3systems these investigations have followed up whole signal chains (including receptors anddownstream components), in most cases single components were functionally analysed to pinpoint pathogenicity – related cascades. The data obtained so far allow some generalconclusions:
There are several examples for signal pathways involved only in pathogenicity; i.e.deletions of the corresponding genes do not effect vegetative properties
Single components (like the different G
subunits or MAPK) are highly conserved, evenhighly homologous to mammalian systems.
The components of a given signal chain might differ considerably between fungi.
The same (or highly homologous) components can be members of cascades regulatingdifferent downstream components (see the results of MAPK knockouts in several fungi).In the following a few selected aspects of this field of research, which has developedrapidly into one of the major foci of molecular phytopathology, will be presented.
Fungi undergo specific differentiation and developmental processes in response to distinct physical and chemical environmental signals. All these events start with an initial"recognition phase" in which specific receptors play an important role by detecting surfacecomponents or other ligands and transmitting this information to one or more downstreamsignaling pathways. So far only one fungal gene encoding a pathogenicity-relatedtransmembrane receptor protein,
, was described. It was identified by a REMI-mutagenesis approach in the rice blast fungus
1999).The REMI mutant tagged in the
gene was almost fully apathogenic (drastic reductionof appressoria formation). The predicted secondary structure of Pth11p suggested that it is anintegral membrane protein; this was confirmed by
localization experiments using aPTH11-GFP-gene fusion. Eukaryotic serpentine receptors have typically seventransmembrane domains (Bockaert and Pin 1999), whereas Pth11p appears to have nine,suggesting an atypical structure. Exogenous cAMP suppressed defects associated with
mutants, suggesting that Pth11p mediates cellular response through the cAMP pathway.
2.2 Heterotrimeric GTP-binding proteins (G-Proteins)
The importance of heterotrimeric G proteins in regulating diverse processes such asdifferentiation, mating, and pathogenicity has been demonstrated in a number of phytopathogenic fungi (a recent compilation in Tudzynski and Tudzynski 2001 lists 7species). In most cases two or more G
subunit genes were detected, only one of which hadsignificant influence on pathogenicity e.g.,
from the chesnut blight fungus
(Liu and Dean 1997).The defects linked to G
knockouts are manifold: e.g., in
the conidia fail to germinate, demonstrating the requirement of this G
- subunit for avery early stage in the life cycle of this pathogen (Truesdell
mutants of thenorthern corn leaf blight fungus
show reduced ability to formappressoria on glass surfaces and corn leaves, but nevertheless are able to induce lesions; inaddition, CGA1 appears to be involved in mating: mutants are female sterile (Horwitz
1999). In the gray mould fungus
, werefunctionally characterized (Tudzynski
2000). Both genes are expressed
at veryearly stages of infection. Knock-out-mutants of both genes caused wild type-like primarynecrotic lesions in the first hours of infection on bean and tomato leaves. However, after twodays, no further development was observed for the lesions caused by the
-mutants produced spreading secondary lesions, albeit retarded.