Postnatal neuronal plasticity of the pyramidal cells of CA1 area of the hippocampus as a reaction to neurotoxic damage.

Kainic acid (KA) was injected into both lateral ventricles of the brain of adult laboratory rats with the aim of verifying whether damage to afferent fibres in the hippocampal CA1 area would also be reflected in changes in the dendritic arborization of the neurones after maturation of these structures was completed. A significant proportion of the afferent fibres ending in area CA1 comes from CA3-4. The neurodegenerative effect of KA on the neurones in CA3-4 thus leads to marked reconstruction of the dendritic network of the pyramidal cells in the CA1 area. In the CA1 area of the experimental animals, there are fewer segments in the proximal part of the basal dendrites and in the lateral branches of the apical dendrites. The total number of segments in the apical dendrites is smaller and the higher order segments are likewise reduced. In the experimental group, the segments of both the basal and the apical dendrites are shorter. In the experimental animals, dendritic spine density in the lateral preterminal branches, the distal part of the apical shaft, the terminal segments of the lateral branches and the apical preterminal branches are smaller than in the controls, whereas in the segments proximal to the soma of the pyramidal cells it is greater. It can be seen from the results that area CA1 of the hippocampus is endowed, even in adulthood, not only with high functional plasticity, but also with surprisingly high morphological plasticity.     

Remodelling during development of the Purkinje cell dendritic tree in the mouse

The development of the Purkinje cells in normal C57 mice was studied from 7-100 d post natum . The growth of the dendritic trees was analysed both metrically and topologically using the method of vertex analysis (Berry & Flinn 1983 a). Granule and PUrkinje cell counts were made so that Purkinje cell segment production could be correlated with the number of parallel fibres deposited. Both topological and metrical results indicate that from 7 to 30 d post natum the Purkinje cell dendritic trees expand massively; accounting for 87% of total segment elaboration, reaching their lateral boundaries by 12-15 d post natum and then advancing towards the pial surface. Continued lateral expansion is constrained by the proximity of dendrites from neighbouring trees. Growth proceeds upwards through the neuropil as a front of prolific random terminal branching with inhibitory forces acting at the edges of the growth corridor and behind the growth front to prevent overlapping of dendrites. By 30 d post natum all boundaries are reached and the size of the dendritic field is fixed. Trees averaged 711.2 segments +/- 21.45 with a mean distance from root to terminal segment of 133.5 +/- 2.9 micrometers. The Va/Vb vertex ratios and the levels of trichotomy during this period indicate that branching patterns deviate from pure random terminal additions in a dichotomous tree. There is opportunity for non-random growth at the areas of inhibitory action. Beyond 30 d post natum remodelling occurs within the arbor which involves segment loss in the subpial region (orders above 16) and segment elaboration within the tree (orders 8-16) causing increased density of dendrites and overlapping of segments. The frequencies of segments and terminals are restored to symmetrical distributions through the orders of the trees from the skewed distributions associated with the frontal advance in earlier growth. During remodelling the Va/Vb vertex ratios and percentage of trichotomous nodes are consistent with growth through dichotomous random terminal branching. Path lengths of 8 micrometers between each order are seen as regular increments throughout entire trees at 100 d post natum . The final tree produced is indistinguishable from a network grown entirely by random terminal dichotomous branching with some 6% trichotomy and a Va/Vb vertex ratios of 0.92. Granule cell number within the granular layer increases rapidly up to 15 d post natum after which cell death causes a decrease to stable levels beyond 30 d post natum .(ABSTRACT TRUNCATED AT 400 WORDS)     

Vertex analysis of Purkinje cell dendritic trees in the cerebellum of the rat

All networks are made up of vertices (points interconnected by segments), which include terminals interconnected by terminal segments, nodes interconnected by link segments and the root point connected to the tree by the root segment. All nodes may be classified into unique types according to the number of terminal and link segments they drain. For example, there are three distinct dichotomous nodes, a 'primary' node draining two terminal segments, a 'secondary' node draining one terminal segment and a link segment, and a 'tertiary' node draining two link segments. The numbers of primary and tertiary nodes approximate to equality in large networks and thus the ratio of primary to secondary nodes defines topology. All higher order nodes ( trichotomous and beyond) may be resolved into dichotomous forms and incorporated into the analysis. Different forms of growth may thus be analysed by comparing the frequency distributions of nodes with those generated by computer simulated growth models. Moreover, all vertices can be ordered so that metrical parameters are easily incorporated and the hierarchical arrangements of vertices of different order discerned. The dendritic trees of 48 Purkinje cells, taken from folia along the primary fissure, were analysed using vertex analysis. The mean number of segments in Purkinje cell trees was 881 +/- 23 (s.e.) and mean total dendritic length 7959 +/- 233 (s.e.) micrometers. Segment lengths were longest over proximal segments but over most of the tree segment lengths were constant at 10 +/- 0.2 (s.e.) micrometers. Vertex, segment and terminal frequency distributions of equivalent orders were all normal with a slight positive skew. Peak frequencies were recorded at the 12th equivalent order. The mean primary/secondary nodal vertex ratio was 0.93 and the proportion of trichotomous branch points in the tree was 5%. Comparison of the frequency distribution of all vertices with computer generated models showed that growth of the Purkinje cell was most closely simulated by a random terminal growth model, incorporating 5% trichotomy , in which the branching of high order terminals was more likely than low order terminals. It was concluded that growth of the Purkinje cell tree could proceed by random terminal branching with growth occurring preferentially over a front composed of terminals that are ascending through a corridor in the molecular layer whose margins are defined by neighbouring trees.     

Localization of Hexokinase in Neural Tissue: Electron Microscopic Studies of Rat Cerebellar Cortex

: The distribution of hexokinase (ATP:d -hexose 6-phosphotransferase, EC 2.7.1.1) in the rat cerebellar cortex has been studied at the electron microscopic level using the peroxidase-antiperoxidase procedure. Extensive staining of cytoplasmic regions, with some increased staining at mitochondrial profiles, was seen in the cell bodies of both neurons (basket, stellate, Lugaro, Golgi, and granule cells) and astrocytes. Oligodendrocytes showed little or no detectable staining. Purkinje cell perikarya were much less intensely stained than were the perikarya of other neurons. The initial portion of the Purkinje dendrite was, like the perikaryon from which it emerged, lightly stained. More intense staining was seen in the secondary and tertiary branches of the Purkinje dendrite, but the terminal branches were devoid of stain. Granule cell dendrites were well stained in their initial portions but devoid of stain in their terminal dendritic digits which form part of the cerebellar glomeruli. In contrast to the unstained granule cell dendritic digits, the central mossy fiber nerve terminal of the glomerulus exhibited intense staining of the mitochondrial profiles and of synaptic vesicles adjacent to the mitochondria. Axons of basket cells showed intense staining in the segments adjacent to the Purkinje cell soma, while terminal twigs of the basket axons in the pinceau surrounding the (unstained) initial segment of the Purkinje axon showed markedly decreased staining intensity. These results indicate that there may be substantial variation in hexokinase levels between the various regions of neuronal processes. Hexokinase was seen at both cytoplasmic and mitochondrial locations in a variety of cells. It does not appear likely that location of hexokinase can be directly correlated with cell type, i.e., with neurons versus glia.     

Statistical evaluation of dendritic growth models

A mathematical model (Kliemann, W. 1987.Bull. math. Biol. 49, 135–152.) that predicts the quantitative branching pattern of dendritic tree was evaluated using the apical and basal dendrites of rat hippocampal neurons. The Wald statistics for χ²-test was developed for the branching pattern of dendritic trees and for the distribution of the maximal order of the tree. Using this statistic, we obtained a reasonable, but not excellent, fit of the mathematical model for the dendritic data. The model's predictability of branching patterns was greatly enhanced by replacing one of the assumptions used for the original model “splitting of branches for all dendritic orders is stochastically independent”, with a new assumption “branches are more likely to split in areas where there is already a high density of branches”. The modified model delivered an excellent fit for basal dendrites and for the apical dendrites of hippocampal neurons from young rats (30–34 days postpartum). This indicates that for these cells the development of dendritic patterns is the result of a purely random and a systematic component, where the latter one depends on the density of dendritic branches in the brain area considered. For apical dendrites there is a trend towards decreasing pattern predictability with increasing age. This appears to reflect the late arrival of afferents and subsequent synaptogenesis proximal on the apical dendritic tree of hippocampal neurons.     

Chronic stress alters dendritic morphology in rat medial prefrontal cortex

Chronic stress produces deficits in cognition accompanied by alterations in neural chemistry and morphology. Medial prefrontal cortex is a target for glucocorticoids involved in the stress response. We have previously demonstrated that 3 weeks of daily corticosterone injections result in dendritic reorganization in pyramidal neurons in layer II-III of medial prefrontal cortex. To determine if similar morphological changes occur in response to chronic stress, we assessed the effects of daily restraint stress on dendritic morphology in medial prefrontal cortex. Male rats were exposed to either 3 h of restraint stress daily for 3 weeks or left unhandled except for weighing during this period. On the last day of restraint, animals were overdosed and brains were stained using a Golgi-Cox procedure. Pyramidal neurons in lamina II-III of medial prefrontal cortex were drawn in three dimensions, and the morphology of apical and basilar arbors was quantified. Sholl analyses demonstrated a significant alteration of apical dendrites in stressed animals: overall, the number and length of apical dendritic branches was reduced by 18 and 32%, respectively. The reduction in apical dendritic arbor was restricted to distal and higher-order branches, and may reflect atrophy of terminal branches: terminal branch number and length were reduced by 19 and 35%. On the other hand, basilar dendrites were not affected. This pattern of dendritic reorganization is similar to that seen after daily corticosterone injections. This reorganization likely reflects functional changes in prefrontal cortex and may contribute to stress-induced changes in cognition.     

The pyramidal cells of Betz within the cingulate and precentral gigantopyramidal field in the human brain

Summary It can be demonstrated with the aid of Golgi-, Nissl-, and pigment preparations that the Betz cells represent a homogeneous class of giant cells within the human brain, which can readily be distinguished from other large pyramids by their densely aggregated lipofuscin deposits. In addition to the primary motor field (4, Brodmann), there exists only a small area on the medial surface of the hemisphere in front of the central sulcus which also contains large Betz pyramids in layer Vb. This recently discovered gigantopyramidal field is almost totally buried in the depth of the cingulate sulcus (Braak, 1976b). Compared with the Betz cells of the primary motor field (4, Brodmann), those of the cingulate area display numerous primitive traits. A small number of short basal dendrites springs off from the cell body. The apical dendrite forks in a short distance from the perikaryon repeatedly but issues only few side branches. A spine-free proximal dendritic segment is poorly developed or lacking. Moreover, numerous spines are encountered along the surface of the soma. In view of their primitive features the large pyramids of the cingulate gigantopyramidal area are interpreted as the forerunners of the precentral Betz pyramids.Supported by the Deutsche Forschungsgemeinschaft (Br 317/7). Dedicated to Prof. Dr. med. Drs. h.c. W. Bargmann in honour of this 70th birthday     

Hippocampal Feedforward Inhibition Focuses Excitatory Synaptic Signals into Distinct Dendritic Compartments

Feedforward inhibition controls the time window for synaptic integration and ensures temporal precision in cortical circuits. There is little information whether feedforward inhibition affects neurons uniformly, or whether it contributes to computational refinement within the dendritic tree. Here we demonstrate that feedforward inhibition crucially shapes the integration of synaptic signals in pyramidal cell dendrites. Using voltage-sensitive dye imaging we studied the transmembrane voltage patterns in CA1 pyramidal neurons after Schaffer collateral stimulation in acute brain slices from mice. We observed a high degree of variability in the excitation-inhibition ratio between different branches of the dendritic tree. Many dendritic segments showed no depolarizing signal at all, especially the basal dendrites that received predominantly inhibitory signals. Application of the GABA_A receptor antagonist bicuculline resulted in the spread of depolarizing signals throughout the dendritic tree. Tetanic stimulation of Schaffer collateral inputs induced significant alterations in the patterns of excitation/inhibition, indicating that they are modified by synaptic plasticity. In summary, we show that feedforward inhibition restricts the occurrence of depolarizing signals within the dendritic tree of CA1 pyramidal neurons and thus refines signal integration spatially.     

Bergmann glia and the recognition molecule CHL1 organize GABAergic axons and direct innervation of Purkinje cell dendrites

The geometric and subcellular organization of axon arbors distributes and regulates electrical signaling in neurons and networks, but the underlying mechanisms have remained elusive. In rodent cerebellar cortex, stellate interneurons elaborate characteristic axon arbors that selectively innervate Purkinje cell dendrites and likely regulate dendritic integration. We used GFP BAC transgenic reporter mice to examine the cellular processes and molecular mechanisms underlying the development of stellate cell axons and their innervation pattern. We show that stellate axons are organized and guided towards Purkinje cell dendrites by an intermediate scaffold of Bergmann glial (BG) fibers. The L1 family immunoglobulin protein Close Homologue of L1 (CHL1) is localized to apical BG fibers and stellate cells during the development of stellate axon arbors. In the absence of CHL1, stellate axons deviate from BG fibers and show aberrant branching and orientation. Furthermore, synapse formation between aberrant stellate axons and Purkinje dendrites is reduced and cannot be maintained, leading to progressive atrophy of axon terminals. These results establish BG fibers as a guiding scaffold and CHL1 a molecular signal in the organization of stellate axon arbors and in directing their dendritic innervation.     

Principles of branch dynamics governing shape characteristics of cerebellar Purkinje cell dendrites

Neurons develop dendritic arbors in cell type-specific patterns. Using growing Purkinje cells in culture as a model, we performed a long-term time-lapse observation of dendrite branch dynamics to understand the rules that govern the characteristic space-filling dendrites. We found that dendrite architecture was sculpted by a combination of reproducible dynamic processes, including constant tip elongation, stochastic terminal branching, and retraction triggered by contacts between growing dendrites. Inhibition of protein kinase C/protein kinase D signaling prevented branch retraction and significantly altered the characteristic morphology of long proximal segments. A computer simulation of dendrite branch dynamics using simple parameters from experimental measurements reproduced the time-dependent changes in the dendrite configuration in live Purkinje cells. Furthermore, perturbation analysis to parameters in silico validated the important contribution of dendritic retraction in the formation of the characteristic morphology. We present an approach using live imaging and computer simulations to clarify the fundamental mechanisms of dendrite patterning in the developing brain.     

Formation of new synaptic contacts by Purkinje axon collaterals in the granular layer of deafferented cerebellar cortex of adult rat

In addition to (i) mossy terminals, (ii) Golgi axons, (iii) granule cell dendrites and (iv), occasionally, Golgi cell dendrites, a third axonal profile identified by morphological criteria as the collateral of Purkinje axons, has been found in 2% of all cerebellar glomeruli. These infrequent components of a few glomeruli, however, were never seen in normal cerebellar cortex to establish specialized synaptic contact with glomerular dendrites. Two to four weeks after surgical isolation of the cerebellar cortex, i.e. following the destruction of both efferent and afferent fibres, the number of glomeruli containing (hypertrophic) axonal branches of Purkinje cells has increased to 13% of all surveyed glomeruli. In addition, the Purkinje axon terminals in the mossy fibre-deprived glomeruli were observed to establish numerous Gray II-type synaptic contacts with surrounding granule cell dendrites. It is suggested that the development of heterologous synapses between hypertrophic, or even intact, Purkinje axon collaterals on the one hand and the mossy fibre-vacated granule cell dendrites on the other, is a compensatory, reactive process to the synaptic "desaturation" of granule neurons, which demonstrate a dormant potential of Purkinje cells to form new synaptic contacts in the adult cerebellum.     

Quantitative studies on the neuron structure of the rat fascia dentata]

1. In 6 month old male rats the structure of dendrites and the distribution of spines on the apical dendrites of granular cells of the dentate gyrus were investigated by light microscopy and statistical methods. 2. The number of dendrites of the first, second and third order of granular cells increases in this sequence in a ratio of 1:2:3; the total length of the dendrites increases correspondently in a ratio of 1:4:5. 3. The mean number of origin points of dendritic branches is 10, the mean number of free dendritic endings is 12. 4. The number of spines per a 25 mum dendritic segment near to the pericaryon (dendritic segment A), in the middle of the dendrite (dendritic segment B) and in the peripheral dendritic part (dendritic segment C) as well as the distribution of spines in the whole apical dendrite was evaluated. The total mean of spines of granular cell apical dendrites of the dentate gyrus (superior respectively inferior) is 12 respectively 10 for the dendritic segment A, 18 respectively 17 for the dendritic segment B and 17 respectively 15 for the dendritic segment C calculated for a dendritic length of 25 mum. 5. The spine density in each case depends upon the distance pericaryondendritic segment and is in close relation to the adjacent layers with their specific afferents. 6. The averaged total number of spines per 1 mum of dendritic length is 0,62 spines/mum for the dentate gyrus (superior) and 0,57 spines/mum for the dentate gyrus (inferior). 7. The granular cells of the dentate gyrus (superior) have a mean dendritic length of a total of 357 mum with a total of 226 visible spines; the granular cells of the dentate gyrus (inferior) have a mean dendritic length of a total of 450 mum with a total of 258 visible spines.     

Morphometry of cat's pulmonary arterial tree

Morphometic data of the pulmonary artery in the cat's right lung are presented. Silicone elastomer casts of cat's right lung were made, and measured, counted and analyzed. The Strahler system is used to describe the branching pattern of the arterial vascular tree. These data are needed for any quantitative approach to the study of the pulmonary circulation. For all the pulmonary blood vessels of the cat lying between the main pulmonary artery and the capillary beds, there are a total of 10 orders of vessels in the right upper lobe, 9 orders of vessels in the right middle lobe and 11 orders of vessels in the right lower lobe. The ratio of the number of branches in successive orders of vessels or the branching ratio, is 3.58. The corresponding average diameter ratio is 1.72, whereas the average length ratio is 1.81.     

Alterations in the morphology of ganglion cell dendrites in the adult rat retina after optic nerve transection and grafting of peripheral nerve segments

Summary Transected ganglion cell axons from the adult retina are capable of reinnervating their central targets by growing into transplanted peripheral nerve (PN) segments. Injury of the optic nerve causes various metabolic and morphological changes in the retinal ganglion cell (RGC) perikarya and in the dendrites. The present work examined the dendritic trees of those ganglion cells surviving axotomy and of those whose severed axons re-elongated in PN grafts to reach either the superior colliculus (SC), transplanted SC, or transplanted autologous thigh muscle. The elaboration of the dendritic trees was visualized by means of the strongly fluorescent carbocyanine dye DiI, which is taken up by axons and transported to the cell bodies and from there to the dendritic branches. Alternatively, retinofugal axons regrowing through PN grafts were anterogradely filled from the eye cup with rhodamine B-isothiocyanate. The transection of the optic nerve resulted in characteristic changes in the ganglion cell dendrites, particularly in the degeneration of most of the terminal and preterminal dendritic branches. This occurred within the first 1 to 2 weeks following axotomy. The different types of ganglion cells appear to vary in their sensitivity to axotomy, as reflected by a rapid degeneration of certain cell dendrites after severance of the optic nerve. The most vulnerable cells were those with small perikarya and small dendritic fields (type II), whereas larger cells with larger dendritic fields (type I and III) were slower to respond and less dramatically affected. Regrowth of the lesioned axons in peripheral nerve grafts and reconnection of the retina with various tissues did not result in a significant immediate recovery of ganglion cell dendrites, although it did prevent some axotomized cells from further progression toward posttraumatic cell death.     

Structure-Dependent Electrical and Concentration Processes in the Dendrites of Pyramidal Neurons of Superficial Neocortical Layers: Model Study

On mathematical models of pyramidal neurons localized in the neocortical layers 2/3, whose reconstructed dendritic arborization possessed passive linear or active nonlinear membrane properties, we studied the effect of morphology of the dendrites on their passive electrical transfer characteristics and also on the formation of patterns of spike discharges at the output of the cell under conditions of tonic activation via uniformly distributed excitatory synapses along the dendrites. For this purpose, we calculated morphometric characteristics of the size, complexity, metric asymmetry, and function of effectiveness of somatopetal transmission of the current (with estimation of the sensitivity of this efficacy to changes in the uniform membrane conductance) for the reconstructed dendritic arborization in general and also for its apical and basal subtrees. Spatial maps of the membrane potential and intracellular calcium concentration, which corresponded to certain temporal patterns of spike discharges generated by the neuron upon different intensities of synaptic activation, were superimposed on the 3D image and dendrograms of the neuron. These maps were considered “spatial autographs” of the above patterns. The main discharge pattern included periodic two-spike bursts (dublets) generated with relatively stable intraburst interspike intervals and interburst intervals decreasing with a rise in the intensity of activation. Under conditions of intense activation, the interburst intervals became close to the intraburst intervals, so the cell began to generate continuous trains of action potentials. Such a repertoire (consisting of two patterns of the activity, periodical dublets and continuous discharges) is considerably scantier than that described earlier in pyramidal neurons of the neocortical layer 5. Under analogous conditions of activation, we observed in the latter cells a variety of patterns of output discharges of different complexities, including stochastic ones. A relatively short length of the apical dendrite subtree of layer 2/3 neurons and, correspondingly, a smaller metric asymmetry (differences between the lengths of the apical and basal dendritic branches and paths), as compared with those in layer 5 pyramidal neurons, are morphological factors responsible for the predominance of periodic spike dublets. As a result, there were two combinations of different electrical states of the sites of dendritic arborization (“spatial autographs”). In the case of dublets, these were high depolarization of the apical dendrites vs. low depolarization of the basal dendrites and a reverse combination; only the latter (reverse) combination corresponded to the case of continuous discharges. The relative simplicity and uniformity of spike patterns in the cells, apparently, promotes the predominance of network interaction in the processes of formation of the activity of pyramidal neurons of layers 2/3 and, thereby, a higher efficiency of the processes of intracortical association.     

THE VISUALIZATION OF CONCANAVALIN A BINDING SITES IN PURKINJE CELL SOMATA AND DENDRITES OF RAT CEREBELLUM

The localization of concanavalin A (Con A) binding sites in Purkinje cell somata and dendrites has been studied using a peroxidase labeling technique. In the somata, the nuclear, Golgi, and endoplasmic reticulum (ER) membranes are rich in Con A binding sites. The hypolemmal cisternae, which are continuous with the ER from the soma and throughout the dendritic tree of Purkinje cells, are also rich in Con A binding sites. Other cisternae seen in these dendrites do not bind detectable amounts of Con A. The results suggest that a cisternal system, rich in carbohydrate, may be continuous from the nuclear envelope to distal dendritic segments of Purkinje cells. Such a system could play a role in the movement of materials from Purkinje somata to dendrites.     

Control of dendritic branching and tiling by the Tricornered-kinase/Furry signaling pathway in Drosophila sensory neurons

To cover the receptive field completely but without redundancy, neurons of certain functional groups exhibit tiling of their dendrites via dendritic repulsion. Here we show that two evolutionarily conserved proteins, the Tricornered (Trc) kinase and Furry (Fry), are essential for tiling and branching control of Drosophila sensory neuron dendrites. Dendrites of fry and trc mutants display excessive terminal branching and fail to avoid homologous dendritic branches, resulting in significant overlap of the dendritic fields. Trc control of dendritic branching involves regulation of RacGTPase, a pathway distinct from the action of Trc in tiling. Timelapse analysis further reveals a specific loss of the ability of growing dendrites to turn away from nearby dendritic branches in fry mutants, suggestive of a defect in like-repels-like avoidance. Thus, the Trc/Fry signaling pathway plays a key role in patterning dendritic fields by promoting avoidance between homologous dendrites as well as by limiting dendritic branching.     

Disynaptic inhibition between neocortical pyramidal cells mediated by Martinotti cells

Reliable activation of inhibitory pathways is essential for maintaining the balance between excitation and inhibition during cortical activity. Little is known, however, about the activation of these pathways at the level of the local neocortical microcircuit. We report a disynaptic inhibitory pathway among neocortical pyramidal cells (PCs). Inhibitory responses were evoked in layer 5 PCs following stimulation of individual neighboring PCs with trains of action potentials. The probability for inhibition between PCs was more than twice that of direct excitation, and inhibitory responses increased as a function of rate and duration of presynaptic discharge. Simultaneous somatic and dendritic recordings indicated that inhibition originated from PC apical and tuft dendrites. Multineuron whole-cell recordings from PCs and interneurons combined with morphological reconstructions revealed the mediating interneurons as Martinotti cells. Martinotti cells received facilitating synapses from PCs and formed reliable inhibitory synapses onto dendrites of neighboring PCs. We describe this feedback pathway and propose it as a central mechanism for regulation of cortical activity.     

Investigation of the geometry of the dendritic tree of retinal ganglion cells

Two parameters are proposed for classifying the structure of the dendritic trees of retinal ganglion cells: the mean distance between the ends of the terminal segments of the dendrites from the soma ( ) and the ratio between the number of horizontal and vertical dendritic segments (a). By using these parameters the dendritic trees of ganglion cells were divided into four types. To describe the two-dimensional structure of dendritic trees the following parameters are used: the distribution of intermediate and terminal dendritic segments in each series of branches depending on the angle of their orientation relative to the boundaries of the inner plexiform layer — p_Z (), q_Z (), and depending on their length — P_Z (1), q_Z (1). Histograms of these functions are given. By means of these histograms the two-dimensional structure of the dendritic tree can be drawn in such a way as to reproduce the structural features of actual dendritic trees with the accuracy of the parameters defining their appearance. The structural and functional significance of the parameters used is discussed.Kaunas Medical Institute. Translated from Neirofiziologiya, Vol. 5, No. 3, pp. 307–314, May–June, 1973.     

Ion Channel Gradients in the Apical Tuft Region of CA1 Pyramidal Neurons

Dendritic ion channels play a critical role in shaping synaptic input and are fundamentally important for synaptic integration and plasticity. In the hippocampal region CA1, somato-dendritic gradients of AMPA receptors and the hyperpolarization-activated cation conductance (I_h) counteract the effects of dendritic filtering on the amplitude, time-course, and temporal integration of distal Schaffer collateral (SC) synaptic inputs within stratum radiatum (SR). While ion channel gradients in CA1 distal apical trunk dendrites within SR have been well characterized, little is known about the patterns of ion channel expression in the distal apical tuft dendrites within stratum lacunosum moleculare (SLM) that receive distinct input from the entorhinal cortex via perforant path (PP) axons. Here, we measured local ion channels densities within these distal apical tuft dendrites to determine if the somato-dendritic gradients of I_h and AMPA receptors extend into distal tuft dendrites. We also determined the densities of voltage-gated sodium channels and NMDA receptors. We found that the densities of AMPA receptors, I_h, and voltage-gated sodium channels are similar in tuft dendrites in SLM when compared with distal apical dendrites in SR, while the ratio of NMDA receptors to AMPA receptors increases in tuft dendrites relative to distal apical dendrites within SR. These data indicate that the somato-dendritic gradients of I_h and AMPA receptors in apical dendrites do not extend into the distal tuft, and the relative densities of voltage-gated sodium channels and NMDA receptors are poised to support nonlinear integration of correlated SC and PP input.