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Electrophysiological experiments have shown that in hypogranular cerebella the Purkinje cells are innervated by several climbing fibres. The aim of this paper is to provide morphological evidence for this multiple innervation and to describe the topographical distribution of the different climbing fibres onto the somadendritic region of the Purkinje cell. Experiments have been performed in hypogranular adult Wistar rats lesioned during the first postnatal week by methylazoxymethanol (MAM) or by X-irradiation. Purkinje cells were labelled by an anti-calbindin antibody, whereas climbing fibres were visualised by means of Phaseolus vulgaris leucoagglutinin. Purkinje cells showed variable degrees of abnormality and displacement. Climbing fibres made contact with the dendrites of all kinds of Purkinje cells, including those ectopically positioned whose dendrites branched in the white matter. This shows that Purkinje cells can develop dendritic branching in the absence of granule cells and maintain the capability of interacting with their proper afferents, even when they are severely affected and displaced. In four Purkinje cells we have been able to follow the course of two climbing fibre terminal arbourisations. Almost no terminal branches were present around the Purkinje cell soma, and the whole arbour covered the proximal two-thirds of the Purkinje cell dendritic tree. These arbourisations, after an initial common course along the primary dendrite, distributed to separate dendritic regions. The observation of a single labelled climbing fibre covering a limited region of the dendritic tree was more common. As this finding is never observed in control material, it is concluded that the remaining region is covered by another unlabelled climbing fibre belonging to a different inferior olive neurone. These results represent a morphological demonstration of multiple climbing fibre innervation of the adult Purkinje cell. The maintenance of polyinnervation in the adult, which is consequent to the loss of granule cells, is not associated with a defect in the peridendritic translocation of the olivary arbour. In addition, the strict segregation of the different climbing fibres to distinct territories of the Purkinje cell dendritic tree suggests that each terminal arbourisation acts as a functionally independent unit and prevents other competitors from invading its own target domain.
Electrophysiological experiments have shown that in hypogranular cerebella the Purkinje cells are innervated by several climbing fibres. The aim of this paper is to provide morphological evidence for this multiple innervation and to describe the topographical distribution of the different climbing fibres onto the somadendritic region of the Purkinje cell. Experiments have been performed in hypogranular adult Wistar rats lesioned during the first postnatal week by methylazoxymethanol (MAM) or by X-irradiation. Purkinje cells were labelled by an anti-calbindin antibody, whereas climbing fibres were visualised by means of Phaseolus vulgaris leucoagglutinin. Purkinje cells showed variable degrees of abnormality and displacement. Climbing fibres made contact with the dendrites of all kinds of Purkinje cells, including those ectopically positioned whose dendrites branched in the white matter. This shows that Purkinje cells can develop dendritic branching in the absence of granule cells and maintain the capability of interacting with their proper afferents, even when they are severely affected and displaced. In four Purkinje cells we have been able to follow the course of two climbing fibre terminal arbourisations. Almost no terminal branches were present around the Purkinje cell soma, and the whole arbour covered the proximal two-thirds of the Purkinje cell dendritic tree. These arbourisations, after an initial common course along the primary dendrite, distributed to separate dendritic regions. The observation of a single labelled climbing fibre covering a limited region of the dendritic tree was more common. As this finding is never observed in control material, it is concluded that the remaining region is covered by another unlabelled climbing fibre belonging to a different inferior olive neurone. These results represent a morphological demonstration of multiple climbing fibre innervation of the adult Purkinje cell. The maintenance of polyinnervation in the adult, which is consequent to the loss of granule cells, is not associated with a defect in the peridendritic translocation of the olivary arbour. In addition, the strict segregation of the different climbing fibres to distinct territories of the Purkinje cell dendritic tree suggests that each terminal arbourisation acts as a functionally independent unit and prevents other competitors from invading its own target domain.
a b s t r a c tThis paper explores how agricultural technology has interacted with recent land use in the UK and how it might do so in the next 50 years. From 1960 to 1985, farmers successfully used technology to increase the output of crop and animal products per unit of land and particularly of labour. This reduced the number of people employed in agriculture, and promoted larger and more specialised farm enterprises. Between 1985 and 2006, food prices were relatively low, and although labour productivity continued to increase, land productivity remained relatively static. However during this period, farmers started to address the effects of agriculture on reduced water quality and habitat loss.For established agricultural products, technological innovation tends to have an incremental effect, working through genetic improvement, the removal of abiotic and biotic stress (e.g. crop nutrition and protection, irrigation and drainage, and animal nutrition, health and housing) and the substitution of labour. Whereas the first two processes tend to be scale-neutral, the substitution of labour is usually easiest to achieve on larger farms. Other key areas for technological innovation include addressing air, soil and water quality, biodiversity, waste reduction, and information management. Over the next 50 years, large step-changes in land use arising from agricultural technology are predicted to arise from the development of new markets for agricultural products. A strong bioenergy sector will strengthen the links between crop commodity and energy prices and will have a major effect on future land use. Climate change and the regulation of greenhouse gas emissions will alter the relative profitability of crop and animal production systems. Lastly, increased public awareness of the links between food, health and the environment could substantially shift the demand for specific agricultural products.Continual improvements in agricultural technology are pivotal to providing society with the flexibility to balance the challenges of improving human well-being with the management of the planet's ecosystem. Increased technological innovation increases the probability that agricultural land can be used for other purposes, but the exact relationship is dependent on trade and environmental policies. The large external effects of agriculture mean that decisions regarding the adoption of future technologies should be taken by farmers working with other stakeholders.
Synapse elimination is considered to be the final step in neural circuit formation, by causing refinement of redundant connections formed at earlier developmental stages. The developmental loss of climbing fiber innervation from cerebellar Purkinje cells is an example of such synapse elimination. It has been suggested that NMDA receptors are involved in the elimination of climbing fiber synapses. In the present study, we probed the NMDA receptor-dependent period of climbing fiber synapse elimination by using daily intraperitoneal injections of the NMDA receptor antagonist MK-801. We found that blockade of NMDA receptors during postnatal day 15 (P15) and P16, but not before or after this period, resulted in a higher incidence of multiple climbing fiber innervation and caused a mild but persistent loss of motor coordination. Neither basic synaptic functions nor cerebellar morphology were affected by this manipulation. Chronic local application of MK-801 to the cerebellum during P15 and P16 also yielded a higher incidence of multiple climbing fiber innervation. During P15-P16, large NMDA receptor-mediated EPSCs were detected at the mossy fiber-granule cell synapse, but not at the parallel fiber-Purkinje cell or climbing fiber-Purkinje cell synapse. It is therefore likely that the NMDA receptors located at the mossy fiber-granule cell synapse mediate signals leading to the elimination of surplus climbing fibers. These results suggest that an NMDA receptor-dependent phase of climbing fiber synapse elimination lasts 2 d at most. During this phase, the final refinement of climbing fiber synapses occurs, and disruption of this process leads to permanent impairment of cerebellar function.
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