Combined Dendritic Length Research Articles

Dendritic morphology of CA1 pyramidal neurones from the rat hippocampus: I. Branching patterns.

The aim of this study was to provide quantitative descriptions of the dendritic branching patterns of pyramidal neurones in the CA1 region of the rat hippocampus. Thirteen adult cells were filled with biocytin and reconstructed by using the light microscope. The number of basal trees arising from the soma of each cell ranged from two to eight. There was wide variation in the number of terminal segments per tree. Six cells had single apical trunks, and seven had trunks that bifurcated in stratum radiatum. The number of apical oblique trees ranged from nine to 30, with each tree usually showing a lower degree of branching than basal trees. Basal and oblique trees had similar branching patterns, with the majority of branch points occurring close to the origin of the tree. Both basal and oblique terminal segments were generally much longer than intermediate segments and constituted up to 90% of the combined dendritic length of the tree. The branching pattern of the apical tuft was different, with many relatively long intermediate segments; terminal segments contributed only some 66% of the combined dendritic length of these trees. The mean total combined dendritic length for six adult cells reconstructed and measured completely was 11,900 +/- 1,000 microns (standard deviation). The relative proportions of the different parts of the dendritic system, although not the total dendritic length, were correlated with the location of the soma relative to the cell body layer. Cells with somata close to the stratum pyramidale/stratum radiatum border had more dendrites terminating in stratum radiatum and fewer in stratum oriens than cells with somata further from it.

Morphology of developing rat genioglossal motoneurons studied in vitro: relative changes in diameter and surface area of somata and dendrites.

This study describes the postnatal change in size of motoneurons in the hypoglossal nucleus that innervate the genioglossus muscle. Such anatomical information is essential for determining the cellular mechanisms responsible for the changes observed in the electrical properties of these motoneurons during postnatal development. The cells analyzed here are part of an earlier study (Núñez-Abades et al. [1994] J. Comp. Neurol. 339:401-420) where 40 genioglossal (GG) motoneurons from four age groups (1-2, 5-6, 13-15, and 19-30 postnatal days) were labeled by intracellular injection of neurobiotin in an in vitro slice preparation of the rat brainstem and their cellular morphology was reconstructed in three-dimensional space. The sequence of postnatal dendritic growth can be described in two phases. The first phase, between birth (1-2 days) and 13-15 days, was characterized by no change in either dendritic diameter (any branch order) or dendritic surface area of GG motoneurons. However, maturation of the dendritic tree produced more surface area at greater distances from the soma by redistributing existing membrane (retracting some terminal branches). During the second phase, between 13-15 days and 19-30 days, the dendritic surface area doubled as a result of an increase in the dendritic diameter across all branch orders and a generation of new terminal branches. In contrast to the growth exhibited by the dendrites, there was little discernible postnatal growth of somata. At all ages, dendrites of GG motoneurons show the largest amount of tapering in the first-and second-order dendrites. The calculated dendritic trunk parameter deviated from a value 1.0, indicating that the dendritic tree of developing GG motoneurons cannot be modeled accurately as an equivalent cylinder. However, the value of this parameter increased with age. Strong correlations were found between the diameter of the first-order dendrite and the dendritic surface area, dendritic volume, combined dendritic length, and, to a lesser extent, the number of terminal dendrites in GG motoneurons. Correlations were also found between somal and dendritic geometry but only when data were pooled across all age groups. These data support earlier studies on kitten phrenic motoneurons, which concluded that postnatal growth of motoneurons was not a continuous process. Based on the fact that there was no growth in the first 2 weeks, the changes in the membrane properties described during this phase of postnatal development (e.g., decrease in input resistance) cannot be attributed to increases in the total membrane surface area of these motoneurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Morphology of developing rat genioglossal motoneurons studied in vitro: Changes in length, branching pattern, and spatial distribution of dendrites

The aim of this study is to describe the postnatal change in dendritic morphology of those motoneurons in the hypoglossal nucleus that innervate the genioglossus muscle. Forty genioglossal (GG) motoneurons from four age groups (1-2, 5-6, 13-15, and 19-30 postnatal days) were labeled by intracellular injection of neurobiotin in an in vitro slice preparation of the rat brainstem and were reconstructed in three-dimensional space. The number of primary dendrites per GG motoneuron was approximately 6 and remained unchanged with age. The development of these motoneurons from birth to 13-15 days was characterized by a simplification of the dendritic tree involving a decrease in the number of terminal endings and dendritic branches. Motoneurons lost their 6th-8th order branches, in parallel with an elongation of their terminal dendritic branches maintaining the same combined dendritic length. The elongation of terminal branches was attributed to both longitudinal growth and the apparent lengthening caused by resorption of distal branches. The elimination of dendritic branches tended to increase the symmetry of the tree, as revealed by topological analysis. Later, between 13-15 days and 19-30 days, there was a reelaboration of the dendritic arborization returning to a configuration similar to that found in the newborn. The length of terminal branches was shorter at 19-30 days, while the length of preterminal branches did not change, suggesting that the proliferation of branches at 19-30 days takes place in the intermediate parts of terminal branches. The three-dimensional distribution of dendrites was analyzed by dividing space into six equal volumes (hexants). This analysis revealed that GG motoneurons have major components of their dendritic tree oriented in the lateral, medial, and dorsal hexants. Further two-dimensional polar analysis (consisting of eight sectors) revealed a reconfiguration of the tree from birth up to 5-6 days involving resorption of dendrites in the dorsal, dorsomedial, and medial sectors and growth in the lateral sector. Later in development (between 13-15 days and 19-30 days), there was growth in all sectors, but of a greater magnitude in the dorsomedial, medial, and dorsolateral sectors.

The Extent of the Dendritic Tree and the Number of Synapses in the Frog Motoneuron.

Frog motoneurons were intracellularly labelled with cobaltic lysine in the brachial and the lumbar segments of the spinal cord, and the material was processed for light microscopy in serial sections. With the aid of the neuron reconstruction system NEUTRACE, the dendritic tree of neurons was reconstructed and the length and surface area of dendrites measured. The surface of somata was determined with the prolate - oblate average ellipsoid calculation. Corrections were made for shrinkage and for optical distortion. The mean surface area of somata was 6710 microm2; lumbar motoneurons were slightly larger than brachial motoneurons. The mean length of the combined dendritic tree of brachial neurons was 29 408 microm and that of lumbar neurons 46 806 microm. The mean surface area was 127 335 microm2 in brachial neurons, and 168 063 microm2 in lumbar neurons. The soma - dendrite surface area ratio was 3 - 5% in most cases. Dendrites with a diameter of </= 1.0 microm constituted approximately 75% of the combined dendritic length in most of the neurons. Unlike in the cat, there was no correlation between the size of stem dendrites and the extent of daughter branches. From the synaptic density estimated in earlier electron microscope investigations of frog motoneuron dendrites (Antal et al., J. Neurocytol., 15, 303 - 310, 1986; 21, 34 - 49, 1992), and from the present data, the number of synapses on the dendritic tree was calculated. The calculations indicated 26 949 synapses on the smallest and 61 519 synapses on the largest neuron if the synaptic density was multiplied by the length of the dendritic tree. If the synaptic density was multiplied by the surface area of the dendritic tree the calculation yielded 23 337 synapses for the smallest and 60 682 synapses for the largest neuron. More than 60% of the combined surface area of dendrites was >600 microm from the soma. This suggests that about two-thirds of the synapses impinged upon distant dendrites >600 microm from the soma. The efficacy of synapses at these large distances is investigated on model neurons in the accompanying paper (Wolf et al., Eur. J. Neurosci., 4 1013 - 1021, 1992).

Restorative effects of reinnervation on the size and dendritic arborization patterns of axotomized cat spinal alpha-motoneurons.

In a preceding paper [Brännström, et al. (1992) J. Comp. Neurol. 318:439-451] a marked reduction in dendritic size was observed in cat spinal motoneurons following permanent axotomy. The aim of the present study was to analyse the possible restorative effects of peripheral reinnervation on the size and dendritic branching patterns of cat spinal motoneurons which had been deprived of neuromuscular contact for an extended period of time. In adult cats the medial gastrocnemius (MG) nerve was transected and ligated. After 6 weeks the nerve was allowed to reinnervate its muscle through a nerve graft. With approximately 6 weeks needed for muscle reinnervation [Foehring, et al. (1986) J. Neurophysiol. 55:947-965], the MG motoneurons were devoid of neuromuscular contact for altogether about 12 weeks. Two years later reinnervated MG alpha-motoneurons were intracellularly labelled with horseradish peroxidase to allow quantitative analyses of the cell bodies and dendritic trees. Comparisons were made with previous data from normal and permanently axotomized MG motoneurons. The reinnervated motoneurons exhibited positive correlations between dendritic stem diameter, on one hand, and combined length, volume, membrane area, and number of end branches of the whole dendrite, on the other. By using the regression equations for these correlations, the total dendritic size of whole reinnervated motoneurons could be estimated. Such calculations showed that in comparison with the reduction in dendritic size found at 12 weeks after permanent axotomy (Brännström et al., see above), peripheral reinnervation caused the dendritic volume and membrane area to return to normal values. However, the values for combined dendritic length and number of dendritic end branches were still reduced by more than 25% as compared to the normal situation. The results indicate that following reinnervation of the target muscle, the axotomized motoneurons did not recover their original number of dendritic branches. The normalization of dendritic membrane area and volume was instead accomplished by two other mechanisms, namely an increase in dendritic diameters and an increased number of dendrites per neuron.

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.

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.

Anatomy of soleus ?-motoneurone dendrites in normal cats and in cats subjected to chronic postnatal tenotomy or overload of the soleus muscle

The anatomy of intracellularly HRP-labeled soleus alpha-motoneurone dendrites was studied both in normal adult cats ("normal soleus", NS) and in adult cats which at a postnatal age of 5-7 days had been subjected to chronic tenotomy of either the soleus muscle ("tenotomized soleus", TS), or all the soleus synergists contributing to the achilles tendon ("overloaded soleus", OS). A set of "structural rules" seemed to govern the architecture of normal soleus alpha-motoneurone dendrites. Thus, the dendrites branched dichotomously and the number of daughter branches originating from a preterminal branch was proportional to the diameter of that parent branch. Branch diameter decreased across branching points according to the "3/2 power rule" of Rall (1959). Branching occurred down to a preterminal branch diameter of about 0.8 micron. Through all branch orders there existed a quite precise relation between the diameter of a preterminal branch and the membrane area of its distal dendritic arborization. The average dendritic path distance from soma to termination was not closely related to the diameter of the stem dendrite, since thick stem dendrites rather generated more profusely branched arborizations than thin stem dendrites. As a corollary of these characteristics close relations existed between the dendritic stem diameter on one hand, and the total number of branches, combined dendritic length, total dendritic membrane area and total volume, on the other. In the OS material, the dendrites were not different from those of normal soleus motoneurone dendrites. In the TS material, the dendrites were less branched and had greater dendritic path lengths, although the relations between various size-parameters within the dendrites were not significantly altered compared with normal dendrites. It was concluded that the change in branching pattern was due to a net elimination of dendritic branches following the muscle tenotomy.

Postnatal development of cat hind limb motoneurons. III: Changes in size of motoneurons supplying the triceps surae muscle.

The postnatal changes of neuronal dimensions were studied in cat triceps surae motoneurons intracellularly labeled with horseradish peroxidase. Systematic correlations were observed in the analysis of single dendrites at each studied stage, from birth to 44-46 days post natum (d.p.n.) age, between size parameters intrinsic to the dendrites as the diameter of a 1st-order dendrite, the combined dendritic length, the dendritic membrane area, and the degree of branching. Some variability among samples was evident in each studied age group. The correlations were, however, sufficiently close to permit indirect estimations of both combined dendritic length and dendritic membrane area for larger samples of neurons from data on dendritic stem caliber. The total postnatal increase in dendritic membrane area was, on the average, 400%, i.e., from close to 100 X 10(3) microns2 to about 500 X 10(3) microns2. The corresponding increase in soma area amounted to 100%. Analysis revealed that there was a time lag between the increase in somatic and dendritic size. Thus, adult somatic dimensions were attained at age 44-46 d.p.n.; however, at this stage, the mean total dendritic membrane area was only about half of the adult value. The postnatal increase in size appeared to vary among neurons, yielding a wider neuronal size spectrum in the adult cat than that observed in kittens. The measured increase in size corresponded to a calculated average addition of dendritic membrane area of 3700 microns2/day from birth to 22-24 d.p.n. and from that stage to 44-46 d.p.n. of 2700 microns2 per day. Likewise, the increase in combined dendritic length could initially be as large as 1 mm/day down to 0.4 mm/day between 22-24 and 44-46 d.p.n., with a mean growth during the first 44-46 d.p.n. of 0.5 to 0.6 mm/day. The ratios of daughters to parent branch diameters (sigmadd1.5: dp1.5) and the dendritic trunk parameter (sigma d1.5) recorded along the proximodistal dendritic path distance revealed transient changes that might impact on the electrotonic properties of the dendrites during postnatal development. Computations from the measured changes in dendritic branch lengths and calibers indicated that if membrane and internal resistivity remain unaltered during postnatal development, the dendritic domain is electrotonically more compact in the newborn kitten than in the adult cat.(ABSTRACT TRUNCATED AT 400 WORDS)

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.

Light microscopic observations on cat Renshaw cells after intracellular staining with horseradish peroxidase. II. The cell bodies and dendrites.

The cell bodies and dendritic trees of five lumbosacral Renshaw cells of adult cats were studied in the light microscope (LM) after intracellular injection with horseradish peroxidase (HRP). The cell bodies were all located in the ventral part of lamina VII. The dendrites extended up to 0.7 mm from the cell body into the neighbouring parts of laminae VIII and IX as well as into more dorsal parts of lamina VII. The dendritic branching was sparse and about half the dendrites were unbranched. The mean diameter of the cell body was positively correlated to both the combined and mean diameters of the first-order dendrites. Between four and eight dendrites originated from the cell bodies. The number of dendritic end-branches, the combined dendritic length, the mean dendritic length from the cell body to the termination of the end branches, the distance from the cell body to the termination of the most remote end-branch, the dendritic surface area, and the dendritic volume all correlated positively with the diameter of the parent first-order dendrite. The dendritic tapering was somewhat more pronounced in the Renshaw cells than previously observed in alpha- and gamma-motoneurons. The present data are discussed in relation to previous morphological observations on Renshaw cells and alpha- and gamma-motoneurons.

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.

Quantitative analysis of the dendrites of cat phrenic motoneurons stained intracellularly with horseradish peroxidase.

All the dendrites (N = 37) generated by four phrenic motoneurons were analyzed following intracellular injection of horseradish peroxidase. The dendritic arbors produced from each of these stem dendrites were studied in detail. The mean number of stem dendrites produced by a phrenic motoneuron was 9.7, their mean diameter was 6.0 micron, and their mean combined diameter was 58.3 micron. The length at which a phrenic motoneuronal dendrite terminated was 1,236 micron, with several end terminals extending more than 2 mm from the cell body. The mean value for the combined lengths of all segments originating from a single stem dendrite was 5.3 mm. A full spectrum of dendritic branching patterns was observed from simple (five unbranched) to complex, the latter producing up to ninth-order branches. Most terminal and nonterminal dendritic segments tapered, producing a mean diameter reduction of 34%, or approximately 9% per 100-micron length. All phrenic motoneurons exhibited a steady decrease in the combined dendritic parameter (sigma d3/2) with distance from the soma as a result of tapering and end-branch termination. The mean surface area and volume of a phrenic motoneuronal dendrite were 35.3 X 10(3) micron 2 and 25.9 X 10(3) micron 3, respectively. The dendrites constituted greater than 97% of the total phrenic motoneuronal surface area, with 75% of this area lying outside of a 300-micron radius from the cell body. The diameter of a stem dendrite was positively correlated with its combined dendritic length, number of terminal branches, dendritic surface area, and volume. Despite this strong correlation, the value of total dendritic surface area calculated using the power equation derived from the dendritic surface area versus stem dendritic diameter plot was not a consistent estimator of the total dendritic surface area directly measured for these four phrenic motoneurons. It is suggested that this inconsistency may be the result of a heterogeneity in the phrenic motoneuronal population and/or in the dendrites projecting to the different terminal fields.

Morphology of lumbar motoneurons innervating hindlimb muscles in the turtle Pseudemys scripta elegans: an intracellular horseradish peroxidase study.

Motoneurons in the turtle lumbar spinal cord, electrophysiologically identified as innervating a muscle belonging to a functional group, were injected with horseradish peroxidase by electrophoresis. A total of 45 motoneurons were reconstructed from transverse sections. Eleven motoneurons were identified as innervating knee extensor muscles, eight as innervating hip retractor and knee flexor muscles, 14 as supplying ankle and/or toe extensors, and 12 as innervating ankle and/or toe flexor muscles. The cell bodies were elongated and spindle-shaped in the transverse plane. The mean equivalent soma diameter was calculated to be 33.4 micrometers. The mean axon conduction velocity was 15.7 m/second. Significant, though rather weak, positive correlations were found between soma diameter, axon diameter, and axon conduction velocity. The axons of the reconstructed motoneurons did not reveal a recurrent axon collateral. However, a few unidentified motoneurons did possess such collaterals. The dendritic trees were restricted to the ipsilateral side of the cord, but reached out in lateral, ventral, and ventromedial directions to the subpial surface. Easily recognizable and characteristic dendrites were found both in the dorsal dendritic tree and in the dorsomedial dendritic tree. Correlations were calculated between the soma diameter and (1) the number of first-order dendrites, (2) the mean diameter of the first-order dendrites, and (3) the combined diameter of the first-order dendrites. In each case no correlations or only weak correlations were found. Fair correlations were observed between the diameter of a first-order dendrite and the number of terminal dendritic branches (r = .61) and the combined dendritic length (r = .78). However, correlations between the combined diameter of all first-order dendrites per neuron and the total number of terminal dendritic branches and the total combined dendritic length of a neuron were extremely weak. The overall appearance of turtle spinal motoneurons is comparable to that observed in other "lower" vertebrates such as frog and lizard. However, similarities are also observed between certain morphometric parameters in turtle and cat lumbar motoneurons.

A quantitative light microscopic study of the dendrites of cat spinal gamma -motoneurons after intracellular staining with horseradish peroxidase.

By use of intracellular staining with horseradish peroxidase (HRP), the dendritic systems of spinal gamma-motoneurons of the adult cat were studied with a light microscope. The dendrites extended in various directions up to 1.5 mm from the cell body. The dendritic branching was sparse and even unbranched dendrites were occasionally seen. The number and combined diameter of the first-order dendrites increased in parallel with the mean cell body diameter. The number of dendritic end branches, the combined dendritic length, the membrane surface area, and the volume of the entire dendrite correlated positively with the diameter of the parent first-order dendrite. In comparison with the alpha -motoneurons (Ulfhake and Kellerth, '81) the gamma -motoneurons had smaller values for mean cell body diameter and mean diameter of the first-order dendrites and they also had a smaller number of first-order dendrites. The dendrites of the gamma-motoneurons were also found to have fewer branching points and larger values for combined dendritic length. The relation between the diameter of the first-order dendrite and the surface area of the entire dendrite was almost identical for the two types of motoneurons.