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9 Interactions Between Epiphyllic Microorganisms and Leaf Cuticles 147<br />

Fig. 2. A scheme of the phyllosphere<br />

as a habitat for<br />

microorganisms showing most<br />

of the relevant climatic and<br />

<strong>plant</strong> parameters determining<br />

the living conditions of the leaf<br />

<strong>surface</strong><br />

atmosphere<br />

waxes<br />

cutin<br />

epidermal<br />

wall<br />

wetting<br />

biofilm<br />

penetration<br />

leaching<br />

(Andrews 1992; Jacques and Morris 1995; Fiss et al. 2000). However, looking<br />

at the leaf <strong>surface</strong> as a microhabitat with very specific boundary conditions,<br />

investigations of the parameters, limitations and interactions between the<br />

lipophilic leaf <strong>surface</strong> and the microorganisms have rarely been carried out<br />

(Fig. 2). This habitat is characterised by an extreme microclimate due to<br />

large variations in climatic parameters like light intensity and temperature<br />

(Andrews and Harris 2000). Due to specific physical, chemical and biological<br />

properties of leaf <strong>surface</strong>s, the phyllosphere is also dominated by a low<br />

availability of water and nutrients (Schönherr and Baur 1996; Beattie and<br />

Lindow 1999). Investigating the microbial ecology of the phyllosphere will be<br />

a combined approach including <strong>plant</strong> ecophysiological and microbiological<br />

tools. In the following, several important aspects of the microbial ecology of<br />

the phyllosphere will be discussed and selected examples for the interactions<br />

occurring between epiphyllic microorganisms and the leaf <strong>surface</strong> will be<br />

given.<br />

2 Physical and Chemical Parameters of the Phyllosphere<br />

water<br />

vapor<br />

gradient<br />

The <strong>plant</strong> cuticle covering the leaf <strong>surface</strong> is a lipophilic, extracellular<br />

biopolymer. It is composed of the cutin polymer (Kolattukudy 2001), which is<br />

a polyester of esterified hydroxy fatty acids, and of cuticular waxes (Walton<br />

1990), deposited as monomeric compounds to the cutin polymer (intracuticular<br />

waxes) and to the cutin <strong>surface</strong> (epicuticular waxes). Cuticular waxes are<br />

basically linear long chain aliphatic compounds of different chain length and<br />

different substance classes. Typical wax constituents are alkanes, aldehydes,<br />

primary and secondary alcohols, acids and esters composed of the respective<br />

acids and alcohols (Bianchi 1995). Besides these linear long-chain aliphatics,

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