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114<br />
5. ORIGIN OF SEEDLING...<br />
Perhaps in cooler environments, to which L. perenne is adapted, the effect of herbivory is<br />
not so strong. Because the benefits related to alkaloid-mediated reduction of competition or<br />
herbivory are not realized, the costs of alkaloid production are limited (Faeth 2002). This explanation<br />
seems also plausible for L. persicum. This species is a noxious weed of oat that is<br />
known from allellopatric properties. As a weed it can take advantage from the host alkaloids<br />
not bearing any costs. A serious difficulty in all explanations of fluorescence in the genus Lolium<br />
is unknown physiological role of annuloline. Its structure is clearly different from perloline,<br />
it is only sparingly soluble in water (Axelrod and Belzile 1958) and presumably it does not act<br />
as an osmoregulator. However, cinnamic acid, the parent compound of annuloline can act as<br />
optical filters by blocking UV-A and B (“Cinnamic acid”, 2006). Perhaps UV light penetrating<br />
soil layers could photoactivate fluorescent annuloine to create a defence response. To cite<br />
one example, it is documented that insecticidal activity of fluorescent alkaloids secreted by<br />
Oxalis tuberosa (oca) is linked to photoactivation. The strongest fluorescence is correlated<br />
with resistance to the larvae of Mycrorypes and it is observed in cultivars from the Andean<br />
highlands, which are subjected to a high incidence of UV radiation (Walker et al. 2003).<br />
Regardless fluorescence is a species specific marker or not, it enables to distinguish<br />
between two different functional types i.e., between annual, fast growing Italian ryegrass<br />
and perennial ryegrass. And despite of variation in the fluorescence test it can be used as<br />
a determinant trait in seed certification schemes to track mechanical seed contamination. Because<br />
the majority of L. perenne plants do not fluoresce, the elimination of fluorescent seeds,<br />
irrespective they are L. perenne or L. multiflorum assures that none undesirable plant will be<br />
in a seed sample. This can be a pragmatic way to prevent from low turf quality. However, the<br />
more basic way to distinguish crops or even cultivars is to identify DNA fragments or genes<br />
that are responsible for differences. The three RAPD markers identified in the present work<br />
and associated with fluorescence can be beneficial to developing such DNA tests. These<br />
tests may reduce the problems caused by environmental variation and subjectivity in scoring<br />
results. A striking example of a DNA marker used in separation of different phenotypes is the<br />
SSR marker, sat309 located only 1-2 cM away from the rhg1 responsible for the resistance to<br />
soybean cyst nematode. Genotypic selection with this marker is 99% accurate in predicting<br />
susceptible lines (Babu et al. 2004). The other successful cases include two RAPD markers<br />
tightly linked to a gall midge resistance gene (Gm2) in rice (Nair et al. 1995), two RAPD<br />
markers flanking the locus Lr19 responsible for the leaf rust resistance in wheat (Gupta et al.<br />
2006) and several markers based on single nucleotide polymorphism of coding and adjacent<br />
noncoding regions of Pm3 alleles conferring resistance to powdery mildew in wheat, Triticum<br />
aestivum and spelt, T. spelta (Tommasini et al. 2006). RAPD markers found in the present<br />
work offer a quick and easy way of discrimination between L. multiflorum and L. perenne.<br />
Nonetheless, they suffer some of the same problems as the seedling root fluorescence e.g.,<br />
they are not valid for all cultivars. Much more accurate and flexible test can be developed<br />
provided that DNA markers linked to key morphological characteristics are identified. There<br />
are some morphological characteristics that differentiate L. multiflorum and L. perenne such<br />
as the 1 st year ear emergence and recovery after winter (Chapter 3). Based on the knowledge<br />
about the chromosome locations of QTLs underlying these characters far more DNA tests<br />
can be developed.