Effects of sensory deprivation on the development of asymmetrical synapses in mouse barrels

Alterations in the numerical density and structure of asymmetrical synapses were examined in thin sections through barrel D4 in six CD/1 mice, including three controls and three sensory deprived animals. Sensory deprivation was effected by once daily trimming of all large mystacial vibrissae on the contralateral side of the snout from P0. The mice were perfuse-fixed at P20, several days following the termination of rapid synaptic growth during barrel development (White et al. , Somatosens Mot Res 14 : 34-55, 1997). Cerebral hemispheres contralateral to the deprived side were osmicated, sectioned at 40 mum and embedded in plastic for thin sectioning. Sterio's ( J Microsc 134 : 127-136, 1984) procedure combined with serial thin section analysis (Braendgaard and Gundersen, J Neurosci Meth 18 : 39-78, 1986), was applied blindly to systematic random samples of neuropil in barrel hollows and septa. No significant difference in the numerical density, estimated total number, or in the proportion of perforated postsynaptic densities was observed. However, a significant decrease in the diameters of asymmetrical postsynaptic densities was observed in hollow (P < 0.05) and septal (P < 0.05) neuropil of deprived animals. These results demonstrate a significant morphological alteration in asymmetrical synapses of a type consistent with a reduction in synaptic activity consequent to sensory deprivation.     

Sustained Enhancement of Lateral Inhibitory Circuit Maintains Cross Modal Cortical Reorganization

Deprivation of one modality can lead to the improvement of other intact modalities. We have previously reported that visual deprivation drives AMPA receptors into synapses from layer4 to 2/3 in the barrel cortex and sharpens functional whisker-barrel map at layer2/3 2 days after the beginning of visual deprivation. Enhanced excitatory synaptic transmission at layer4-2/3 synapses is transient and returns to the base line level a week after the beginning of visual deprivation. Here we found that sharpened whisker-barrel function is maintained at least for a week in visually deprived animals. While increased AMPA receptor-mediated synaptic transmission at layer4-2/3 synapses dropped to the base line a week after the beginning of visual deprivation, lateral inhibitory synaptic transmission onto the neighboring barrel was kept strengthened for a week of visually deprived animals. Thus, transient strengthening of excitatory synapses at layer4-2/3 in the barrel cortex could trigger the enhancement of inhibitory inputs to neighboring barrel, and sustained lateral inhibition can maintain the sharpening of whisker-barrel map in visually deprived animals.     

The effect of vibrissa deprivation pattern on the form of plasticity induced in rat barrel cortex

Plasticity was induced in the barrel cortex of adolescent rats by depriving every second vibrissa on the contralateral vibrissa pad.This produced a chessboard pattern of barrels in the cortex where each barrel receiving its principal input from a spared vibrissa was surrounded by barrels for which the principal vibrissa had been deprived and conversely, each barrel receiving its principal input from a deprived vibrissa was surrounded by barrels for which the principal vibrissa had been spared. After 7 days' deprivation, responses to the regrown vibrissae were depressed in layers II/III (49% of control levels) and IV (60%). Depression was far greater than that seen with "all vibrissa" deprivation, suggesting that activity in the spared vibrissae accentuated the depression of the deprived vibrissae. Depression was not due to subcortical changes as thalamic Ventral Posterior Medial (VPM) responses to deprived vibrissa were unchanged. The short latency responses in layer IV (5-7 ms) were unaffected by deprivation, but the number of cells responding at intermediate latencies (8-13 ms) was markedly reduced (to 66% of control). Potentiation of the spared vibrissa response was substantial in the near side of the neighbouring barrel (2.2-fold increase in layers II/III, 2.9-fold in layer IV) but had not spread to the far side after 7 days' deprivation. Sparing multiple vibrissae may increase the rate of potentiation since 7 days is insufficient time for potentiation in single vibrissa spared animals. Potentiation was not due to subcortical changes as thalamic VPm responses to the spared vibrissa were normal. However, in the spared barrel the response latency decreased by 1-2 ms. Only the cells responding at short latency exhibited potentiated responses (39% increase) suggesting that some thalamocortical plasticity is still possible at P28-35. These results show that chessboard pattern deprivation is capable of inducing substantial plasticity over a wide area of barrel cortex. All the major forms of plasticity seen with other vibrissa deprivation patterns were present, although no other single deprivation pattern studied so far causes the complete repertoire seen with chessboard deprivation.     

Rapid development and plasticity of layer 2/3 maps in rat barrel cortex in vivo

Cortical synaptic circuitry develops rapidly in the second postnatal week, simultaneous with experience-dependent turnover of dendritic spines. To relate the emergence of sensory maps to synaptogenesis, we recorded synaptic potentials evoked by whisker deflection in layer 2/3 neurons from postnatal day (P) 12 to 20. At P12, synaptic responses were undetectable. Only 2 days later in life (P14), receptive fields had mature organization. Sensory deprivation, if initiated before P14, disrupted receptive field structure. In layer 4, responses and maps were already mature by P12 and insensitive to deprivation, implying that barrel cortex develops from layer 4 to layer 2/3. Thus, P12-14 is a critical period shared by layer 2/3 synapses and their spines, suggesting that spine plasticity is involved in the refinement of maps.     

Serotonin mediates cross-modal reorganization of cortical circuits

Loss of one type of sensory input can cause improved functionality of other sensory systems. Whereas this form of plasticity, cross-modal plasticity, is well established, the molecular and cellular mechanisms underlying it are still unclear. Here, we show that visual deprivation (VD) increases extracellular serotonin in the juvenile rat barrel cortex. This increase in serotonin levels facilitates synaptic strengthening at layer 4 to layer 2/3 synapses within the barrel cortex. Upon VD, whisker experience leads to trafficking of the AMPA-type glutamate receptors (AMPARs) into these synapses through the activation of ERK and increased phosphorylation of AMPAR subunit GluR1 at the juvenile age when natural whisker experience no longer induces synaptic GluR1 delivery. VD thereby leads to sharpening of the functional whisker-barrel map at layer 2/3. Thus, sensory deprivation of one modality leads to serotonin release in remaining modalities, facilitates GluR1-dependent synaptic strengthening, and refines cortical organization.     

A survey of morphogenesis during the early postnatal period in PMBSF barrels of mouse SmI cortex with emphasis on barrel D4

The long term goal of this work is to understand synaptogenesis in homologous regions of the cerebral cortex, i.e. a whisker barrel. Hemispheres of aldehyde perfused mice, at various ages from P6 to P65 (DOB = P0; three each), were osmicated and sectioned at 40mm parallel to the pia. Barrels were identified, mapped and measured in sections through mid-level layer IV, and then embedded for electron microscopy. The main findings were: (1) Cell bodies and large diameter dendrites thin out in barrel hollows from P6 to P8. (2) Degeneration occurs primarily from P6 to P11, peaking on P8. (3) Single synapses from narrow, tubular axons predominate before P14; afterwards, multiple synapses from bag-like terminals increase in number. (4) The number of spines increases dramatically between P9 and P12. (5) Asymmetrical and symmetrical synapses occur at all ages studied; their junction lengths are not significantly different at any age. (6) Asymmetrical synapse density increases rapidly from P6 to P8, slowly from P9 to P 12, sharply between P13 and P14 along with patterned whisking, slowly to P20 and drops in adults. (7) Synapses onto spiny and non-spiny stellate cell bodies increase markedly from P10 to P20. (8) Changes in density of asymmetrical synapses in neuropil and of symmetrical synapses on spiny stellate cell bodies follow similar sequences but the sequence in neuropil is 72 h earlier. (9) When barrel size is taken into account, synaptogenesis is monotonic, increasing sharply in the second postnatal week followed by a slower increase into adulthood.     

Development of the axon cap neuropil of the Mauthner cell in the goldfish

Summary Development of the axon cap neuropil of the Mauthner neuron in post-hatching larval goldfish brains was observed electron-microscopically. The axonal initial segment of newly hatched (day-4) larvae is completely covered with synaptic terminals containing clear spherical synaptic vesicles. Profiles of thin terminal axons, the spiral fibers, containing similar synaptic vesicles, rapidly increase in number around the initial segment and form glomerular neuropil similar to the central core of the adult axon cap by day 7. Three types of synapses are formed in the core neuropil. Bouton-type synapses contacting the initial segment are most abundant in day-4 to-14 larvae; they decrease thereafter and are rare on the distal half of the initial segment of day-40 larvae. Asymmetric axo-axonic synapses are commonly observed between spiral fibers in the core neuropil of day-7 to -19 larvae, but become fewer by day 40. Unique symmetrical axo-axonic synapses showing accumulation of synaptic vesicles on either side of apposed membrane thickenings first appear in day-14 core neuropil, gradually increase in number, and become the predominant type in day-40 core neuropil. Thick myelinated axons, which lose their myelin sheaths in the glial cap cell layer, start to penetrate into the axon cap on day 10. They gradually increase in number and form the peripheral part of the axon cap together with the cap dendrites, which finally grow into the axon cap from the axon hillock region of the Mauthner cell by day 40.     

Organization of myelin in the mouse somatosensory barrel cortex and the effects of sensory deprivation

In rodents, the barrel cortex is a specialized area within the somatosensory cortex that processes signals from the mystacial whiskers. We investigated the normal development of myelination in the barrel cortex of mice, as well as the effects of sensory deprivation on this pattern. Deprivation was achieved by trimming the whiskers on one side of the face every other day from birth. In control mice, myelin was not present until postnatal day 14 and did not show prominence until postnatal day 30; adult levels of myelination were reached by the end of the second postnatal month. Unbiased stereology was used to estimate axon density in the interbarrel septal region and barrel walls as well as the barrel centers. Myelin was significantly more concentrated in the interbarrel septa/barrel walls than in the barrel centers in both control and sensory‐deprived conditions. Sensory deprivation did not impact the onset of myelination but resulted in a significant decrease in myelinated axons in the barrel region and decreased the amount of myelin ensheathing each axon. Visualization of the oligodendrocyte nuclear marker Olig2 revealed a similar pattern of myelin as seen using histochemistry, but with no significant changes in Olig2+ nuclei following sensory deprivation. Consistent with the anatomical results showing less myelination, local field potentials revealed slower rise times following trimming. Our results suggest that myelination develops relatively late and can be influenced by sensory experience. © 2012 Wiley Periodicals, Inc. Develop Neurobiol, 2013     

A survey of the cytology and synaptic organization of the insular trigeminal-cuneatus lateralis nucleus in the rat, including an identification of spinal afferent inputs

The cytology and synaptic organization of the insular trigeminal-cuneatus lateralis (iV-Cul) nucleus was examined in the rat. In addition, the ultrastructural morphology and synaptic connectivity of anterogradely labeled spinal afferent axons terminating in iV-Cul were examined following injection of horseradish peroxidase (HRP) into the cervical spinal cord. The uniformity of the ultrastructural features of iV-Cul neurons supports the presence of a homogeneous neuronal population. The most prominent feature of the iV-Cul neuropil is the presence of numerous interdigitating astrocytic processes, which extensively isolate neuronal somata and processes. iV-Cul contains a heterogeneous population of axonal endings that can be separated into three categories, depending upon whether they contain predominantly spherical-shaped agranular synaptic vesicles (R endings), predominantly pleomorphic-shaped agranular synaptic vesicles (P endings), or a heterogeneous population of dense-core vesicles (DC endings). The R endings represent the majority of axonal endings in iV-Cul and establish asymmetrical axodendritic and axospinous synaptic contacts, primarily along the distal portions of the dendritic tree. P endings establish symmetrical axosomatic, axodendritic, and axospinous synaptic contacts and exhibit a more generalized distribution along the somadendritic tree. DC terminals establish asymmetrical axodendritic synaptic contacts with distal dendritic processes and are the least frequently observed endings in the iV-Cul neuropil. Numerous synaptic glomeruli, exhibiting a single large central R bouton that establishes multiple axodendritic or axospinous synapses, characterize the iV-Cul neuropil. Axoaxonic synapses are conspicuously absent from the iV-Cul neuropil and glomeruli. The anterograde HRP labeling of spinal afferent axons that terminate in iV-Cul indicates that the terminals along these fibers are of the R type and that they are engaged predominantly in synaptic glomeruli. The results of this study indicate that the synaptic organization of iV-Cul is distinctly different from that of neighboring somatosensory nuclei, and supports the recent suggestion that this nucleus should be considered a separate precerebellar spinal relay nucleus in the lateral medulla.     

A Survey of the Cytology and Synaptic Organization of the Insular Trigeminal—Cuneatus Lateralis Nucleus in the Rat,Including an Identification of Spinal Afferent Inputs

The cytology and synaptic organization of the insular trigeminal—cuneatus lateralis (iV-Cul) nucleus was examined in the rat. In addition, the ultrastructural morphology and synaptic connectivity of anterogradely labeled spinal afferent axons terminating in iV-Cul were examined following injection of horseradish peroxidase (HRP) into the cervical spinal cord. The uniformity of the ultrastructural features of iV-Cul neurons supports the presence of a homogeneous neuronal population. The most prominent feature of the iV-Cul neuropil is the presence of numerous interdigitating astrocytic processes, which extensively isolate neuronal somata and processes. iV-Cul contains a heterogeneous population of axonal endings that can be separated into three categories, depending upon whether they contain predominantly spherical-shaped agranular synaptic vesicles (R endings), predominantly pleomorphic-shaped agranular synaptic vesicles (P endings), or a heterogeneous population of dense-core vesicles (DC endings). The R endings represent the majority of axonal endings in iV-Cul and establish asymmetrical axodendritic and axospinous synaptic contacts, primarily along the distal portions of the dendritic tree. P endings establish symmetrical axosomatic, axodendritic, and axospinous synaptic contacts and exhibit a more generalized distribution along the somadendritic tree. DC terminals establish asymmetrical axodendritic synaptic contacts with distal dendritic processes and are the least frequently observed endings in the iV-Cul neuropil. Numerous synaptic glomeruli, exhibiting a single large central R bouton that establishes multiple axodendritic or axospinous synapses, characterize the iV-Cul neuropil. Axoaxonic synapses are conspicuously absent from the iV-Cul neuropil and glomeruli. The anterograde HRP labeling of spinal afferent axons that terminate in iV-Cul indicates that the terminals along these fibers are of the R type and that they are engaged predominantly in synaptic glomeruli. The results of this study indicate that the synaptic organization of iV-Cul is distinctly different from that of neighboring somatosensory nuclei, and supports the recent suggestion that this nucleus should be considered a separate precerebellar spinal relay nucleus in the lateral medulla.     

Axonal Dynamics of Excitatory and Inhibitory Neurons in Somatosensory Cortex

Cortical topography can be remapped as a consequence of sensory deprivation, suggesting that cortical circuits are continually modified by experience. To see the effect of altered sensory experience on specific components of cortical circuits, we imaged neurons, labeled with a genetically modified adeno-associated virus, in the intact mouse somatosensory cortex before and after whisker plucking. Following whisker plucking we observed massive and rapid reorganization of the axons of both excitatory and inhibitory neurons, accompanied by a transient increase in bouton density. For horizontally projecting axons of excitatory neurons there was a net increase in axonal projections from the non-deprived whisker barrel columns into the deprived barrel columns. The axon collaterals of inhibitory neurons located in the deprived whisker barrel columns retracted in the vicinity of their somata and sprouted long-range projections beyond their normal reach towards the non-deprived whisker barrel columns. These results suggest that alterations in the balance of excitation and inhibition in deprived and non-deprived barrel columns underlie the topographic remapping associated with sensory deprivation.     

THE DISTRIBUTION OF SYNAPSES ON A PHYSIOLOGICALLY IDENTIFIED MOTOR NEURON IN THE CENTRAL NERVOUS SYSTEM OF THE LEECH : An Electron Microscope Study after the Injection of the Fluorescent Dye Procion Yellow

The fine structure of a physiologically identified motor neuron in the segmental ganglion of the leech central nervous system and the morphology of synapses on it were studied after injection of the fluorescent dye Procion yellow as a marker. The injected cell and its processes within the neuropil were located in thick or thin sections with fluorescence optics after initial fixation with glutaraldehyde and brief treatment with osmium tetroxide. The same or adjacent thin sections could then be examined in the electron microscope. Comparison with uninjected cells showed that the general features of the injected cell are retained although some organelles are distorted. The main features of the geometry of this neuron are the same from animal to animal: a single large process runs from the soma through the neuropil to bifurcate and enter the contralateral roots. Within the neuropil the main process gives off long branches (up to 150 µ), but these are greatly outnumbered by short branches and spines, one or a few microns in length, which were not appreciated in previous light microscope studies after injection of Procion yellow. Serial thin sections of selected areas along the main process within the neuropil showed that there are synapses on most of the shorter branches and spines; occasional synaptic contacts were also made on the main process itself and on longer branches. At least two morphologically distinct types of synapse could be recognized. A minimum estimate of the total number of synapses on the motor cell is 300, based on their occurrence in reconstructed segments.     

The cerebral ganglia of Milnesium tardigradum Doyère (Apochela, Tardigrada): Three dimensional reconstruction and notes on their ultrastructure

Differential interference contrast micrographs from stretched animals, serially sectioned semi-thin and ultrathin sections revealed that the cerebral ganglia (supraoesophageal mass) of the eulardigrade Milnesium tardigradum lie above the buccal tube and adjacent tissue like a saddle. It has an anterior indentation which is penetrated by two muscles that arise from the cuticle of the forehead. The cerebral ganglia consist of lateral outer lobes bearing an eye on each side, and two inner lobes which extend caudally. Between the inner lobes a cone-like projection tapers into a nerve bundle. Each outer lobe is joined with the first ventral ganglion. From the outer lobe near the eye the ganglion for a posterolateral sensory field extends to the epidermis. Anterior to the supraoesophageal mass are three dorsal ganglia for the upper three peribuccal papillae. Two additional ganglia attached to the cerebral mass supply the lateral cephalic papillae. The cerebral ganglia are covered by a thin neural lamella. The pericarya which surround the neuropil have large nuclei. Near the axons in the centre of the supraoesophageal mass the cytoplasm is crowded with vesicles of different size and appearance. Some of them resemble synaptic vesicles while others resemble dense core bodies. Structurally different types of synapses and axons can be distinguished within the neuropil.     

Paternal deprivation induces dendritic and synaptic changes and hemispheric asymmetry of pyramidal neurons in the somatosensory cortex

Similar to maternal care, paternal care is a source of neonatal sensory stimulation, which in primates and rodents has been shown to be essential for developing structure and function of sensory cortices. The aim of our study in the biparental rodent Octodon degus was to assess the impact of paternal deprivation on dendritic and synaptic development in the somatosensory cortex. We (i) quantified the amount of paternal care in relation to total parental investment and (ii) compared dendritic and synaptic development of pyramidal neurons in the somatosensory cortex of animals raised by a single mother or by both parents. On the behavioral level we show that paternal care comprises 37% of total parent‐offspring interactions, and that the somatosensory stimulation provided by the fathers primarily consists of huddling, licking/grooming, and playing. On the morphological level we found that, compared with offspring raised by both parents (mother and father), the father‐deprived animals displayed significantly reduced spine numbers on the basal dendrites of pyramidal neurons. Furthermore, paternal deprivation induces hemispheric asymmetry of the dendritic morphology of somatosensory pyramidal neurons. Father‐deprived animals show shorter and less complex basal dendrites in the left somatosensory cortex compared with the right hemisphere. These findings indicate that paternal deprivation results in delayed or retarded dendritic and synaptic development of somatosensory circuits. © 2009 Wiley Periodicals, Inc. Develop Neurobiol, 2009     

Ultrastructure of neurons and synapses in the tentacle gastrodermis of the sea anemone Calliactis parasitica

Little is known about gastrodermal neurons and synapses in the tentacles of sea anemones. Using transmission electron microscopy of serial thin sections of Calliactis parasitica, we have identified both a sensory cell and a ganglion cell with granular vesicles originating from the Golgi complex and have identified four types of synapses in the tentacular gastrodermal nerve plexus. The sensory cell has a recessed apical cilium with a basal body and a perpendicularly oriented centriole, below which are several strands of striated rootlets surrounded by mitochondria. The ganglion cell lacks a cilium and resembles a bipolar neuron, with oppositely directed processes lying parallel to the basally located circular smooth muscle. Both one-way and two-way interneuronal synapses are present with 60- to 90-nm granular vesicles of various densities aligned at the paired electron-dense membranes and fine cross filaments in the intervening 13-nm cleft. Two types of neuroeffector synapses have been located. Dense granular vesicles are present at neuromuscular synapses, whereas less dense vesicles are present at neuroglandular synapses. Most of the synaptic vesicles range from 60 to 120 nm in diameter. Two types of nerve cells and a variety of synaptic loci provide morphological substrates for the spontaneous SS2 conduction pulses in the tentacular gastrodermis of C. parasitica. J Morphol 231:217–223, 1997. © 1997 Wiley-Liss, Inc.     

Effect of auditory deafferentation on the synaptic connectivity of a pair of identified interneurons in adult field crickets

In adult crickets, Teleogryllus oceanicus, unilateral auditory deafferentation causes the medial dendrites of an afferent-deprived, identified auditory interneuron (Int-1) in the prothoracic ganglion to sprout and form new functional connections in the contralateral auditory neuropil. The establishment of these new functional connections by the deafferented Int-1, however, does not appear to affect the physiological responses of Int-1's homolog on the intact side of the prothoracic ganglion which also innervates this auditory neuropil. Thus it appears that the sprouting dendrites of the deafferented Int-1 are not functionally competing with those of the intact Int-1 for synaptic connections in the remaining auditory neuropil following unilateral deafferentation in adult crickets. Moreover, we demonstrate that auditory function is restored to the afferent-deprived Int-1 within 4-6 days following deafferentation, when few branches of Int-1's medial dendrites can be seen to have sprouted. The strength of the physiological responses and extent of dendritic sprouting in the deafferented Int-1 progressively increase with time following deafferentation. By 28 days following deafferentation, most of the normal physiological responses of Int-1 to auditory stimuli have been restored in the deafferented Int-1, and the medial dendrites of the deafferented Int-1 have clearly sprouted and grown across into the contralateral auditory afferent field. The strength of the physiological responses of the deafferented Int-1 to auditory stimuli and extent of dendritic sprouting in the deafferented Int-1 are greater in crickets deafferented as juveniles than as adults. Thus, neuronal plasticity persists in Int-1 following sensory deprivation from the earliest juvenile stages through adulthood.     

Neural pathways and innervation of cnidocytes in tentacles of sea anemones

Our previously published studies are here reviewed detailing neuro-cnidocyte synapses, demonstrating putative neurotransmitter substances, and identifying complex neural pathways in sea anemones. Synapses were traced to their contacts on nematocytes and spirocytes by transmission electron microscopy of serial thin sections of tentacles. In five animals, cells containing microbasic p-mastigophores had synapses with clear vesicles, whereas cells containing basitrichous isorhizas had synapses with dense-cored vesicles, providing preliminary evidence for a selectivity of neurotransmitter types for different nematocysts. Either clear or dense-cored synaptic vesicles were also present at neuro-spirocyte contacts. Antho-RFamide immunoreactivity occurred in some anthozoan synaptic vesicles and immunogold labeling of serotonin was found at a neuro-spirocyte synapse. Neural pathways included direct innervation of spirocytes by sensory cells, sequential neuro-neuro-spirocyte and neuro-neuro-nematocyte synapses and reciprocal synapses involving axons of both sensory cells and ganglion cells. Such synaptic patterns resemble neuro-effector pathways found in higher animals and lay to rest the independent effector hypothesis for cnidocyte discharge in tentacles of sea anemones.     

Whisker Deprivation Drives Two Phases of Inhibitory Synapse Weakening in Layer 4 of Rat Somatosensory Cortex

Inhibitory synapse development in sensory neocortex is experience-dependent, with sustained sensory deprivation yielding fewer and weaker inhibitory synapses. Whether this represents arrest of synapse maturation, or a more complex set of processes, is unclear. To test this, we measured the dynamics of inhibitory synapse development in layer 4 of rat somatosensory cortex (S1) during continuous whisker deprivation from postnatal day 7, and in age-matched controls. In deprived columns, spontaneous miniature inhibitory postsynaptic currents (mIPSCs) and evoked IPSCs developed normally until P15, when IPSC amplitude transiently decreased, recovering by P16 despite ongoing deprivation. IPSCs remained normal until P22, when a second, sustained phase of weakening began. Delaying deprivation onset by 5 days prevented the P15 weakening. Both early and late phase weakening involved measurable reduction in IPSC amplitude relative to prior time points. Thus, deprivation appears to drive two distinct phases of active IPSC weakening, rather than simple arrest of synapse maturation.     

Cofilin1 Controls Transcolumnar Plasticity in Dendritic Spines in Adult Barrel Cortex

During sensory deprivation, the barrel cortex undergoes expansion of a functional column representing spared inputs (spared column), into the neighboring deprived columns (representing deprived inputs) which are in turn shrunk. As a result, the neurons in a deprived column simultaneously increase and decrease their responses to spared and deprived inputs, respectively. Previous studies revealed that dendritic spines are remodeled during this barrel map plasticity. Because cofilin1, a predominant regulator of actin filament turnover, governs both the expansion and shrinkage of the dendritic spine structure in vitro, it hypothetically regulates both responses in barrel map plasticity. However, this hypothesis remains untested. Using lentiviral vectors, we knocked down cofilin1 locally within layer 2/3 neurons in a deprived column. Cofilin1-knocked-down neurons were optogenetically labeled using channelrhodopsin-2, and electrophysiological recordings were targeted to these knocked-down neurons. We showed that cofilin1 knockdown impaired response increases to spared inputs but preserved response decreases to deprived inputs, indicating that cofilin1 dependency is dissociated in these two types of barrel map plasticity. To explore the structural basis of this dissociation, we then analyzed spine densities on deprived column dendritic branches, which were supposed to receive dense horizontal transcolumnar projections from the spared column. We found that spine number increased in a cofilin1-dependent manner selectively in the distal part of the supragranular layer, where most of the transcolumnar projections existed. Our findings suggest that cofilin1-mediated actin dynamics regulate functional map plasticity in an input-specific manner through the dendritic spine remodeling that occurs in the horizontal transcolumnar circuits. These new mechanistic insights into transcolumnar plasticity in adult rats may have a general significance for understanding reorganization of neocortical circuits that have more sophisticated columnar organization than the rodent neocortex, such as the primate neocortex.