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52<br />

3. MORPHOLOGICAL VARIATION...<br />

different environments in addition to the lack of reproductive barriers - all these argue for taking<br />

them as a single entity.<br />

The history of crops has proved that many of more dramatic differences in plant morphology<br />

are due to differences in one, or just a few genes and many have been induced by<br />

mutagenic treatment. Some of these mutants fall outside the accepted morphological limits<br />

not merely of the species, but of the genus. However, the classification keys rarely take them<br />

into account that resulted in a kind of “unnatural” classification. Crosses between these different<br />

forms readily give rise to true-breeding intermedium forms as it is observed in barley<br />

after crossing six-rowed with two-rowed cultivars or in Lolium as a result of hybridization<br />

between L. multiflorum and L. perenne. The latter is often classified as Lolium x hybridum.<br />

As such, it is intermediate in many traits; it is less winter hardy but higher yielding than perennial<br />

ryegrass. Regularly producing new species by crossing different forms seems absurd. In<br />

barley, for example, following a re-classification all mutants, ecotypes and cultivated forms<br />

were included in a single entity Hordeum vulgare (Briggs 1978). Such approach is also presented<br />

for L. multiflorum and L. perenne in the Integrated Taxonomic System of the USA<br />

(2007). This seems especially justified by human activities, which have probably caused the<br />

observed diversification of both taxa. This conclusion is supported by the distinction of L. multiflorum<br />

cultivars from the remaining populations. They are characterized by extremely high<br />

values for generative characters. Italian ryegrass is primary used as an annual forage crop<br />

to provide high quality grazing for dairy cattle. Therefore, it should be easy to establish, grow<br />

rapidly and produce high yields through the summer (Cosgrove 1999). The demand for improvements<br />

through breeding programmes directed to rapid grow, high quality forage and die<br />

after heading has resulted in lower winter hardiness, greater number of heading plants in the<br />

1st year, longer spikes and greater number of spikelets in L. multiflorum cultivars in comparison<br />

with ecotypes. Most importantly, these predicted increases may be small but could pose<br />

a serious challenge to taxonomists. Cultivars represent the transgressive segregation and as<br />

such they fall easily outside the species mean. If selection of perennial ryegrass is directed<br />

into opposite direction i.e., to more persistent, later flowering cultivars, the separation of<br />

L. multiflorum and L. perenne may be produced artificially. This scenario seems plausible as<br />

confirmed by the lack of differences between both species at the ecotype level.<br />

Several authors argued that breeding activities have increased the level of hybridization<br />

between L. multiflorum and L. perenne and resulted in huge amount of introgressive forms<br />

in nature (Bennett et al. 2002). This further will cause the morphological unification of both<br />

species. However, the results from this study do not support this view. Taking into account<br />

higher similarity between ecotypes of both species in comparison with cultivars, it can be<br />

speculated that differences between them have been produced recently by genetic manipulations<br />

during breeding programmes especially in L. multiflorum. It should be noted that in<br />

the most distinct cultivars some unusual characters were observed - thick stems, very wide,<br />

stiff and rough leaves and branched spikes. The introgression from Festuca species can not<br />

be therefore, excluded. Introgression is a backcrossing procedure, which allows a limited<br />

number of alien genes to be introduced from a donor species into the reconstituted genome<br />

of the recurrent species by recombination and selection. The diversity of demands on grasslands<br />

agriculture has led to increasing effort in widening the Lolium gene pool through the<br />

exploitation of the close relationship between Lolium and Festuca. Although less digestible

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