Postnatal development of cat hind limb motoneurons. I: Changes in length, branching structure, and spatial distribution of dendrites of cat triceps surae motoneurons

The postnatal development of length, branching structure, and spatial distribution of dendrites of triceps surae motoneurons, intracellularly stained with horseradish peroxidase, was studied from birth up to 44-46 days of postnatal (d.p.n.) age in kittens and compared with corresponding data from adult cats. The number of dendrites of a triceps surae motoneuron was about 12, and the arborization of each dendrite generated an average of 12-15 terminal branches. There was no net change in the number of dendrites of a neuron or in the degree of branching of the dendrites despite the occurrence of both a transient remodeling of the dendritic branching structure and changes of the spatial distribution of the dendritic branches during postnatal development. The perisomatic territory in the transverse plane occupied by the dendritic branches of a motoneuron increased in parallel with the overall growth of the spinal cord. Thus, the relative size of the dendritic territory in this plane was kept almost constant, whereas dendritic branches projecting in the rostrocaudal direction grew much faster than the spinal cord and also became more numerous. At birth the rostro-caudal dendritic span of individual motoneurons bridged 1:6 to 1:5 of the L7 spinal cord segment length; this figure was 1:3 at 22-24 d.p.n. Hence, in this direction, the growing dendritic branches invaded novel dendritic territories. The change in dendritic branch length from birth to 6 weeks of age corresponded to an average growth rate of 2 to 4 microns per dendritic branch and day, which implies that the total increase in length of the dendrites of a neuron could amount to 1 mm/day. The increase in branch length did not occur in a uniform or random manner; instead, it followed a spatiotemporal pattern with three phases: From birth to 22-24 d.p.n., growth was particularly prominent in greater than or equal to 3rd order preterminal and 2nd through 6th order terminal branches. From 22-24 to 44-46 d.p.n., a large increase in branch length confined to terminal branches of greater than or equal to 3rd branch orders was observed. As indicated by topological analysis, this length increase was probably due in part to a resorption of peripheral dendritic branches during this stage of development. From 44-46 d.p.n. to maturity, the increase of dendritic branch length was restricted to preterminal branches of low (less than or equal to 4th) branch order.(ABSTRACT TRUNCATED AT 400 WORDS)     

Morphometric analysis of phrenic motoneurons in the cat during postnatal development.

The dendritic geometry of 20 phrenic motoneurons from four postnatal ages (2 weeks, 1 and 2 months, and adult) was examined by using intracellular injection of horseradish peroxidase. The number of primary dendrites (approximately 11-12) remained constant throughout postnatal development. In general, postnatal growth of the dendrites resulted from an increase in the branching and in the length and diameter of segments at all orders of the dendritic tree. There was one exception. Between 2 weeks and 1 month, the maximum extent of the dendrites increased in parallel with the growth of the spinal cord; however, there was no increase in either combined dendritic length or total membrane surface area. In addition, there was a significant decrease in the number of dendritic terminals per cell (59.8 +/- 9.3 vs. 46.4 +/- 7.4 for 2 weeks and 1 month, respectively). The distance from the soma, where the peak number of dendritic terminals per cell occurred, ranged from 700-900 microns at 2 weeks and 2 months to 1,300-1,700 microns in the adult. The diameter of dendrites as a function of distance from the soma along the dendritic path increased with age. The process of maturation tended to increase the distance from the soma over which the surface area and dendritic trunk parameter (sigma d1.5/D1.5) remained constant. The three-dimensional distribution of dendrites was analyzed by dividing space into six equal volumes or hexants. This analysis revealed that the postnatal growth in surface area in the rostral and caudal hexants was proportionately larger than that in either the medial, lateral, dorsal, or ventral hexants. Strong linear correlations were found between the diameter of the primary dendrite and the combined length, surface area, volume, and number of terminals of the dendrite at all ages studied.     

Postnatal development of cat hind limb motoneurons. II: In vivo morphology of dendritic growth cones and the maturation of dendrite morphology

The maturation of dendrite morphology was studied by light and electron microscopy in cat spinal alpha-motoneurons intracellularly labeled with horseradish peroxidase. Alpha-motoneurons supplying the triceps surae (TS) and the intrinsic foot sole (SP) muscles were investigated in kittens from birth to 44-46 days of postnatal (d.p.n.) age. At birth, a large number of dendritic branches displayed growth cones, filopodia, and fusiform processes. The growth cones were of lamellipodial and filopodial types, but intermediate forms also occurred. The growth cones shared several morphological features with the neuritic growth cones studied in vitro. It was suggested that the occurrence of different types of growth cones--even in the same dendrite--may reflect their transformation from one type to the other and the level of growth activity could be inferred from the number and form of the growth cones. About 50-70% of the terminal branches in the dendrites of newborn kittens possessed growth cones, filopodia, and/or fusiform processes. The corresponding figure for preterminal branches was 20-30%, with a clear decrease in incidence when approaching the soma. During the period under study, most of these growth-associated processes disappeared from the dendrites so that at 44-46 d.p.n. of age only about 10% of the terminal and less than 1% of the preterminal branches had growth-associated processes. Analysis of the three-dimensional distribution of dendritic branches with such processes disclosed that they were relatively more frequent in the medial, rostral, and caudal dendritic territories. It was concluded that the pattern of distribution and disappearance of growth cones, filopodia, and fusiform processes coincided with postnatal longitudinal dendritic growth and the development of the adult dendritic territories described in a preceding paper (Ulfhake et al., '88). Dendritic growth, with respect to length and caliber, also occurred in the absence of growth cones and filopodia. It is suggested that the important role of these processes may be to act as a steering device in establishing the adult distribution and synaptology of the dendrites. Comparison of TS and SP alpha-motoneuron dendrite morphology at birth and at 22-24 d.p.n. age showed that the SP neurons lagged in the maturation process. Light and electron microscopic observations indicated that postnatally direct contacts might exist between dendrites and fine blood vessels in the neuropil without any interposing glial sheath. The number of such suspected contacts diminished during the period under study, indicating that the glial ensheathment of the blood vessel takes place, in part, postnatally.     

Dendritic maturation of displaced putative cholinergic amacrine cells in the rabbit retina

The dendritic trees of Cb, cholinergic, amacrine cells in the ganglion cell layer of the developing rabbit retina are revealed by intracellular injection with Lucifer yellow to have the adult dendritic branching pattern at birth. It is demonstrated that these cells maintain a constant number of dendritic branches throughout postnatal development and that their dendritic trees increase in size by the growth and subsequent elongation of all branches. Proximal and distal dendrites increase in length by almost the same proportions between birth and adulthood. Although the adult pattern of dendritic branching of Cb amacrine cells is established by birth, dendrites in the young possess numerous short appendages (1-5 microns in length) resembling the "dendritic spines" of immature cat retinal ganglion cells. Some of these structures remain on the dendrites of adult cells but the majority are lost at the end of the third postnatal week. As dendritic spines disappear, the dendrites of Cb amacrine cells, especially the distal portion of the tree, acquire numerous varicosities. At each stage after P10, the gain in the number of varicosities greatly exceeds the loss in spines; this is not consistent with the hypothesis that all varicosities are retracted dendritic spines. The rapid increase in the number of varicosities on distal dendrites of Cb amacrine cells during the first 3 postnatal weeks coincides with the maturation of amacrine cell physiological responses. There is no distinct centroperipheral gradient in the postnatal dendritic maturation (acquisition of varicosities, loss of spines, attainment of the adult number of branches) of Cb amacrine cells from the visual streak to the peripheral retina. However, the area of their dendritic tree increases relatively more in the retinal periphery compared to that in the visual streak.     

Postnatal development of cat hind limb motoneurons supplying the intrinsic muscles of the foot sole.

The postnatal development of dendrite anatomy in alpha-motoneurons intracellularly labeled with horseradish peroxidase (HRP), innervating the intrinsic muscles of the sole of the foot (IFS MNs) in the cat, was investigated. The number of dendrites per neuron was about 11 and did not change from birth to adult. The number of branches per dendrite decreased during the same period by 20-25%. The net elimination of dendritic branches appeared to occur at distal branching points, as revealed by topological analysis. The dendritic branching pattern tended to be asymmetric at birth and the net decrease in dendritic branching postnatally did not alter this pattern. The length of preterminal branches (PTB) increased by a factor of 2, while terminal branch (TB) length increased by a factor of 3.3 postnatally. The large increase in TB length was attributed to both longitudinal growth and an apparent lengthening caused by resorption of distal branches during development. Dendritic length in the transverse spinal cord plane increased in parallel with the overall growth of the parent spinal cord segment, while dendritic growth along the rostro-caudal axis exceeded, by about one order of magnitude, dendritic growth in the transverse plane. Average branch diameter doubled from birth to adult. The decrease in branch diameter across branching points did not obey satisfactorily to the 'power rule' of Rall. However, the 1.5 power ratio of daughters-to-parents branch dropped from 1.18 to 1.08 between 3 weeks of age and adult. Tapering was evident in both PTBs and TBs. The rate of taper did not change postnatally. From birth onwards, 'local' branch diameter correlated closely with amount of membrane area and combined length of the dendritic branches located distal to the 'supporting' parent branch. These relations were similar in all age groups and are suggested to be properties intrinsic to the IFS MNs. The local branch diameter also co-varied with the number of distal dendritic branches, but in this case there was a systematic shift in the relationship with increasing postnatal age. It appears that the local diameter in IFS MN dendrites is a key indicator of the size of the distal dendritic arborization.     

Differential growth and remodelling of ganglion cell dendrites in the postnatal rabbit retina

The postnatal dendritic maturation of small field type 1 (SF1), medium field type 1 (MF1) and type 2 (MF2), and large field type 1 (alpha) ganglion cells in the rabbit retina was compared qualitatively and quantitatively. Dendritic tree structure was revealed by intracellular injection of the fluorescent dye Lucifer yellow, and the stained cells were then morphologically separated on the basis of some area, dendritic field size, total dendritic length, number of nodes, and mean internodal distance. Cells in the visual streak and an area inferior to the streak were sampled from retinae between birth and adulthood. The dendrites of all studied classes of rabbit ganglion cells were extensively covered by short spine-like appendages. As in cat retina, many dendritic spines disappeared by the end of the third postnatal week, at which stage the adult dendritic form could be recognised. However, there was differential loss in the number of spines from the dendrites of the four cell classes. In both the streak and inferior retina, adult SF1 cells had the same number of spines/dendritic unit length throughout postnatal life, whereas MF1 and MF2 ganglion cells lost at least half of their number of spines/unit dendritic length by maturity. Alpha ganglion cells lost virtually all their dendritic spines by adulthood. In both retinal locations, there were small changes in the number of nodes (dendritic branch points) of small field and medium field ganglion cells but alpha cells lost between 70 to 80% of their nodes by adulthood. The dendrites of ganglion cells with contrasting morphology thus undergo differential remodelling during postnatal maturation. The completion of the period of dendritic remodelling coincided with the first appearance of adult receptive field organisation, suggesting that structural remodelling, in particular that involving dendritic spines, may be associated with the development of the cell's synaptic circuitry. The dendrites of neighbouring postnatal ganglion cells in the rabbit retina also grow by different amounts; the increase in dendritic tree area, total dendritic length, and mean internodal distances of alpha cells exceeded that of small field and medium field cells in corresponding retinal positions. This implies that retinal dendrites elongate by active growth rather than by "passive stretching."     

Membrane area and dendritic structure in type-identified triceps surae alpha motoneurons

The size and branching structure of the dendritic tree were studied in nine type-identified triceps surae alpha-motoneurons that were labeled intracellularly with horseradish peroxidase and reconstructed from serial sections in the light microscope. The average total membrane area (AN) for motoneurons of type S (slow-twitch) motor units was about 22% smaller than AN for cells of type F units (including both FF and FR motor unit types in this category) (480.1 X 10(3) microns 2 vs. 617.7 X 10(3) microns 2, respectively). Systematic correlations were found between stem dendrite diameter and three measures of dendritic size: dendrite membrane area, combined dendritic length, and number of terminations. All of these correlations were significantly different for the dendrites of F and S motoneurons. Power-function relations between stem diameter and dendritic membrane area were used to estimate AN for a sample of 79 type-identified motoneurons. Mean estimated AN values were significantly different for the F and S motoneuron groups, despite a large overlap in AN values between these groups. The branching structure of dendrites of F and S motoneurons also showed clear differences. Type S motoneuron dendrites showed less-profuse branching and a more-even radial distribution of branch points than found in type F cells. Examination of two forms of the "3/2 power rule" for the relation between the diameters of parent and daughter dendritic branches at branch points showed that the dendrites of type S motoneurons conform less well with the anatomical constraints necessary to represent binary branching trees as equivalent cylinders than do dendrites of type F cells. There was no systematic difference between F and S motoneuron dendrites in the degree of asymmetry of first-order daughter trees. The results overall indicate that the dendrites of F and S motoneuron groups are structurally different, giving rise to a systematic difference in AN between these groups. Such structural differences suggest that the F and S groups of alpha-motoneurons can be viewed as intrinsically distinct cell types and not just large vs. small variants of the same cell species.     

A quantitative analysis of the geometry of cat motoneurons innervating neck and shoulder muscles

The geometry of the somata and dendritic trees of motoneurons innervating neck and shoulder muscles was investigated by using intracellular injections of HRP. In general, these motoneurons did not belong to a homogeneous population of motoneurons. Differences in average primary dendritic diameter, number of primary dendrites, and other measures of dendritic tree size were found between different neck and shoulder motoneuron groups. Several indices of proximal dendritic tree size (number of primary dendrites, sum of dendritic diameters, Rall's dendritic trunk parameter, and the sum of dendritic holes) were weakly correlated with the diameter or surface area of the soma. Some of these correlations depended on the muscle supplied by the motoneuron. The total combined dendritic length ranged from 66,660 to 95,390 microns. There was a weak, but positive, correlation between the diameter of primary dendrites and combined dendritic length. This relationship varied from motoneuron to motoneuron. The diameters of all dendrites of three trapezius motoneurons were examined in detail. The total dendritic surface area examined ranged from 415,000 to 488,100 microns 2 and represented approximately 99% of the total neuronal surface area. Last-order dendrites showed a high degree (39.9%) of taper. Dendritic tapering, by itself, was a major factor in the decrease of the (sum of dendritic diameters)3/2 measured at progressively distal sites from the soma. Although few parent and daughter dendrites obeyed the "three-halves law," the average exponent was 1.57. The diameters of primary dendrites and dendritic surface area were weakly correlated. The correlation between dendritic diameter and combined dendritic length or surface area improved if the weighted average of the diameter of second-order dendrites was used as a measure of dendrite size. Second-order dendrites, whose branches terminated in different regions of the spinal cord, showed different relationships between dendritic diameter and combined dendritic length or surface area. Comparisons between the motoneurons examined in the present study and motoneurons innervating other muscles indicate that, although all spinal motoneurons share several common features (e.g., long dendrites, dendritic tapering), each motoneuron group has a set of unique features (e.g., soma shape, relationship between primary dendrite diameter and dendritic surface area). Thus, the rules governing motoneuron dendritic geometry are not fixed but depend on the species of the motoneuron.     

Postnatal development of peroneal motoneurons in the kitten

In 1- to 72-day-old kittens, motoneurons of the 3 peroneal muscle nuclei were labeled by retrograde axonal transport of horseradish peroxidase from individual muscles. At birth, the locations of peroneal nuclei were similar to those of the adult cat. Counts of motoneurons at different ages indicated that postnatal cell death does not occur in peroneal motor nuclei. Primary dendrites were as numerous in motoneurons of newborn kittens as in adult motoneurons but they were thinner, shorter and poorly ramified. The number of recurrent axon collaterals was higher in the first postnatal week than at later stages. The growth of motoneurons followed similar rates in the 3 peroneal nuclei. Distributions of cell body diameters and volumes were unimodal at birth and became bimodal between 15 and 20 days postnatal. The separation of peroneal motoneurons in two size subgroups, presumably corresponding to alpha and gamma populations, was followed by an increase in growth rate which became faster for alpha than for gamma motoneurons.     

Changes in size and dendritic arborization patterns of adult cat spinal alpha-motoneurons following permanent axotomy.

This study was performed to analyse quantitatively the changes in dimensions and dendritic branching patterns of adult cat spinal alpha-motoneurons following permanent axotomy, i.e., in a situation in which the transected motoraxons are prevented from reinnervating their peripheral target muscle. After transection and ligation of the medial gastrocnemius nerve of adult cats, homonymous alpha-motoneurons were intracellularly labelled with horseradish peroxidase and subjected to quantitative light microscopic analyses. The cell bodies and proximal dendrites were studied at 3, 6, and 12 weeks after the axotomy. An initial increase in cell body size at 3 weeks was followed by a gradual return towards normal values. The mean diameter of the stem dendrites was decreased at all time periods studied, and the combined diameter of the stem dendrites was reduced at 12 weeks after the axotomy. Entire dendritic trees were reconstructed at 12 weeks postoperatively, and the regression equations describing the correlations between dendritic stem diameter, on one hand, and the size of the entire dendrite, on the other, were used to calculate the total dendritic length, volume, and membrane area of whole axotomized motoneurons. The dendritic branching patterns were also analysed. In comparison with normal medial gastrocnemius alpha-motoneurons, the dendritic membrane area and volume of the axotomized cells had decreased by 36% and 29%, respectively, at 12 weeks after the axotomy. This reduction in dendritic size was due to a loss of preterminal and terminal dendritic segments. Abnormal dendritic elongations were observed in 2 of 16 completely reconstructed dendrites.     

Anatomy of dendrites in motoneurons supplying the intrinsic muscles of the foot sole in the aged cat: evidence for dendritic growth and neo-synaptogenesis.

Motoneurons (MNs) supplying the intrinsic muscles of the foot sole (IFS) were studied in the aged cat (greater than 15y). Axon conduction velocity of IFS MNs was 30-40% slower in the aged than in young adult cats. IFS MNs that appeared intact during intracellular recordings and labeling with horseradish peroxidase (HRP) were subjected to anatomical investigation of their dendrites. The results were compared with corresponding data from young adult (less than 3y) cats. The average number of dendrites per IFS MN was twelve in both the aged and young adults. However, the branching was significantly more extensive in the aged cat, thus indicating that proliferation of dendritic branches may occur during the later part of life. Topological analysis revealed a significant difference in the frequency distributions of nodal vertices between young adult and aged cats. In the young adult, the dendritic branching pattern was compatible with trees generated by outgrowth from terminal segments, while in the aged there was a clear indication of collateral outgrowth of branches. The dendritic path distance and the length of terminal branches were similar in young adults and aged. The length of preterminal branches was shorter in the aged, while the combined dendritic length of a dendrite was larger compared to young adults. These data are consistent with the topological data, and add further evidence that the proliferation of branches in the aged cat may also take place from preterminal branches. Light microscopic analysis revealed the presence of "growth cone-like" extensions in the dendrites of the aged cats. Such profiles were not encountered in dendrites from young adults. Electron microscopic observations showed that these "growth cone-like" formations were not artifacts and that they were apposed by numerous axonal boutons, of which a number made synaptic contact. A distinct feature of the extensions was their rich content of mitochondria and membranous elements. It was suggested that these "growth cone-like" formations were sites at which novel synaptic connections are established, and that they may represent the initial stage of an outgrowth of new dendritic branches in the aged cat. Local dendritic branch diameter related closely to the amount of dendritic membrane area located distally in both young adults and aged. Curve fitting disclosed that this relationship was quite similar for both age groups, despite concurrent differences in combined dendritic length and branching degree.     

Axon-bearing amacrine cells of the macaque monkey retina

A new and remarkable type of amacrine cell has been identified in the primate retina. Application of the vital dye acridine orange to macaque retinas maintained in vitro produced a stable fluorescence in the somata of apparently all retinal neurons in both the inner nuclear and ganglion cell layers. Large somata (approximately 15-20 microns diam) were also consistently observed in the approximate center of the inner plexiform layer (IPL). Intracellular injections of horseradish peroxidase (HRP) made under direct microscopic control showed that the cells in the middle of the IPL constitute a single, morphologically distinct amacrine cell subpopulation. An unusual and characteristic feature of this cell type is the presence of multiple axons that arise from the dendritic tree and project beyond it to form a second, morphologically distinct arborization within the IPL; these cells have thus been referred to as axon-bearing amacrine cells. The dendritic tree of the axon-bearing amacrine cell is highly branched (approximately 40-50 terminal dendrites) and broadly stratified, spanning the central 50% of the IPL so that the soma is situated between the outermost and innermost branches. Dendritic field size increases from approximately 200 microns in diameter within 2 mm of the fovea to approximately 500 microns in the retinal periphery. HRP injections of groups of neighboring cells revealed a regular intercell spacing (approximately 200-300 microns in the retinal periphery), suggesting that dendritic territories uniformly cover the retina. One to four axons originate from the proximal dendrites as thin (less than 0.5 microns), smooth processes. The axons increase in diameter (approximately 1-2 microns) as they course beyond the dendritic field and bifurcate once or twice into secondary branches. These branches give rise to a number of thin, bouton-bearing collaterals that extend radially from the dendritic tree for 1-3 mm without much further branching. The result is a sparsely branched and widely spreading axonal tree that concentrically surrounds the smaller, more highly branched dendritic tree. The axonal tree is narrowly stratified over the central 10-20% of the IPL; it is approximately ten times the diameter of the dendritic tree, resulting in a 100 times greater coverage factor. The clear division of an amacrine cell's processes into distinct dendritic and axonal components has recently been observed in other, morphologically distinct amacrine cell types of the cat and monkey retina and therefore represents a property common to a number of functionally distinct cell types. It is hypothesized that the axon-bearing amacrine cells, like classical neurons,     

Physiological changes accompanying anatomical remodeling of mammalian motoneurons during postnatal development

The development of respiratory motoneurons provides unique data that may be generalized to other mammalian motoneuron populations. Like other motoneurons, respiratory motoneurons undergo developmental changes in the shape of the action potential and their repetitive firing. The unique observations concern the postnatal change in the recruitment pattern of cat phrenic motoneurons that is correlated with a halving of mean input resistance, a stasis of growth in the cell membrane and a reduction in the complexity of the dendritic tree. A similar pattern of change was observed for hypoglossal motoneurons studied in rat brainstem slices. Without an increase in total membrane surface area, the decreased resistance must result from a reduced specific membrane resistance. Two mechanisms are proposed to explain this decrease in resistance: proliferation and redistribution of either synaptic inputs and/or potassium channels. Although there was a significant contribution of synaptic input in determining input resistance throughout postnatal development, it was the density of cesium- or barium-sensitive potassium conductances that differentiated low resistance from high resistance motoneurons. Low resistance motoneurons had more cesium- and barium-sensitive channels than their high resistance counterparts. Based on the variations in the relative changes observed in input resistance versus membrane time constant with these two potassium channel blockers (cesium and barium), it is proposed that the distribution of these potassium channels change with age. Initially, their distribution is skewed toward the dendrites but as development progresses, the distribution becomes more uniform across the motoneuron membrane. During postnatal development, the rapid decrease in input resistance results from a proliferation of potassium channels in the membrane and of synaptic inputs converging onto developing respiratory motoneurons while the membrane is being spatially redistributed but not expanded.     

Postnatal development of lamina III/IV nonpyramidal neurons in rabbit auditory cortex: quantitative and spatial analyses of Golgi-impregnated material

We have studied the postnatal development of lamina III/IV spine-free nonpyramidal neurons in the auditory cortex of the New Zealand white rabbit. The morphology and dendritic branching pattern of single cells impregnated with a Golgi-Cox variant were analyzed with the aid of camera lucida drawings and three-dimensional reconstructions obtained with a computer microscope. Sample sizes of 20 neurons were obtained at birth (day 0), postnatal day (PD) 3, 6, 9, 12, 15, 21, and 30 days of age. Normative data were also available from PD-60 and young adult rabbits studied previously. At birth, lamina II-IV have not yet emerged from the cortical plate; immature nonpyramidal neurons at the bottom of the cortical plate (presumptive layer IV) are characterized by short, vertically oriented dendrites. Growth-cone-like structures are present along the shafts and at the tips of the dendrites. At birth, soma area and total dendritic length are, respectively, 34 and 10% of adult values. The cortical plate acquires a trilaminar appearance at PD-3. The six-layered cortex is present by PD-6. During the first postnatal week dendritic length increases fourfold and is accompanied by a significant increase in both terminal and preterminal dendritic growth cones. At the onset of hearing at PD-6, there is a significant proliferation of dendrites and branches to 144 and 200% of adult levels, respectively. These supernumerary dendrites are rapidly lost during the second postnatal week, at which time the somata and dendrites become covered with spines. The loss of higher-order dendrites occurs more gradually; the number of dendritic branches is still 116% of adult values at PD-30. Spine density peaks between days PD-12 and PD-15, and then gradually diminishes until the cells are sparsely spined or spine free by PD-30. Total dendritic length increases in a linear fashion up to PD-15, at which time it is 80% of adult values. An analysis of terminal and intermediate branches demonstrated that the increase in total dendritic length after PD-6 is due entirely to the growth of terminal dendrites. Total dendritic length attains adult levels by PD-30. Spatial analyses revealed that a vertical orientation of dendrites is present at birth. Associated with the onset of hearing at PD-6, there is an explosive elaboration of dendrites toward the pial surface.(ABSTRACT TRUNCATED AT 400 WORDS)     

Androgenic regulation of dendritic trees of motoneurons in the spinal nucleus of the bulbocavernosus: reconstruction after intracellular iontophoresis of horseradish peroxidase

Androgen-sensitive motoneurons in the spinal nucleus of the bulbocavernosus (SNB) in adult male rats were labeled after intracellular iontophoresis of horseradish peroxidase, after which they were fully reconstructed in three dimensions in order to measure their dendritic trees. Three groups of rats were compared: intact adult male rats and male rats castrated as adults and given Silastic tube implants containing either testosterone or nothing. In the high-androgen groups (intact males and testosterone-treated castrates), soma size and the diameter of the first-order dendrites were larger than in blank-treated castrates. Moreover, the terminal dendrites in all groups possessed growth cones, implying that the dendrites of these motoneurons are capable of growth in adulthood. However, there were no statistically significant group differences in the length, membrane surface area, or volume of the dendritic trees, or in the orientation or branching symmetry of dendrites. In general, there were positive correlations between the size of the motoneuronal soma and various measures of the size of the dendritic tree and between the diameter of individual stem (first-order) dendritic branches and the size of remainder of that dendrite. These data suggest that there may be a modest effect of androgen on the size of the dendritic trees of SNB motoneurons in adulthood, although the effect is much smaller than has previously been reported.     

Dendritic plasticity in the early postnatal feline retina: quantitative characteristics and sensitive period

Retinal lesions were made in kittens between 3 and 60 days postnatal age and in adult cats. After postlesion survival times ranging from 4 to 11 months the dendritic morphology of retinal ganglion cells was revealed by retrograde labeling with horseradish peroxidase or with neurofibrillar staining techniques. After retinal lesions on the third postnatal day changes of dendritic morphology were observed in retinal ganglion cells adjacent to regions of retrograde degeneration. Originating from eccentrically positioned somata the dendritic fields extended into the regions that were free of neighboring cells. The dendrites oriented toward the ganglion-cell-free region were elongated and thicker than normal. The density of dendrites per unit area was increased in this part of the dendritic trees. Lesions on the 20th, 38th, and 56th postnatal days elicited increasingly weaker changes of dendritic morphology. The sensitive period for the type of dendritic plasticity described ends between 40 and 60 days postnatally.     

Distribution of dendrites from biventer cervicis and complexus motoneurons stained intracellularly with horseradish peroxidase in the adult cat

The three-dimensional distribution of dendrites from the dorsal neck muscles biventer cervicis (BC) and complexus (CM) was examined in the adult cat using intracellular staining techniques. Motoneurons were electrophysiologically identified, stained with injection of horseradish peroxidase, and reconstructurcted from serial histological sections. The dendritic distributions of all motoneurons examined followed an orderly pattern. Many dendrites extended rostrally and caudally to form a complex parallel collection of dendrites in the ventromedial nucleus. Other dendrites projected dorsolaterally into the spinal accessory nucleus and lateral parts of lamina VII and VIII. Dorsomedial dendrites followed a path parallel to the medial border of the ventral horn and frequently terminated near the central canal. A few scattered dendrites were usually found directly dorsal to the soma in lamina VIII. This pattern of dendritic distribution differed distinctly from the dendritic distribution of motoneurons in other spinal regions. However, in all spinal regions, including the upper cervical spinal cord where BC and CM motoneurons were found, the pattern of dendritic distribution from different motoneurons was similar if their somata were located in the same region. For 15 motoneurons with well-stained dendrites, the mean rostral-caudal extent of the dendritic tree was 2,860 μm. The mean total dendritic length of three of these motoneurons measured 73,100 μm, almost four times larger than hindlimb motoneurons involved in planter reflexes. Despite the large size of the dendritic trees of BC and CM motoneurons, the surface areas of BC and CM cell bodies were smaller than most large hindlimb motoneurons. These quantitative differences in motoneuron dimensions may in turn be reflected by differences in the electrotonic properties of motoneurons in different motoneuron nuclei.     

Theta ganglion cell type of cat retina

We define a new bistratified ganglion cell type of cat retina using intracellular staining in vitro. The theta cell has a small soma, slender axon, and delicate, highly branched dendritic arbor. Dendritic fields are intermediate in size among cat ganglion cells, with diameters typically two to three times those of beta cells. Fields increase in size with distance from the area centralis, ranging in diameter from 70 to 150 microns centrally to a maximum of 700 microns in the periphery. Theta cells have markedly smaller dendritic fields within the nasal visual streak than above or below it and smaller fields nasally than temporally. Dendritic arbors are narrowly bistratified. The outer arbor lies in the lower part of sublamina a (OFF sublayer) of the inner plexiform layer where it costratifies with the dendrites of OFF alpha cells. The inner arbor occupies the upper part of sublamina b (ON sublayer), where it costratifies with ON alpha dendrites. The outer and inner arbors are composed of many relatively short segments and are densely interconnected by branches that traverse the a/b sublaminar border. Experiments combining retrograde labeling with intracellular staining indicate that theta cells project to the superior colliculus and to two components of the dorsal lateral geniculate nucleus (the C laminae and medial interlaminar nucleus). Theta cells project contralaterally from the nasal retina and ipsilaterally from the temporal retina. They apparently correspond to a sluggish transient or phasic W-cell with an ON-OFF receptive field center.     

Development of spiny and aspiny neurons in the caudate nucleus of the dog during the first postnatal month

The adult and developmental morphology of spiny and aspiny neurons in the dog caudate nucleus was examined using the Golgi-Kopsch technique. In the adult, three types each of spiny and aspiny neurons were identified based upon dendritic morphology and cell soma size. They corresponded in large part to those neurons described previously in the caudate nuclei of the rat, cat, and monkey. At birth, dendrites of spiny neurons possessed varicosities, filopodia, and thick proximal dendritic stumps—all characteristic of immaturity. Maturation of these processes involved the thinning of proximal dendrites, lengthening of dendritic shafts, and growth of dendritic spines. Although most of the dendritic maturation occurred during the first postnatal month, spine densities and dendritic lengths of spiny I neurons at 30 days were still less than those seen in the adult. Aspiny I neurons were also immature at birth but lacked the filopodia and thicker proximal dendrites that characterized immature spiny neurons. Aspiny dendritic development involved primarily the lengthening of dendritic processes; by 30 days the aspiny I neurons were indistinguishable from those seen in the adult. These results suggest that dendritic development of spiny I neurons may extend well past the end of the first postnatal month and that studies investigating functional development in the caudate nucleus should consider the relatively extended time period required for maturation of these primary synaptic sites.     

Postnatal dendritic development of Y-like geniculocortical relay neurons.

We describe the dendritic development of neurons in the dorsal lateral geniculate nucleus (LGNd) projecting to cortical area 18 in the postnatal cat. LGN neurons were identified by retrograde labeling from area 18 with fluorescent latex microspheres and injected in the fixed slice with Lucifer yellow (LY) and horseradish peroxidase (HRP) to visualize their dendritic arborizations. Both topological (measures of the patterns of dendritic branching and their territorial coverage) and metric parameters (measures of the quantitative parameters describing the size, length, extent and diameter of the dendritic arbors) were measured in three-dimensions for 25 LGN neurons in cats between 1 and 18 postnatal weeks. In addition, dendritic growth was compared to the changing dimensions of the LGNd. At all ages, neurons projecting to area 18 have large somata and radiate dendrites. From 1 to 18 weeks neurons increase in size--both soma area and the length of all dendritic segments double during this period. Intermediate and terminal dendritic segments show comparable growth until 5 weeks. However, only terminal segments continue to grow significantly from 5 until 18 weeks. Dendrites become straighter during development, the angle between daughter branches decreases and dendritic segment diameter increases, with terminal segments showing a greater increase relative to intermediate segments. The density of dendritic appendages increases transiently at 5 weeks and a differential redistribution occurs, so that by 18 weeks dendrites further from the soma have a greater density of appendages than those near the soma. Some dendritic relationships remain invariant during development--intermediate segments are always shorter, thicker and straighter than terminal segments. During these changes however, area 18 projecting neurons maintain a constant number of primary dendrites and have, on average, a constant branching pattern. The relative volume of the LGNd occupied by an area 18 projecting neuron increases 2.4-fold between 1 and 18 weeks as the dendrites grow with the result that the coverage of a given point of the LGNd by dendrites of area 18 projecting nearly doubles from 24 to 45 neurons per unit volume. This increased net dendritic overlap provides a substrate for enhanced numerical synaptic divergence of the Y-cell pathway from a point source in the retina to the visual cortex.