Veinte años de Encuentros - Encuentros en la Biología

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Invierno 2012-2013
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Salvador Guirado
Catedrático de Biología Celular de la Universidad de Málaga
Primer Editor y Director de la revista Encuentros en la Biología
[email protected]
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Vol.5 ¦ Nº 141
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Vol.5 ¦ Nº 141
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