Remodelling during Development of the Purkinje Cell Dendritic Tree in the Mouse
The development of the Purkinje cells in normal C57 mice was studied from 7-100 d post natum. The growth of the dendritic trees was analysed both metrically and topologically using the method of vertex analysis (Berry & Flinn 1983a). Granule and Purkinje cell counts were made so that Purkinje cell segment production could be correlated with the number of parallel fibres deposited. Both topological and metrical results indicate that from 7 to 30 d post natum the Purkinje cell dendritic trees expand massively; accounting for 87% of total segment elaboration, reaching their lateral boundaries by 12-15 d post natum and then advancing towards the pial surface. Continued lateral expansion is constrained by the proximity of dendrites from neighbouring trees. Growth proceeds upwards through the neuropil as a front of prolific random terminal branching with inhibitory forces acting at the edges of the growth corridor and behind the growth front to prevent overlapping of dendrites. By 30 d post natum all boundaries are reached and the size of the dendritic field is fixed. Trees averaged 711.2 segments ± 21.45 with a mean distance from root to terminal segment of 133.5 ± 2.9 μ m. The Va/Vb vertex ratios and the levels of trichotomy during this period indicate that branching patterns deviate from pure random terminal additions in a dichotomous tree. There is opportunity for non-random growth at the areas of inhibitory action. Beyond 30 d post natum remodelling occurs within the arbor which involves segment loss in the subpial region (orders above 16) and segment elaboration within the tree (orders 8-16) causing increased density of dendrites and overlapping of segments. The frequencies of segments and terminals are restored to symmetrical distributions through the orders of the trees from the skewed distributions associated with the frontal advance in earlier growth. During remodelling the Va/Vb vertex ratios and percentage of trichotomous nodes are consistent with growth through dichotomous random terminal branching. Path lengths of 8 μ m between each order are seen as regular increments throughout entire trees at 100 d post natum. The final tree produced is indistinguishable from a network grown entirely by random terminal dichotomous branching with some 6% trichotomy and a Va/Vb vertex ratios of 0.92. Granule cell number within the granular layer increases rapidly up to 15 d post natum after which cell death causes a decrease to stable levels beyond 30 d post natum. Purkinje cell number is constant throughout the entire study. However, there is no correlation between the numbers of granule cells (parallel fibres) per Purkinje cell and the number of Purkinje cell branches elaborated beyond early development.
Proceedings of the Royal Society of London Series B
- Pub Date:
- May 1984