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PubMed1979

Quantitative analysis of dendritic branching. I. Simple formulae for the quantitative analysis of dendritic branching.

Dendritic branching of neurons may be quantitatively studied using applications of graph theory. Dendritic ramifications may be considered as 'forests' of planted, stemmed, binary 'trees' with simple mathematical properties. Two fundamental numbers, that of dendritic tips and that of dendritic stems, usually suffice to quantify the dendritic branching of one neuron as, in most cases, they allow the computation of all other numbers of dendritic points and segments.

Animals↗
PubMed1979

Dendro axonic neurotransmission. II. Morphological sites for the synthesis, binding and release of neurotransmitters in dopaminergic dendrites in the substantia nigra and cholinergic dendrites in the neostriatum.

Morphological evidence is presented indicating sites of synthesis, storage, and release of neurotransmitters in dendrites of dopaminergic cells of the substantia nigra and cholinergic cells of the neostriatum. Smooth endoplasmic reticulum can be identified in dopaminergic neurons touching the dendritic surface. The false transmitter for dopamine, 5-hydroxydopamine (5-OHDA), is localized to smooth endoplasmic reticulum or large vesicular structures which approach the dendritic surface. The dopamine synthesizing enzyme, tyrosine hydroxylase (TH), is localized to microtubules and smooth endoplasmic reticulum which approach the postsynaptic membrane. In the neostriatum, dopaminergic nerve endings make asymmetrical axospinous contacts. The postsynaptic spines often contain a few 'vesicles' near the postsynaptic thickenings. The surface and subsurface structures stain preferentially for choline acetyltransferase (CAT), the synthesizing enzyme for acetylcholine. It is hypothesized that neurotransmitters are released from dendrites as a general phenomenon in the CNS and that they can act upon axonal endings.

Acetylcholine↗
PubMed1979

Quantitative analysis of dendritic branching. II. Fundamental dendritic numbers as a tool for the study of neuronal groups.

The fundamental dendritic numbers [14] were studied in various neuronal groups from different species. Their mean values allow group characterization on a quantitative basis. This allows group comparisons and classification. Four sets of polydendritic neuronal groups are isolated: fewly branched Golgi I groups (A), mainly stemmed Golgi I groups (B), Golgi II groups (C) and highly branched Golgi I groups (D). Numerical interspecific comparisons allow quantitative phylogenetic studies. Only groups of the D set exhibit a significant evolution. Dendritic numbers are an efficient tool for neuronal group studies.

Animals↗
PubMed1979

Organization and development of brain stem auditory nuclei of the chicken: dendritic gradients in nucleus laminaris.

Nucleus laminaris (NL) is a third-order auditory nucleus in the avian brain stem which receives spatially-segregated binaural inputs from the second-order magnocellular nuclei. The organization of dendritic structure in NL was examined in Golgi-impregnated brains from hatchling chickens. Quantitative analyses of dendritic size and number were made from camera lucida drawings of 135 neurons sampled from throughout the nucleus. The most significant results of this study may be summarized as follows: (1) The preponderant neuron in n. laminaris may be characterized as having a cylindrical-to-ovoid cell body, about 20 micrometer in diameter. The neurons comprising NL were found to be nearly completely homogeneous in issuing their dendrites in a bipolar fashion: one group of dendrites is clustered on the dorsal surface of the cells, the other group on the ventral. The dendrites of NL are contained within the glia-free neuropil surrounding the nucleus. From the rostromedial to the caudolateral poles of NL there is a gradient of increasing extension of the dendrites, increasing number of tertiary and higher-order dendrites, and increasing distance from the somata of the occurrence of branching. (2) The total dendritic size (sum of the dorsal) and ventral dendritic lengths of the cells) increases 3-fold from the rostromedial to the caudolateral poles of NL. About 50% of the variance in dendritic size is accounted for by the position of the cells in NL, and the gradient of dendritic size increase has the same orientation across NL as the tonotopic gradient of decreasing characteristic frequency in NL. (3) From the rostromedial pole to the caudolateral pole of NL there is an 11-fold decrease in the number of primary dendrites along a gradient coinciding with the length and frequency gradients. Sixty-six percent of the variance in dendrite number is accounted for by position in the nucleus. (4) The correlation of dorsal and ventral dendritic size on a cell-by-cell basis is not high (r = 0.47), indicating a fair amount of variability on the single-cell level. On the other hand, the average dorsal dendritic length within an isofrequency band in NL correlates very highly with the average ventral dendritic length. Thus, on an areal basis, the amount of dendritic surface area offered to the dorsal and ventral afferents is tightly regulated. (5) The dorsal and ventral dendrites have separate gradients of increasing length and number across NL. The dorsal gradients are skewed toward the rostrocaudal axis, while the ventral dendritic gradients are skewed mediolaterally. (6) There was no correlation between either dendritic size or number of primary dendrites and the size of the somata in NL, which remains relatively constant throughout the nucleus. Several hypotheses about the ontogenetic control of dendritic structure are examined in light of the above data. Of these, the hypotheses that the ontogeny of dendritic size and number is largely under afferent control receives a great deal of circumstantial support.

Animals↗
PubMed1979

A technique for estimating total spine numbers on Golgi-impregnated dendrites.

The functional significance of dendritic spines and their morphological sensitivity to a wide spectrum of experimental manipulations and pathological states have led to a number of studies in which counts of dendritic spine number have been carried out. These studies have, for the most part, involved the enumeration of only those spines which protrude from the opaque shafts of Golgi-impregnated dendrites into the clear zones flanking the dendrite. Such counts, limited to only those spines which are visible, underrepresent the true total number of spines borne by the dendrites. The magnitude of underrepresentation correlates positively with dendritic shaft diameter and negatively with spine length. This seriously restricts the usefulness of comparisons of spine density between dendrites, or even between segments of the same dendrite. In the present report, a geometrically based method is presented whereby total dendritic spine numbers can be estimated with reasonable accuracy, taking into account factors such as dendrite diameter and spine length. The technique entails the following principal steps: a determination, for a given length of dendrite over which spines are to be enumerated, of the volume of the flanking zones in which spines are visible and can be counted; a determination of the volume of the entire zone which encircles the dendritic shaft and which contains all spines, both visible and not visible; and a proportional extrapolation from the number of visible spines to obtain an estimate of the true total spine number. Tests of the predictive accuracy of the technique using dendrites of known total spine number suggest that estimates which deviate from true total spine numbers by less than 10% can be achieved.

Animals↗
PubMed1975

Some aspects of the electroanatomy of dendrites.

An understanding of the neuronal function requires the knowledge of the electroanatomy of dendrites, which comprise the major area and receive the main input in most neurons. Some simplifying assumptions are necessary to describe the electrical characteristics of the dendritic tree. The applicability of the simplified model of a combined equivalent dendritic cylinder proposed by Rall, was tested and verified by a combined analysis of anatomic and electrical data from the same spinal motoneurons. Assuming a uniform somadendritic membrane, estimates of the specific membrane resistance (RM: 2,700 +/- 920 omegacm2) were made by relating the neuronal input resistance with the combined dendritic trunk parameter (sigmaD3/2: 320 +/- 150-10(-6) CM3/2). From these combined anatomic and electrical data the dendritic electrotonic lengths (Lgeom: 1.5 +/- 0.3 times the length constant) were derived. Comparable L values (Ltrans: 1.5 +/- 0.3) resulted independently from analysis of membrane voltage transients during current steps. The linear dendritic cable model has proved its applicability for the analysis of small voltage deflections during current step applications at the soma as well as for the analysis of the majority of minimal postsynaptic potentials (PSP's). During the transmission along the dendritic cable the PSP undergoes changes in shape. These changes often permit a determination of the distance of the dendritic input from the soma. Unfortunately, the attenuation of the dendritic signal cannot be directly assessed. Dendritic synaptic transmission can be observed in isolation in chromatolytic motoneurons because the somal synapses are peeled off from the soma by proliferating glial cells in the course of retrograde reaction. These observations support the prediction that the PSP's with relatively short rise-times and duration originate from synapses near the soma. It may be questioned as to whether the linear dendritic cable approximation also applies to the larger voltage displacements during excitatory synaptic action. Particularly interesting is an increase of the apparent membrane resistance during depolarization known as anomalous rectification. The anomalous rectification could be reversibly eliminated and turned into a normal rectification by the application of cobalt ions or other calcium antagonists. Therefore, it appears likely that this phenomenon is caused by a voltage-(and time-) dependent reaction of the membrane, consisting of a smoothly increased calcium conductance during depolarizations that are even subthreshold for eliciting action potentials. Such a process would result in a shortening of the dendritic electrotonic length and in facilitating the postsynaptic excitatory transmission.

Animals↗
PubMed1977

Age-related deterioration of pyramidal cell basal dendrites in rat auditory cortex.

The basal dendritic trees of layer V pyramidal cells in the rat auditory cortex were examined quantitatively in a group of 3-month-old and a group of 34- and 36-month-old rats. Two forms of analysis were used on the Golgi preparations: (1) the number of intersections between the basal dendrites and a series of concentric circles whose common center lies over the perikaryon center, and (2) the number of dendritic branches, by order, per neuron. The data indicate that in the old animals the density of the dendritic tree has decreased significantly within a radius of about 150mu of the perikaryon, yet the extent of the dendritic domain has not changed appreciably. Analysis of the dendritic branching suggests that there has been a deterioration not only in the peripheral branches of the dendritic tree, but also that entire dendrites have been lost. This loss of primary branches was confirmed through the reconstruction of layer V neuronal perikarya and their proximal dendrites from 1-mu plastic serial sections of auditory cortex. Concomitant with the loss of dendrites which accompanies advancing age is a tendency for the perikaryon to be smaller, but not distorted, in the old animals.

Aging↗
PubMed1979

Golgi and EM studies of the formation of dendritic and axonal arbors: the interneurons of the substantia gelatinosa of Rolando in newborn kittens.

Golgi studies in newborn kittens show that the two most prevalent interneurons in Rexed's lamina II of the dorsal horn of the medulla, the stalked cell and islet cell (Gobel, '75a,b, '78b) form their dendritic arbors in a similar fashion. At birth, both cell types are present in forms ranging from immature, in which numerous short dendrites radiate from the cell body in all directions, to relatively mature in which their dendritic arbors have elongated in specific directions and the adult branching pattern is already evident. During postnatal maturation, many dendrites are lost while only a few go on to lengthen. The unmyelinated axons of both cells are first recognized in forms in which lengthening dendrites have taken on their preferred direction of orientation. The two parts of Rexed's lamina II, i.e., layers IIa and IIb have already reached their adult mediolateral width at birth and the neuropil has nearly achieved its adult compactness. Space in the compact neuropil for elongating neuronal and astrocytic processes becomes available through the disintegration of many existing dendrites and by an overall fourfold increase in the rostrocaudal length of the dorsal horn of the medulla during postnatal maturation. At birth, the lengthening of the plasma membranes of elongating neuronal and astrocytic processes proceeds as vesicles (addition vesicles) found in aggregates throughout dendrites, unmyelinated axons and astrocytic processes fuse with and become incorporated into the existing plasma membranes. In addition, many dendrites in layers IIa and IIb are beading up and disintegrating. Within the beads, neurotubules are lost and addition vesicles fuse with each other to form small cavities. These cavities continue to enlarge, hollowing out the beads. The cavities ultimately open to the intercellular space as their membranes fuse with the plasma membrane of the beads. Finally, the beads disintegrate and their plasma membranes fragment. The thread-like segments between adjacent disintegrating beads shrivel until they ultimately disappear. Disintegration of beaded dendrites results in very little debris and does not provoke a phagocytic glial reaction. The disintegration of a dendritic branch takes place in spite of synaptic input from non-primary axons and is thought to occur from a failure to establish synaptic connections with primary axonal endings.

Animals↗
PubMed1979

Tangential orientation and spatial order in dendrites of cat auditory cortex: a computer microscope study of Golgi-impregnated material.

In the tangential plane (parallel to the pial surface) dendrites in the primary auditory cortex (A1) of cat were found to exhibit preferentially oriented growth. This was shown by means of a computer microscope study of Golgi-Cox stained neurons as seen in 100 micrometers and 300 micrometers thick tangential sections. Two techniques were used to represent the 3-dimensional structure of dendrites: the "dendritic stick" and the "dendritic trumpet". The former dismembers a dendrite into its individual segments; the latter considers a dendrite as an entity and represents it by its centroid, its moments and the spatial dispersionof its branches. Both statistical and Fourier analyses of the data show that within the tangential plane there is a significant and consistent orientation of the dendritic sticks in a dorso-ventral direction which seems correlated with the cortical isofrequency contours observed in electrophysiological maps of the A1 region. The dendritic trumpet analyses also show a distinctly non-random vertical distribution of pyramidal cell basal dendrites but not of stellate cell dendrites.

Animals↗
PubMed1975

Motoneuron dendrites: role in synaptic integration.

Dendrites constitute over 80 per cent of the receptive surface area in cat motoneurons. Calculations based on matched electrical and gemoetrical measurements in these neurons indicate that the specific resistance of dendritic membranes in resting motoneurons is at least 2,000 ohm-cm2. When the specific membrane resistance is this high, even the most distal dendritic synapses can contribute significantly to the depolarization of the soma, and hence influence the rate of action potential generation. However, dendritic membrane resistance depends strongly on the level of background synaptic activity. The conductance changes associated with excitatory synaptic activity on a dendrite can be great enough to reduce significantly both the excitatory synaptic driving potential and the effective membrane resistance on that dendrite, and thus greatly reduce the effectiveness of synapses on the dendrite. Inhibitory synaptic activity produces an even greater reduction in dendritic membrane resistance. Thus the relative effectiveness of dendritic synapses depends on the type, distribution, and intensity of background synaptic activity, as well as on dendritic geometry and resting membrane properties.

Action Potentials↗
PubMed1979

Golgi studies on Purkinje cell development in the frog during spontaneous metamorphosis. II. Details of dendritic development.

The development of Purkinje cell dendrites was studied in the bullfrog from premetamorphic tadpoles to 10-week-old postmetamorphic frog-lets by the Golgi-Kopsch method. In this species two distinct patterns of arbor formation may be seen, which appear to be related to differences in the timing of initial dendritic development. In Purkinje cells that begin development in early tadpole stages, the dendritic tree is elaborated by continuous and concomitant growth and branching, a process by which the developing arbor expands in both height and width. Arbor formation in Purkinje cells that begin development in metamorphosing tadpoles proceeds in two separate steps. Initially, dendrites of such cells elongate, but form only a few poorly developed branches; only when the arbor reaches near-adult height does branching become extensive. Additional differences present in Purkinje cells are reflected in the paucity of growth cones and filopodia in the tadpole, and numerous filopodia and growth cones in the metamorphic period. An interesting feature of dendritic development in this species is a tendency to alter the arboreal domain by the formation of extra-arboreal dendrites, and possibly by the occasional resorbtion of other partially formed dendrites. The pattern of dendritic development in the frog is different than in mammals and is difficult to interpret. Such unusual development may be due to disturbances in the timing of the formation of Purkinje cell dendrites and of the establishment of the external granular layer (EGL).

Animals↗
PubMed1979

Morphometrical synaptology of Clarke cells and of distal dendrites in the nucleus dorsalis: an electron microscopic study in the cat.

The fine structural synaptology of large Clarke cells in L3 has been investigated from a morphometrical point of view in both normal and adult cats which received horseradish peroxidase (HRP) injections in the cerebellum. This marking method made it possible to distinguish small or distal dendrites of large Clarke cells from those of interneurons and the marginal cells of Clarke's column. A total of 1036 boutons was observed on the perikarya of 21 large Clarke cells; 81.9% (848/1036) were small-sized boutons, the cross-sectional areas of which ranged between 0.3 and 2.9 sq. micrometer, while 18.1% (186/1036) were giant boutons ranging between 3.0 and 8.0 sq. micrometer. From 1075 boutons on 17 primary dendrites of Clarke cells, 72.4% (778/1075) were small-sized boutons and 27.6% (297/1075) were giant boutons. From 1679 boutons contacting 366 distal or small HRP-labeled dendrites, 89.9% (1507/1679) were small boutons and 10.1% were giant boutons. The giant boutons were more frequently located on the proximal dendrites than on the cell bodies or small distal dendrites of Clarke cells. The proportion of S- and F-type boutons was different in 3 parts of large Clarke cells. F-type boutons were more frequent on soma (55.0% 570/1036) and primary dendrites (59.4%, 635/1075). S-type boutons outnumbered the F-type on small or distal dendrites (62.6%, 1952/1679). The S/F ratio seemed to increase from the cell body toward the distal dendrites. The results suggest that Clarke cells receive predominantly small S-type boutons since the total receptive area of the dendrites is supposed to exceed that of the cell body.

Animals↗
PubMed1979

Dendritic spikes in Purkinje cells of the guinea pig cerebellum studied in vitro.

Extracellular spikes were recorded simultaneously from dendrites and somata of Purkinje cells in thin cerebellar sections. Spontaneously occurring dendritic spikes were biphasic with the initial phase positive. Triphasic dendritic spikes with a large negative phase appeared during electrophoretic application of glutamate. In media containing procaine, tetrodotoxin, or high concentrations of KCl, negative dendritic spikes occurred whereas soma spikes were abolished. The negative dendritic spikes were suppressed by CoCl2 or MnCl2. Electrical stimulation elicited climbing fibre responses in somata and large negative waves in dendrites. Spikes of dendritic origin were different from those reflecting electrotonic spread of soma spikes. The relation between soma spikes and active dendritic spikes is discussed.

Animals↗
PubMed1978

Gap junctions between dendrites and somata of neurons in the primate sensori-motor cortex.

Gap junctions have been found infrequently between two dendrites or a dendrite and a cell soma in the deep layers of both the motor and somatic sensory cortices of the primate. At these junctions the outer leaflets of the plasma membranes of both profiles are intimately apposed with a gap of 2 nm between them which shows a structure of hexagonal subunits in tangential sections. These gap junctions occur mainly between the dendrites or dendrites and somata of large stellate cells but are also associated in some examples with a dendro-dendritic synapse and thus occur between large stellate dendrites and presynaptic dendrites; a desmosome may also occur in association with a gap junction and dendro-dendritic synapse. Gap junctions have been identified as sites of electrical transmission between cells in a number of sites and it is therefore suggested that some neurons in the sensori-motor cortex are electrotonically couples.

Animals↗
PubMed1979

Postnatal dendritic development in the rabbit visual cortex.

Golgi preparations of rabbit visual cortex aged 1-25 days, as well as similar tissues from adults, were examined for the growth of the dendritic arbor, and in particular the development of dendritic spines. The layer 5 pyramidal neurons and layer 4 stellate neuron were chosen as representatives of larger classes of neurons in the visual cortex. It was determined that the growth of the dendritic arbor, determined by counts of total number of dendritic and total dendritic length, is quite similar for pyramidal and stellate neurons. Dendritic spine development, however, is more rapid in pyramidal neurons than in stellate. This disparity in the rate of dendritic spine development is discussed in the light of physiologic studies on the development of receptive field properties in the rabbit visual cortex.

Age Factors↗
PubMed1979

Distribution of thalamic input to different dendrites of a spiny stellate cell in mouse sensorimotor cortex.

A Golgi impregnated, gold-toned [2] spiney stellate cell from layer IV of mouse SmI cortex was reconstructed in three dimensions from serial thin sections to assess the apatial relationships of the synapses onto its dendrites. The distribution of thalamocortical (TC) synapses with the reconstructed dendrites is presented in this report. Thalamocortical axon terminals were labeled by lesion induced degeneration which, in mouse SmI cortex, may reliably indicate the numbers of thalamocortical axon terminals. Results indicate that thalamocortical synapses, which are distributed over most regions of the dendritic tree, are arranged in a regular, periodic fashion on parts of two of the reconstructed dendrites. In these regions, the necks of spines receiving thalamocortical input attach to the dendrite shaft at intervals of about 5 micrometers. In many other regions of the dendritic tree, two spines receiving thalamocortical synapses are separated by a similar interval. Further studies are expected to determine the extent to which dendrites of spiny stellate cells and of other kinds of cortical neurons are contacted in a periodic fashion by thalamocortical axon terminals.

Animals↗