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Cajal's Achievment in the Field of Development of Dendritic Arbors Pablo Garcia Lopez

Cajal's Achievment in the Field of Development of Dendritic Arbors Pablo Garcia Lopez

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Garcia-Lopez et al., 2009 1
 TitleCajal´s achievements in the field of Development of Dendritic ArborsFull names of authors
Pablo Garcia-Lopez, Virginia Garcia-Marin, Ricardo Martinez-Murillo, and Miguel Freire.
Institutional affiliation of all authors
Instituto Cajal, CSIC. Ave. Dr. Arce 37, 28002 Madrid, Spain
Running title
 Cajal and development of dendritic arbors
Keywords
 Cajal, development, dendritic arbors, dendritic spines.
List of frequently used abbreviations and symbolsComplete postal address
*
Correspondence to: Virginia Garcia Marin,Instituto Cajal (CSIC) Avda. Doctor Arce, 3728002 Madrid (Spain)e-mail: vgmarin@cajal.csic.es Tel.: +34 91 585 47 34Fax: +34 91 585 47 54
 The email addresses of all authors
Pablo Garcia-Lopez: caravaca1@gmail.comRicardo Martinez-Murillo: r.martinez@cajal.csic.esMiguel Freire: mfreire2@gmail.com
 
Garcia-Lopez et al., 2009 2
 ABSTRACT 
In 1909 Cajal published an up-dated version in French (Cajal, 1909-1911) of his main work 
Textura del Sistema Nervioso del Hombre y los Vertebrados 
(Cajal, 1899-1904), considered the mostimportant book devoid to the nervous system. Owing that this year is the centenary of thispublication, we have decided to divulge an article focused on Cajal´s description of the morphologicalchanges that undergo dendritic trees during development. We will emphasize his brilliant hypothesesexplaining the modelling of dendritic trees (the neurotropic hypothesis and the role of neuronalactivity in the patterning of the dendritic trees), and the status of this topic in present day Neuroscience. Many cellular and molecular mechanisms, have been discovered in recent years, andsignificant advances have been made in our understanding of activity-dependent plasticity. Here, we will show photographs taken from Cajal´s slides housed in the Cajal´s Museum (Instituto Cajal, CSIC)illustrating the principal changes in neuronal morphology at different stages of development of thespinal cord, cerebellum and cerebral cortex. We will also discuss Cajal’s initial proposals regarding theinfluence of neurotropic substances (chemoatactic hypothesis) and neural activity in the modelling of the dendritic tree, as well as the evidences that later confirmed these theories.
 
Garcia-Lopez et al., 2009 3
1. INTRODUCTION
Cajal´s contributions to the Nervous Systems represent the roots of modern Neuroscience. This immense and exceptional body of work carried out mainly with the Golgi method, wassummarized by Cajal in his
opus magnum 
book 
Textura del Sistema Nervioso del Hombre y los Vertebrados 
(Cajal, 1899-1904). Ten years later, a French larger and updated translation of this work entitled
Histologie du Système Nerveux de l’Homme et des Vertébrés 
(Cajal, 1909-1911), allowed Cajal to publish agreater exposure of his findings. Actually, a centenary of this publication is hold and we felt theopportunity to review some of the important facts on dendritic development that were discussed inhis book and that are valid to date.Dendritic morphology is a critical determinant of neuronal function since the patterns of dendrite arborization determine the kinds of input that neurons receive. The dendritic arborization islargely established during development through a combination of an intrinsic genetic program, growthpromoting signals that regulate the size of the arbor, chemotropic cues that guide dendrites to theirproper location, and neurite-neurite contacts that ensure proper representation of the dendritic fieldand the establishment of the appropriate synaptic contacts (Parrish et al., 2007). These developmentalsignals may interact to create the definitive arborization of the dendritic tree, which will then only beslightly modified during adulthood.Cajal was one of the first scientists to describe the morphological changes in the dendritic treeduring development. One of the principal sources of Cajal´s successes was to apply the
ontogenic method 
 to study the nervous system. Instead of studying the adult nervous system, he decided to work withembryos, young animals or simple animals, thereby reducing the complexity of the structures. Thisapproach had additional important advantages, both improving the consistency of the results obtained with the Golgi method (Golgi, 1873) and making possible to nerve fibers to be visualized. Embryonicaxons are mostly unmyelinated which permitted their stain following the Golgi method. By applying the ontogenic and Golgi methods, Cajal described the successive steps in the development of neuronsand how they connect with their targets. In his opera magna,
“Textura del Sistema Nervioso del Hombre y los Vertebrados” 
1
(Cajal, 1899-1904), after the chapters dealing with the spinal cord, cerebellum and
1
 Translated from the french version “Histologie” into english by:Neely Swanson and Larry Swanson (1995).
Histology of the Nervous system of Man and Vertebrales 
. New York, Oxford Universty Press.Pedro Pasik and Tauba Pasik (1999).
Texture of the Nervous System of Man and Vertebrales 
. SpringerWien, New Cork, SpringerBarcelona.Other Cajal´s works translated into english are: Javier DeFelipe and Edgard G. Jones, (1991).
Cajal´s Degeneration and Regeneration of the Nervous System 
. Translated by Raoul M.May. Edited by DeFelipe and Jones. Oxford Universty Press.

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