects are ascribed to post-receptor adaptation. Such post-receptor effects were not obtained when adapting wavelengths did not straddle the presumed "crosspoints" of opponent-colors mechanisms, and, when only one opponent mechanism was adapted, effects were absent for test wavelengths at the crosspoint of that mechanism. For the red vs green system, increasing the intensities of adapting wavelengths from 2.2 to 5.5 log td did not appreciably increase the magnitudes of postreceptor effects. Quantitative accounts for the results are derived from a vector model for color vision. The results provide general support for opponent-colors interpretations of normal and dichromatic vision and suggest that the flicker/fused adaptation paradigm is a useful tool for probing postreceptor mechanisms of vision.

Adaptation, Ocular↗

Link-specific adaptation in the luminance and chromatic channels.

We present evidence that adaptation occurs in the separate links that connect the receptors to the luminance and the chromatic channels. The relative effectiveness of the L and M cone inputs to the luminance and red/green chromatic channels was determined by using, respectively, heterochromatic flicker photometry and a cancellation technique which maintained a pure yellow. The findings provide evidence for a link-specific adaptation model, wherein one receptor system provides two separate links to the luminance and chromatic channels and the outputs of an individual cone to these two postreceptoral channels can be separately attenuated or weighted at each of these links by colored adapting lights. One line of evidence for link-specific adaptation is that colored adapting fields selectively suppress L and M cone inputs to the red/green chromatic channel by a smaller factor than the luminance channel. A second line of evidence is that there is not only a magnitude difference but also a dynamic difference between adaptive processes operating in the luminance and chromatic channels: the luminance channel has a faster gain change and recovery from adaptation than does the chromatic channel. The results, together with other evidence, make it plausible that an important component of light adaptation in cone vision occurs at the cone-bipolar synapse.

Adaptation, Ocular↗

The lateral spread of light adaptation in cat horizontal cell responses.

To investigate the sites of light adaptation processes in the mammalian distal retina, we studied the lateral spread of adaptation signals in cone-driven cat horizontal (H-) cell responses. The size of the adaptation pool is compared to the receptive field for H-cell responses. H-cell activity was recorded intracellularly in the optically intact, in vivo eye. It is demonstrated that light adaptation as measured in H-cells is not a strictly local process. Background light falling outside a central test region effectively modulates the responses to a small test light, flashed on the receptive field center. The integration area for adaptation signals was quantitatively compared to the H-cell receptive field size by measuring the desensitizing effect of background light on the responses to a small centered test spot, as a function of background spot size. The area-adaptation function is comparable to the area-response function but has a slightly smaller length constant. Light adaptation in H-cell responses, therefore, reveals spread of adaptation over a large distance and is probably mediated through lateral interactions in the H-cell network rather than in the cones.

Adaptation, Ocular↗

The influence of contrast adaptation on color appearance.

Most models of color vision assume that signals from the three classes of cone receptor are recoded into only three independent post-receptoral channels: one that encodes luminance and two that encode color. Stimuli that are equated for their effects on two of the channels should be discriminable only to the remaining channel, and are thus assumed to isolate the responses of single channels. We used an asymmetric matching task to examine whether such models can account for changes in color appearance following adaptation to contrast--to temporal variations in luminance and chromaticity around a fixed mean luminance and chromaticity. The experiments extend to suprathreshold color appearance the threshold adaptation paradigm of Krauskopf, Williams and Heeley [(1982) Vision Research, 32, 1123-1131]. Adaptation changes the perceived color of chromatic test stimuli both by reducing their saturation (contrast) and by changing their hue (direction within the equiluminant plane). The saturation losses are largest for test stimuli that lie along the chromatic axis defining the adapting modulation, while the hue changes are rotations away from the adapting direction and toward an orthogonal direction within the S and L-M plane. Similar selective changes in both perceived color and perceived lightness occur following adaptation to stimuli that covary in luminance and chromaticity. The selectivity of the aftereffects for multiple directions within color-luminance space is inconsistent with sensitivity changes in only three independent channels. These aftereffects suggest instead that color appearance depends on channels that can be selectively tuned to any color-luminance direction, and that there are no directions that invariably isolate responses in only a single channel. We use the perceived color changes to examine the spectral sensitivities of the chromatic channels and to estimate the distribution of channels. We also examine how adaptation alters the contrast-response function, how it affects reaction times for luminance and chromatic contrast, the extent to which the aftereffects exhibit interocular transfer, and the way in which the perceived color changes differ from those induced by conventional light adaptation.

Adaptation, Ocular↗

The time course of adaptation in macaque retinal ganglion cells.

The time course of adaptation of cells of the parvocellular (PC) and magnocellular (MC) pathways has been characterized following changes in retinal illuminance or chromaticity. Adaptation state was cycled between high and low luminance levels or between backgrounds with wavelengths metameric to 630 and 570 nm. Cell responsivity was probed with brief bursts of luminance or chromatic modulation. After a change in luminance, adaptation of both MC-cells (tested with a luminance probe) and red-green PC-cells (tested with a chromatic probe) was relatively rapid and largely complete within 100 msec or less. After a change in chromaticity, recovery of responsivity in red-green PC-cells was dependent on cell type. Recovery of responsivity with backgrounds elevating maintained firing was complete within a few seconds, but with backgrounds suppressing cell firing, recovery took many tens of seconds. This very slow time course may be due to a threshold effect. In experiments with backgrounds which selectively adapted one cone type, use of cone-isolating probes indicated that the time course of PC-cell chromatic adaptation may be determined at a site after the subtraction of cone signals. Recovery of responsivity of MC-cells was also prolonged over several seconds following a chromatic change. Our data suggest that adaptation in macaque ganglion cells depends on mechanisms both before and after the site of cone interaction, and that these mechanisms may differ in time course between MC- and PC-cells. The results indicate that it may be important in psychophysical adaptation experiments to consider the presence of multiple postreceptoral mechanisms with different adaptation characteristics.

Adaptation, Ocular↗

Enduring stereoscopic motion aftereffects induced by prolonged adaptation.

This study investigated the effects of prolonged adaptation on the recovery of the stereoscopic motion aftereffect (adaptation induced by moving binocular disparity information). The adapting and test stimuli were stereoscopic grating patterns created from disparity, embedded in dynamic random-dot stereograms. Motion aftereffects induced by luminance stimuli were included in the study for comparison. Adaptation duration was either 1, 2, 4, 8, 16, 32 or 64 min and the duration of the ensuing aftereffect was the variable of interest. The results showed that aftereffect duration was proportional to the square root of adaptation duration for both stereoscopic and luminance stimuli; on log-log axes, the relation between aftereffect duration and adaptation duration was a power law with the slope near 0.5 in both cases. For both kinds of stimuli, there was no sign of adaptation saturation even at the longest adaptation duration.

Adaptation, Ocular↗

Mechanism of general adaptation.

An analysis of theories and results of corresponding studies indicate that the adaptive processes in organisms have to be discriminated to specific and nonspecific adaptive responses. The integrated sum of specific adaptive responses constitute homeostatic regulation in order to maintain a constant level of rigid constants of the body's internal milieu. The constancy of rigid constants (temperature, pH, osmotic pressure, and contents of ions, water and p0(2)) is necessary to ensure the optimal activity of enzymes. The nonspecific adaptive responses are directed towards the mobilization of the organism's reserves for energy and protein synthesis. Additionally, a general activation of the body's defence faculties is included into the nonspecific adaptive responses. The nonspecific adaptive responses constitute a coordinated mechanism of general adaptation. The mobilization of the reserve for protein synthesis is connected with induction of the adaptive synthesis of the enzyme and structural proteins in order to restore and develop the functional capacity of cellular structures that were highly active during acute influence of various stressors.

Adaptation, Physiological↗

Long-term adaptation in hearing impaired ears.

Recovery from adaptation was measured in acoustically-traumatized ears for pure-tone adapters 2-15 min in duration at levels of 70-110 dB SPL. The whole-nerve potential was recorded in cats. The decrease in response to a probe stimulus (normalized decrement) was used as a measure of adaptation and, for a particular adapter, was assessed at different probe frequencies. Impaired animals showed generally less effect of the adapting stimuli at a given adapter level (in dB SPL) and faster recovery than in normal ears. The form of the normalized decrement vs probe frequency functions were generally normal. We interpret these observations to mean that while the effectiveness of the adapter is reduced the spread of adaptation to different frequency regions is relatively normal.

Acoustic Stimulation↗

Odorant-specific adaptation pathways generate olfactory plasticity in C. elegans.

Following prolonged exposure to an odorant, C. elegans exhibits a diminished response to the odorant for several hours. This olfactory adaptation is odorant selective; animals can adapt independently to different odorants sensed by a single pair of olfactory neurons, the AWC neurons. The mechanism of olfactory adaptation is genetically complex, with different genes required for adaptation to different odorants. Animals mutant for the gene adp-1 fail to adapt to a subset of AWC-sensed odorants; adp-1 affects a calcium-dependent process required for adaptation. Mutations in another gene, osm-9, affect adaptation to a different but overlapping subset of AWC-sensed odorants. Mutations in adp-1 and osm-9 do not diminish the ability of unadapted animals to respond to odorants, indicating that odorant sensation and odorant adaptation are distinct processes.

Adaptation, Physiological↗

Response of the osteocyte syncytium adjacent to and distant from linear microcracks during adaptation to cyclic fatigue loading.

Cyclic loading induces fatigue in bone and initiates a complex, functionally adaptive response. We investigated the effect of a single period of fatigue on the histologic structure and biomechanical properties of bone. The ulnae of 40 rats were subjected to cyclic fatigue (-6000 microepsilon) unilaterally until 40% loss of stiffness developed, followed by 14 days of adaptation. The contralateral ulna served as a treatment control (n = 20 rats), and a baseline loaded/non-loaded group (n = 20 rats/group) was included. Bones from 10 rats/group were examined histologically and the remaining bones (10 rats/group) were tested mechanically. The following measurements were collected: volumetric bone mineral density (vBMD); ultimate force (Fu); stiffness (S); energy-to-failure (U); cortical area (Ct.Ar); microcrack density (Cr.Dn); microcrack mean length (Cr.Le); microcrack surface density (Cr.S.Dn); osteocyte density (Ot.N/T.Ar and Ot.N/TV); bone volume fraction (B.Ar/T.Ar); resorption space density (Rs.N/Ct.Ar); and maximum and minimum area moments of inertia (IMAX and IMIN). Using confocal microscopy, the bones were examined for diffuse matrix injury, canalicular disruption, and osteocyte disruption. The adapted bones had increased B.Ar, IMAX, and IMIN in the mid-diaphysis. Fatigue loading decreased structural properties and induced linear microcracking. At 14 days, adaptation restored structural properties and microcracking was partially repaired. There was a significant nonlinear relationship between Ot.N/T.Ar and B.Ar/T.Ar during adaptation. Disruption of osteocytes was observed adjacent to microcracks immediately after fatigue loading, and this did not change after the period of adaptation. In fatigue-loaded bone distant from microcracks, diffuse matrix injury and canalicular disruption were often co-localized and were increased in the lateral (tension) cortex. These changes were partially reversed after adaptation. Loss of canalicular staining and the presence of blind-ends in regions with matrix injury were suggestive of rupture of dendritic cell processes. Taken together, these data support the general hypothesis that the osteocyte syncytium can respond to cyclic loading and influence targeted remodeling during functional adaptation. Changes in the appearance of the osteocyte syncytium were found in fatigue-loaded bone with and without linear microcracks. We hypothesize that the number of dendritic cell processes that experience load-related disruption may determine osteocyte metabolic responses to loading and influence targeted remodeling.

Adaptation, Physiological↗

Information transmission rate changes of retinal ganglion cells during contrast adaptation.

During adaptation to high-contrast stimulation, retinal ganglion cell's responsiveness change is characterized by decreased firing rate and declined sensitivity. In order to examine the modification of information transmission properties of the ganglion cell during this adaptation process, neural activities were recorded extracellularly from the chicken retina using a multi-electrode recording system, and the information transmission rate of the retinal ganglion cells was estimated. The results show that the response entropy and noise entropy of the ganglion cells both decreased during the adaptation process, which resulted in a modest decline of information transmission rate of ganglion cells after several seconds' adaptation. However, due to the decrease of the neuron's firing rate during the adaptation, it is revealed that the information carried by each spike was increased as compared to pre-adaptation, suggesting that retinal ganglion cells' information processing strategies during contrast adaptation may reflect economical principle by promoting each spike more informative. These results also suggest that contrast adaptation and sensitivity rescaling of the visual neurons provide an efficient manner in information transmission and save the system's metabolic cost in the meantime.

Action Potentials↗