Publisher Summary This chapter discusses the response characteristics of neocortical graft neurons to host somatosensory input. The neocortical grafts placed in the primary somatosensory cortex of adult rats respond to host sensory inputs... more
Publisher Summary This chapter discusses the response characteristics of neocortical graft neurons to host somatosensory input. The neocortical grafts placed in the primary somatosensory cortex of adult rats respond to host sensory inputs under certain restricted conditions. The responses in these innervated transplants are substantial but far below the level of activity seen in normal neocortical neurons. The decrement in graft responses is because of 2 factors: (i) abnormal distribution and development of various fiber systems found in neocortex after normal development and (ii) a failure of innervated graft neurons to develop normal excitability because of the imbalance of extrathalamic modulatory influences. The effect of pharmacological agents on graft responses is discussed in the chapter. In an experiment described in the chapter, some of the graft neurons isolated for response measurements were held long enough to quantify the effects of transiently decreasing intra-graft inhibition with bicuculline ( 5 mM, pH 3.3, 40-90 nA) or increasing the post-synaptic excitability with iontophoresis of glutamate (0.5 M, pH 7.2, 20-65 nA). The changes seen in the response profile following iontophoresis of glutamate and bicuculline can be seen. The response probability following glutamate and bicuculline application increased to 0.42 and 0.72, respectively, without changing the latency to onset of response. Many graft cells that were unresponsive to peripheral stimulation became so following the application of these drugs. This effect is reflected in the percentage of cells isolated that were responsive to peripheral stimulation.
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In rats, navigating through an environment requires continuous information about objects near the head. Sensory information such as object location and surface texture are encoded by spike firing patterns of single neurons within rat... more
In rats, navigating through an environment requires continuous information about objects near the head. Sensory information such as object location and surface texture are encoded by spike firing patterns of single neurons within rat barrel cortex. Although there are many studies using single-unit electrophysiology, much less is known regarding the spatiotemporal pattern of activity of populations of neurons in barrel cortex in response to whisker stimulation. To examine cortical response at the population level, we used voltage-sensitive dye (VSD) imaging to examine ensemble spatiotemporal dynamics of barrel cortex in response to stimulation of single or two adjacent whiskers in urethane-anesthetized rats. Single whisker stimulation produced a poststimulus fluorescence response peak within 12–16 ms in the barrel corresponding to the stimulated whisker (principal whisker). This fluorescence subsequently propagated throughout the barrel field, spreading anisotropically preferentially...
Research Interests: Neuroscience, Materials Science, Chemistry, Electrophysiology, Neurophysiology, and 13 moreMedicine, Animals, Somatosensory Cortex, Rats, Time Factors, Barrel Cortex, Local Field Potential, Stimulation, Action Potentials, Whisker, Psychology and Cognitive Sciences, Somatosensory system, and Medical and Health Sciences
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Crabtree and colleagues report newly discovered connections between nuclei of the somatosensory thalamus, challenging the traditional view that thalamic nuclei do not communicate.
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DefinitionThe 5 region of the mRNA that is not translated into protein. It extends from the transcription start site to the translation ATG start site, and contains regulatory sequences that control mRNA stability and translation... more
DefinitionThe 5 region of the mRNA that is not translated into protein. It extends from the transcription start site to the translation ATG start site, and contains regulatory sequences that control mRNA stability and translation efficiency.
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Previous electrophysiological experiments have documented the response of neurons in the adult rat somatic sensory ("barrel") cortex to whisker movement after normal experience and after periods of experience with all but two... more
Previous electrophysiological experiments have documented the response of neurons in the adult rat somatic sensory ("barrel") cortex to whisker movement after normal experience and after periods of experience with all but two whiskers trimmed close to the face (whisker "pairing"). To better understand how the barrel cortex adapts to changes in the flow of sensory activity, we have developed a computational model of a single representative barrel cell based on the Bienenstock, Cooper, and Munro (BCM) theory of synaptic plasticity. The hallmark of the BCM theory is the dynamic synaptic modification threshold, theta M, which dictates whether a neuron's activity at any given instant will lead to strengthening or weakening of the synapses impinging on it. The threshold theta M is proportional to the neuron's activity averaged over some recent past. Whisker pairing was simulated by setting input activities of the cell to the noise level, except for two inputs t...
Research Interests: Genetics, Neuroscience, Physics, Electrophysiology, Plasticity, and 15 moreMedicine, Multidisciplinary, Synaptic Plasticity, Visual Cortex, Computer Simulation, Cerebral Cortex, Computer Model, Animals, Neuronal Plasticity, Synaptic Transmission, Neurons, Rats, Experience-Dependent Plasticity, Barrel Cortex, and Somatosensory system
Research Interests: Biology, Microcirculation, Oxidative Stress, Medicine, Biological Sciences, and 14 moreAnatomy, Female, Animals, Male, Parietal Cortex, Rats, Single Photon Emission Computed Tomography, Occlusion, Blood Flow, Model System, Red blood cell, Middle Cerebral Artery, Vascular Diseases, and Medical and Health Sciences
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Page 1. Two-photon microscopy reveals that cortical blood flow reverses direction at the first branch that lies downstream from a localized photothrombotic clot Chris B. Schaffera, Nozomi Nishimuraa, Beth Friedmanb, Philbert ...
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Histochemical localization of acetylcholine esterase (AChE) shows that the reaction product is in the outer half of the molecular layer of dorsal cortex of the turtle Pseudemys. Thalamic and noradrenergic locus coeruleus fibers are found... more
Histochemical localization of acetylcholine esterase (AChE) shows that the reaction product is in the outer half of the molecular layer of dorsal cortex of the turtle Pseudemys. Thalamic and noradrenergic locus coeruleus fibers are found in the same location. Two hypotheses could account for this apparent overlap of inputs. First, a cholinergic fiber system could exist in turtle cortex that occupies the same portion of the molecular layer. On the other hand, the AChE enzyme could be associated with a non-cholinergic fiber system, for example the adrenergic fibers. In the latter alternative nlo cholinergic fiber input would need to be present in turtle cortex at all. Our experiments analyzed the levels of acetylcholine (ACh) and the activity of its synthetic enzyme, choline acetyltransferase (ChAT) in adult turtle thalamic input to cortex as a first step toward distinguishing between these alternatives. The results show that turtle cortex contains ACh and exhibits ChAT activity. These biochemical results support the idea that the AChE staining pattern in the outer half of the molecular layer may reflect the laminar distribution of cholinergic fiber activity in this simple cortex.
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We model experience-dependent plasticity in the cortical representation of whiskers (the barrel cortex) in normal adult rats, and in adult rats that were prenatally exposed to alcohol. Prenatal exposure to alcohol (PAE) caused marked... more
We model experience-dependent plasticity in the cortical representation of whiskers (the barrel cortex) in normal adult rats, and in adult rats that were prenatally exposed to alcohol. Prenatal exposure to alcohol (PAE) caused marked deficits in experience-dependent plasticity in a cortical barrel-column. Cortical plasticity was induced by trimming all whiskers on one side of the face except two. This manipulation produces high activity from the intact whiskers that contrasts with low activity from the cut whiskers while avoiding any nerve damage. By a computational model, we show that the evolution of neuronal responses in a single barrel-column after this sensory bias is consistent with the synaptic modifications that follow the rules of the Bienenstock, Cooper, and Munro (BCM) theory. The BCM theory postulates that a neuron possesses a moving synaptic modification threshold, θ M , that dictates whether the neuron's activity at any given instant will lead to strengthening or w...
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During tactile learning there is a transformation in the way the primary somatosensory cortex integrates, represents, and distributes information from the skin. To define this transformation, the site of earliest modification has been... more
During tactile learning there is a transformation in the way the primary somatosensory cortex integrates, represents, and distributes information from the skin. To define this transformation, the site of earliest modification has been identified in rat somatosensory cortex after a change in sensory experience. Afferent activity was manipulated by clipping all except two whiskers on one side of the snout ("whisker pairing"), and the receptive fields of neurons at different cortical depths were mapped 24 hours later. Neurons in layer IV, the target of the primary thalamic pathway, were unaltered, whereas neurons located above and below layer IV showed significant changes. These changes were similar to those that occur in layer IV after longer periods of whisker pairing. The findings support the hypothesis that the layers of cortex contribute differently to plasticity. Neurons in the supragranular and infragranular layers respond rapidly to changes in sensory experience and may contribute to subsequent modification in layer IV.
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Trimming all whiskers except two on one side of an adult rat's face results in cortical plasticity in which the spared whiskers, D2 and one D-row surround whisker (either D1 or D3), evoked responses containing more spikes than the... more
Trimming all whiskers except two on one side of an adult rat's face results in cortical plasticity in which the spared whiskers, D2 and one D-row surround whisker (either D1 or D3), evoked responses containing more spikes than the response evoked by the cut whisker (called whisker pairing plasticity). Previously we have reported that acetylcholine (ACh) depletion in cortex prevents surround D-row whisker plasticity from developing within the barrel cortex. In this study we examined whether the animal's active use of its two intact whiskers can restore some aspects of plasticity in the ACh-depleted cortex. To achieve this goal, ACh was depleted from barrel field cortex, and 14 days after the depletion surgery, whiskers were trimmed and animals were trained on a whisker-dependent gap crossing task. After 7 days of training, animals were anesthetized with urethan and prepared for single-unit recording. Training the ACh-depleted, whisker-paired animals resulted in a significant ...
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Trimming all but two whiskers in adult rats produces a predictable change in cortical cell-evoked responses characterized by increased responsiveness to the two intact whiskers and decreased responsiveness to the trimmed whiskers. This... more
Trimming all but two whiskers in adult rats produces a predictable change in cortical cell-evoked responses characterized by increased responsiveness to the two intact whiskers and decreased responsiveness to the trimmed whiskers. This type of synaptic plasticity in rat somatic sensory cortex, called "whisker pairing plasticity," first appears in cells above and below the layer IV barrels. These are also the cortical layers that receive the densest cholinergic inputs from the nucleus basalis. The present study assesses whether the cholinergic inputs to cortex have a role in regulating whisker pairing plasticity. To do this, cholinergic basal forebrain fibers were eliminated using an immunotoxin specific for these fibers. A monoclonal antibody to the low-affinity nerve growth factor receptor 192 IgG, conjugated to the cytotoxin saporin, was injected into cortex to eliminate cholinergic fibers in the barrel field. The immunotoxin reduces acetylcholine esterase (AChE)-positiv...
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The current view of whisker movement is that approximately 25 whiskers on each side of the face move in synchrony. To determine whether whiskers are constrained to move together, we trained rats to use two whiskers on the same side of the... more
The current view of whisker movement is that approximately 25 whiskers on each side of the face move in synchrony. To determine whether whiskers are constrained to move together, we trained rats to use two whiskers on the same side of the face in simple behavioral tasks and videotaped the whiskers during the task. Here we report that the movement of adjacent whiskers is usually synchronous but can diverge: 1) the distance between whiskers can vary dramatically during movement; 2) one whisker can move while the second one remains stationary; 3) two whiskers can simultaneously move in opposite directions; and 4) one whisker can be maintained in contact with an object while the other is retracted and protracted. The frequency of whisker movement during the task falls within the previously reported range for rats whisking freely into air or performing roughness discrimination with their whiskers. Our data also suggest that whisker movement can be divided into three distinct phases: prot...
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Whisker deflection typically evokes a transient volley of action potentials in rat somatic sensory (SI) barrel cortex. Postexcitatory inhibition is thought to quickly terminate the cortical cell response to whisker deflection. Using dual... more
Whisker deflection typically evokes a transient volley of action potentials in rat somatic sensory (SI) barrel cortex. Postexcitatory inhibition is thought to quickly terminate the cortical cell response to whisker deflection. Using dual electrode extracellular recording in awake rats, we describe an infrequent type of cell response in which stimulation of single hairs consistently blocks the ongoing discharge of neurons without prior excitation (I-only inhibition). Reconstruction of the recording sites indicates that I-only inhibition occurs most frequently when the recording site is clearly in the septum or at the barrel-septum junction. The same cells that respond with I-only inhibition to one whisker can show an excitatory discharge to other whiskers, usually followed by inhibition. Stimulation of either nose hairs or the large mystacial vibrissa can evoke I-only inhibition in SI cortex. I-only inhibition is most commonly observed at low stimulus frequencies ( approximately 1 Hz...
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Modulation of receptive field properties of thalamic somatosensory neurons by the depth of anesthesia. The dominant frequency of electrocorticographic (ECoG) recordings was used to determine the depth of halothane or urethan anesthesia... more
Modulation of receptive field properties of thalamic somatosensory neurons by the depth of anesthesia. The dominant frequency of electrocorticographic (ECoG) recordings was used to determine the depth of halothane or urethan anesthesia while recording extracellular single-unit responses from thalamic ventral posterior medial (VPM) neurons. A piezoelectric stimulator was used to deflect individual whiskers to assess the peak onset latency, magnitude, probability of response, and receptive field (RF) size. There was a predictable increase in the dominant ECoG frequency from deep stage IV to light stage III-1 anesthetic levels. There was no detectable frequency at stage IV, a 1- to 2-Hz dominant frequency at stage III-4, 3-4 Hz at stage III-3, 5-7 Hz at stage III-2, and a dual 6- and 10- to 13-Hz pattern at stage III-1. Reflexes and other physical signs showed a correlation with depth of anesthesia but exhibited too much overlap between stages to be used as a criterion for any single s...
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A short period of combined deficiency of vitamins E and C causes profound central nervous system (CNS) dysfunction in guinea pigs. For this report, CNS histopathology was studied to define the nature and extent of injury caused by this... more
A short period of combined deficiency of vitamins E and C causes profound central nervous system (CNS) dysfunction in guinea pigs. For this report, CNS histopathology was studied to define the nature and extent of injury caused by this double deficiency. Weanling guinea pigs were fed a vitamin E-deficient or -replete diet for 14 d. Then vitamin C was withdrawn from the diet of some guinea pigs. Four diet groups were thus formed: replete, vitamin E deficient, vitamin C deficient, and both vitamin E and C deficient. From 5 to 11 d after institution of the doubly deficient diet, 9 of 12 guinea pigs developed paralysis, and 2 more were found dead. The remaining guinea pig in the doubly deficient group and all animals in the other 3 groups survived without clinical impairment until the experiment was terminated at 13-15 d. Brains and spinal cords were serially sectioned and stained for examination. Only the combined deficiency produced damage in the CNS. The damage consisted mainly of ne...
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1. Changes in the receptive field (RF) properties of thalamic VPM neurons were assessed quantitatively using single-unit recording techniques following a selective excitotoxic lesion of the ipsilateral thalamic reticular nucleus (TRN).... more
1. Changes in the receptive field (RF) properties of thalamic VPM neurons were assessed quantitatively using single-unit recording techniques following a selective excitotoxic lesion of the ipsilateral thalamic reticular nucleus (TRN). The response profiles to controlled deflections of the contralateral vibrissae were obtained from 97 VPM neurons in normal and 102 VPM neurons in TRN-lesioned animals. 2. Histological signs of TRN lesions were detectable in Nissl-stained sections as early as 20 h after the release of kainic acid into TRN. 3. The average RF size of VPM neurons in normal animals was 2.39 +/- 0.18 whiskers (mean +/- SE). Immediately after the lesion of TRN, the average RF size in VPM was enlarged significantly and remained expanded for as long as 1 mo after the destruction of TRN (7.64 +/- 0.47 whiskers, P < 0.001). 4. Subsequent lesions of trigeminal subnucleus interpolaris (SpVi) in TRN-lesioned animals produced a marked reduction in the RF size of VPM neurons. The ...
Research Interests: Chemistry, Neurophysiology, Medicine, Brain Mapping, Female, and 15 moreAnimals, Male, Reaction Time, Mechanoreceptors, Receptive Field, Synaptic Transmission, Neurons, Baclofen, Rats, Thalamic Reticular Nucleus, Neural pathways, Bicuculline, Psychology and Cognitive Sciences, Medical and Health Sciences, and Nerve degeneration
It is known that sensory deprivation, including postnatal whisker trimming, can lead to severe deficits in the firing rate properties of cortical neurons. Recent results indicate that development of synchronous discharge among cortical... more
It is known that sensory deprivation, including postnatal whisker trimming, can lead to severe deficits in the firing rate properties of cortical neurons. Recent results indicate that development of synchronous discharge among cortical neurons is also activity influenced, and that correlated discharge is significantly impaired following loss of bilateral sensory input in rats. Here we investigate whether unilateral whisker trimming (unilateral deprivation or UD) after birth interferes in the same way with the development of synchronous discharge in cortex. We measured the coincidence of spikes among pairs of neurons recorded under urethane anesthesia in one whisker barrel field deprived by trimming all contralateral whiskers for 60 days after birth (UD), and in untrimmed controls (CON). In the septal columns around barrels, UD significantly increased the coincident discharge among cortical neurons compared with CON, most notably in layers II/III. In contrast, synchronous discharge w...
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Contribution of supragranular layers to sensory processing and plasticity in adult rat barrel cortex. J. Neurophysiol. 80: 3261-3271, 1998. In mature rat primary somatic sensory cortical area (SI) barrel field cortex, the... more
Contribution of supragranular layers to sensory processing and plasticity in adult rat barrel cortex. J. Neurophysiol. 80: 3261-3271, 1998. In mature rat primary somatic sensory cortical area (SI) barrel field cortex, the thalamic-recipient granular layer IV neurons project especially densely to layers I, II, III, and IV. A prior study showed that cells in the supragranular layers are the fastest to change their response properties to novel changes in sensory inputs. Here we examine the effect of removing supragranular circuitry on the responsiveness and synaptic plasticity of cells in the remaining layers. To remove the layer II + III (supragranular) neurons from the circuitry of barrel field cortex, N-methyl--aspartate (NMDA) was applied to the exposed dura over the barrel cortex, which destroys those neurons by excitotoxicity without detectable damage to blood vessels or axons of passage. Fifteen days after NMDA treatment, the first responsive cells encountered were 400-430 micro...
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The daily ingestion of alcohol by pregnant mammals exposes the fetal brain to varying levels of alcohol through the placental circulation. Here we focus on the lingering impact on cortical function of 6.5% alcohol administered in a liquid... more
The daily ingestion of alcohol by pregnant mammals exposes the fetal brain to varying levels of alcohol through the placental circulation. Here we focus on the lingering impact on cortical function of 6.5% alcohol administered in a liquid diet to pregnant rats throughout gestation, followed by 3 alcohol-free months before brain function was analyzed in the offspring. Both spontaneous activity of the neurons in the barrel cortex and the level of response to test stimuli applied to the whiskers remained reduced by >75% after alcohol exposure. Whisker pairing, a type of cortical plasticity induced by trimming all but two whiskers in adult rats, occurred in <1 d in controls, but required 14 d to reach significance after alcohol exposure. These long-term neuronal deficits are present in all layers of cortex and affect neurons with both fast and slow action potentials. Plasticity is first seen in the total sample of neurons at 14 d; however, by 7 d, neurons in layer II/III already s...
Research Interests: Neuroscience, Aging, Animal Husbandry, Pregnancy, Cerebral Cortex, and 15 moreFemale, Animals, Ethanol, Neuronal Plasticity, Neurons, Cortical Plasticity, Prenatal Alcohol Exposure, Barrel Cortex, Action Potentials, Brain Function, Environment, Cortical Reorganization, Cortical Excitability, NMDA receptor, and Medical and Health Sciences
The capacity of adult barrel cortex to show experience-dependent plasticity after early restricted neonatal sensory deprivation was analyzed in barrel field cortex neurons. Selective sensory deprivation was induced by trimming two... more
The capacity of adult barrel cortex to show experience-dependent plasticity after early restricted neonatal sensory deprivation was analyzed in barrel field cortex neurons. Selective sensory deprivation was induced by trimming two whiskers from postnatal day 0 (P0) to P21, namely, the principal D2 whisker plus one adjacent surround whisker (D3). At maturity (P90), responses of supragranular (layer II/III) and barrel (layer IV) neurons, all located in the D2 barrel column, were analyzed for modified responses to the deprived principal whisker (D2) and the nondeprived (D1) and deprived (D3) adjacent surround whiskers. For supragranular neurons, the responses to both principal and surround whiskers were reduced at maturity, whereas the barrel neurons showed mildly elevated responses to the principal whisker but a reduced response to the deprived surround whisker. In normal adult rats, trimming all but the principal D2 whisker and an adjacent D3 whisker for 3 d (whisker pairing) produce...
Research Interests: Chemistry, Medicine, Learning, Animals, Male, and 13 moreNeuronal Plasticity, Receptive Field, Synaptic Transmission, Neurons, Cortical Development, Neural plasticity, Somatosensory Cortex, Rats, Sensory Deprivation, Experience-Dependent Plasticity, Barrel Cortex, Psychology and Cognitive Sciences, and Medical and Health Sciences
Lesions of primary sensory cortex produce impairments in brain function as an outcome of the direct tissue damage. In addition, indirect lesion effects have been described that consist of functional deficits in areas sharing neural... more
Lesions of primary sensory cortex produce impairments in brain function as an outcome of the direct tissue damage. In addition, indirect lesion effects have been described that consist of functional deficits in areas sharing neural connections with the damaged area. The present study characterizes interhemispheric deficits produced as a result of unilateral lesions of the entire vibrissa representation of S-I barrel field cortex (BFC) in adult rats using single-neuron recording under urethane anesthesia. After unilateral lesions of adult BFC, responses of neurons in the contralateral homotopic BFC are severely depressed. Background (spontaneous) activity is reduced by approximately 80%, responses to test stimuli applied to the whiskers are reduced by approximately 50%, and onset of synaptic plasticity induced by trimming all but two whiskers ("whisker-pairing plasticity") is delayed over sevenfold compared with sham-lesion control animals. These deficits persist with only ...
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The daily ingestion of alcohol by pregnant mammals exposes the fetal brain to varying levels of alcohol through the placental circulation. Here we focus on the lingering impact on cortical function of 6.5% alcohol administered in a liquid... more
The daily ingestion of alcohol by pregnant mammals exposes the fetal brain to varying levels of alcohol through the placental circulation. Here we focus on the lingering impact on cortical function of 6.5% alcohol administered in a liquid diet to preg- nant rats throughout gestation, followed by 3 alcohol-free months before brain function was analyzed in the offspring. Both spontaneous
Research Interests: Neuroscience, Aging, Animal Husbandry, Pregnancy, Cerebral Cortex, and 15 moreFemale, Animals, Ethanol, Neuronal Plasticity, Neurons, Cortical Plasticity, Prenatal Alcohol Exposure, Barrel Cortex, Action Potentials, Brain Function, Environment, Cortical Reorganization, Cortical Excitability, NMDA receptor, and Medical and Health Sciences
Brain tissue transplantation is a versatile technique with the potential to identify cellular mechanisms that control developing as well as mature functions of neurons and glia. We are currently interested in how the continued... more
Brain tissue transplantation is a versatile technique with the potential to identify cellular mechanisms that control developing as well as mature functions of neurons and glia. We are currently interested in how the continued differentiation of embryonic donor tissue affects the ingrowth of adult host axons into these transplants. Two important variables are the type of host fiber and the age of the donor tissue at the time of transplantation. For example, axons labeled by acetylcholinesterase (AChE) histochemistry always start to grow into the transplants within a few days and ultimately achieve numerical densities comparable to the normal cortex (Hohmann and Ebner 1982, Park, et al. 1984). Monoamine fibers visualized by histofluorescence, in contrast, are not detectable for 3–4 weeks and never reach normal densities in these neocortical transplants (Park et al. 1984). Only rare thalamic and commissural fibers elongate across the interface zone into the transplants without special pretreatment (Smith et al. 1984). Host axons arising from peptidergic neurons have never been observed entering the transplants at any donor age (Ebner et al. 1984). Host GABAergic neurons grow into transplants taken from very young donors, embryonic day 14 or younger, but not at all into older donor tissue (Smith et al. 1984). All types of host fiber systems are damaged to some extent at the time of transplantation, but these results indicate that each responds to the damage in very different ways. The precise molecular composition of the donor tissue may be crucial to successful transplant innervation. For example, cholinergic axons that initiate an immediate response to damage encounter a set of cell surface and matrix molecules that are different from those encountered by slower growing fibers because the embryonic donor tissue is differentiating rapidly after implantation. The type of connections that develop between donor and host tissue therefore may depend upon synchrony of host axon growth with the degree of maturity of transplant cells.
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The capacity of adult barrel cortex to show experience-dependent plasticity after early restricted neonatal sensory deprivation was analyzed in barrel field cortex neurons. Selective sensory deprivation was induced by trimming two... more
The capacity of adult barrel cortex to show experience-dependent plasticity after early restricted neonatal sensory deprivation was analyzed in barrel field cortex neurons. Selective sensory deprivation was induced by trimming two whiskers from postnatal day 0 (P0) to P21, namely, the principal D2 whisker plus one adjacent surround whisker (D3). At maturity (P90), responses of supragranular (layer II/III) and barrel