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6. DO ANY SPECIES BOUNDARIES....<br />
on AFLP or SSR maps. Furthermore, several gaps are usually found. For example, the AFLP<br />
map derived from p150/112 contains five gaps (Bert et al. 1999) while as many as five chromosomes<br />
are deficient of SSR markers on the map based on the F 2<br />
population derived from<br />
contrasting cultivars of L. perenne (Gill et al. 2006).<br />
Certainly the uniform distribution of markers on the present map resulted from the application<br />
of various techniques. These findings raise the more fundamental point that a well<br />
saturated genetic map can only be constructed based on several marker types. A vast of different<br />
markers ensures that various types of genomic sequences are matched. Undoubtedly,<br />
the application of transposon-based markers is one of the major improvements preventing<br />
from marker clustering. Successful transposable elements insert in both plant and animal<br />
species in the centric heterochromatin as well as into the others transposons. Although the<br />
former are tagged to some extent by AFLP markers, the latter are rarely recognised due to<br />
high accumulation of LTR sequences. However, both regions can be successfully filled by<br />
transposon based markers. The low efficiency of AFLP markers to discover clusters of transposons<br />
is indirectly confirmed by their more frequent clustering with RAPD markers than with<br />
transposons.<br />
Difficulties in alignment of all maps based on a single p150/112 population but employing<br />
a single marker category in a given experiment illustrate well that high quality genetic<br />
map can only be produced if various markers are used at once. The most dramatic example<br />
includes the SSR and AFLP maps, both constructed in p150/112 by the same authors (Jones<br />
et al. 2002a; b) but both with different linkage group assigning and moreover any attempts to<br />
align both maps have not been undertaken. Similarly, at least two maps are based on the F 2<br />
population derived from a cross between two cultivars of L. perenne, Perma and Aurora. The<br />
first employs RFLP (Armstead et al. 2002) while the second SSR (Gill et al. 2006). Unfortunately,<br />
no consensus map is created. Each map with different marker categories is surely<br />
advantageous over those mono-markers and it has greater utility both in QTL mapping and<br />
evolutionary studies. Even fewer markers ensure quite uniform genome coverage (Hayward<br />
et al. 1998; Warnke et al. 2004). Employment of different marker categories is of the highest<br />
importance if a map has to be used for evolutionary considerations. For example, among<br />
all markers employed, those based on transposons (SSAP) proved to be the most powerful<br />
tool for mapping of species boundaries. Strong directional distortions demonstrate the extent<br />
and nature of evolutionary processes that would be difficult to deduce from the other marker<br />
categories also because all others segregate according to the expected Mendelian ratios<br />
typical of intraspecific crosses. Furthermore, mapping of insertion sites demonstrates clearly<br />
that distorted regions are predominantly associated with transposons. Transposon markers<br />
are among the most unique features of the present maps.<br />
6.4.3. Location of selected traits on the map of L. multiflorum and L. perenne<br />
One of the most obvious outcomes of genetic mapping is the location of genes of interests.<br />
In comparison with major crops including rice, maize or barley, the knowledge about genetics<br />
of morphological characters in Lolium is scarce. Only recently some traits have been<br />
mapped including flowering time, self-incompatibility locus and few enzymatic loci (Yamada<br />
et al. 2005). However, existing information is relatively underdeveloped. The current genetic