Involvement of polyamines in root development

Root development is under the control of hormonal, metabolic, and environmental cues that can act on genetically-controlled developmental programmes and thus affect the plasticity of root architecture. These processes involve not only the five 'classical' plant hormones, but also other growth regulators, such as polyamines. The present review emphasises the importance of polyamines in the different aspects of root development: primary root growth and lateral and adventitious root formation. Free (agmatine, putrescine, spermidine, spermine), conjugated (such as hydroxycinnamate conjugates) and macromolecule-bound polyamines are reported to be present in root systems. Modifications of their endogenous levels by inhibitor treatment, by mutation, by gene manipulation, or by exogenous treatment can have drastic effects on root development and subsequent architecture. These effects may be related to the involvement of polyamines in the control of cell division and differentiation, which plays an important role in the root apex and during lateral and adventitious root formation. The exact mechanisms of action remain to be elucidated, but accumulating evidence in plant and animal cells supports the idea that, besides biophysical effects on membranes and nucleic acids, polyamines interact with protein kinases and transcription factors and are thus involved in signal transduction pathways. The high flexibility of polyamine metabolism in response to environmental stress and the metabolic link between polyamine and ethylene synthesis strongly suggest that polyamines may play a role in environmentally-induced plasticity of root development. Moreover, polyamines may be implicated in the establishment of biotic interactions between roots and rhizospheric micro-organisms.