Breeding and Cultivation of Japanese Species of Genus Coix As a Fodder Crop
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Breeding and Cultivation of Japanese Species of Genus Coix as a Fodder Crop By MICHIO MURAKAMI Faculty of Agriculture, Kyoto Prefectural University Juzudama and polyploidization with an aim Introduction of developing perennial fodder coix with high yielding potential. The genus Coix, believed to be originated in India and Southeast Asia, is a relative of Experimental results maize which belongs to Maydeae, and is one of the five oriental genera of Tripsaceae, i.e. The experiment was carried out at the Coix, Trilobachne, Polytocci, Sclemchne and Kyoto Prefectural University during a period Chionachne. Coix was cultivated in India in from 1955 to 1976. Results are summarized ancient days as indicated in Vedic literature as follows: (B.C. 1500-B.C. 800). It is now widely dis tributed over tropical, subtropical, and tem 1) Geogravhical ·vari<ltions in perate zones as wild species, and is cultivated morphological characte1·s in some districts. Seven species of Coix have Many ecotypes (hereafter referred to lines) been identified so far, based on the shape of of Hatomugi and Juzudama were collected leaves and seeds, although no generally ac from all parts of Japan to examine their mor cepted taxonomy has been established yet. In phological characters. Juzudama showed a C. Lacry1na-Jobi L, seven varieties have been greater variation in plant height than Hato reported so far. Studies on the genus Coix mugi, ranging from 72 to 181 cm as compared have been confined only to morphological and to 115 to 148 cm in the later. Juzudama is cytological aspects,. ,1,'ls>. shorter in plant height at higher latitudes, In Japan, two major types of Coix have while such a tendency was not observed with been known, i.e. Hatomugi ( C. M<l-yuen Ro Hatomugi. J uzudama exceeded Hatomugi in man, C. Lcicrym<l-Jobi L. var. Ma-yuen Stapf. number of tillers. At lower latitudes, Hato etc.) and Juzudama (C. Lacryma-Jobi L). mugi headed earlie1· than Juzudama, but at Hatomugi is an annual type and its seeds higher latitudes some lines of Juzudama have been used as drug and fodder, while headed earlier than Hatomugi, although early Juzudama, a perennial wild type, is grown maturing types of Hatomugi were also observ under unfavorable environments, especially ed. ill-drained conditions. Though it is not clear Variations within a line ( coefficient of vari when Juzudama was introduced to Japan, it ation) in plant height, number of tillers and has been known since fairly early days as a heading date indicated that Juzudama has wild plant, while some old documents indicated significantly smaller coefficients with increas that Hatomugi was introduced from China at ing latitude, especially for plant height. Such the beginning of the 19th century. geographical or ecological differences between The author carried out experiments on the wild and cultivated species of Coix in Japan interspecific crossing between Hatomugi and seem to be caused by the difference in the date 57 of introduction and the way of utilization13>. of pollen mother cells of the Ft plants 82% of the total number of cells examined showed 2) Pollen morphology 10 II, whereas 14% showed 9 II +2 I, and a As pollen morphology is a major field of few cells 4 I and 6 I. The occurrence of uni palynology, the surface structure of pollen valent chromosomes mostly due to the E chro membrane was compared with electron micro mosome is regarded the cause of the reduced scope among Hatomugi, Juzudama, F1 hybrids pollen fertility in F1 hybrids5>. between them, tetraploid Coix, and corn (Zea Mays L) with an aim of knowing the kinship 5) Segregation in F2 povulation of these plants. It was found that the sexine Early heading and late heading plants were of pollen membrane is covered by fine spinules, segregated with a ratio of 52 to 285, which and there is no recognizable difference in the leads an assumption that the time of heading surface structure among these plants, except is determined by 3 genes. Plant height show a slight difference in size and shape of spine, ed a trimodal distribution consisted of paren and a considerable difference in the structure tal and F1 modes (Fig. 2), indicating that of germ pores of pollens. This result suggests plant height is determined by a few genes with a considerable similarity among the species relatively large effectiveness. Penotypic and of Coix, as well as between Coix and Zea·1 >. genetic correlations between plant height and o~).ler characters showed that plant height has S) Characters of F1 plants hi°k h positive conelations with heading date, Seed fertility of Hatomugi and Juzudama leaf length, plant weight, and pollen fertility, was more than 90% in natural pollination, and but is negatively correlated with numbe1· of their selfing rate was proved to be as high as tillers (Fig. 3). As tall plants were all late 85% by isolated cultures. Interspecific cross maturing, the breeding for an early maturing ability between them was fairly high, showing tall type would be difficult. Winter-survival 43.1 and 50.3% in reciprocal crosses. Signifi ability was apparently greater with late cant heterosis was recognized in many char maturing tall plants than early-maturing acters of F 1 hybrids, especially with plant short plantsG>. height, number of tillers, leaf length and plant weight. In addition, the F1 plants showed par 6) F3 lines ticularly good regrowth after cutting, in terms From 43 F2-plants taken at random F:1 of plant height and number of tillers. How lines were derived. Mean plant heights of F:1 ever, the pollen and seed fertilities of the F1 lines ranged from 110 to 250 cm continuously. plants were only 60% and 30% respectively. Coefficient of variation of plant height was All the F1 plants survived winter in open smaller in tall lines, giving the correlation of field, and their growth in the next year was - 0.823 between plant height and the coeffi as good as in the first year. These facts cient of variation. Phenotypic and genetic cor suggest the usefulness of the F1 hybrid as a relations between plant height and some other fodder crop2>. characters were calculated by the analysis of variance and covariance. Plant height was 4) Cytogenetical features negatively correlated with number of tillers, Cytogenetical study was carried out to find but was positively correlated with heading out the cause of the decreased pollen fertility elate (Table 1). in the F1 hybrids. Both Hatomugi and Juzu Heritability of characters was estimated by dama have 20 somatic chromosomes with using correlation and regression between F2 karyotype formulae shown in Fig. 1. The individuals and F:1 lines and further by the shape of E chromosome is clearly different analysis of variance. Heritability estimated between the two species, and in the meiosis by correlation for plant height and for head- 58 JARQ Vol. 13, No. l, 1979 A Un11111111u111111 · Uuu11111111nn11 A I I I I I I I I I Length (11) IG. ,I 5. 3 ·1. 7 4. 5 4. 4 3. 9 3. 7 3. 7 3. 5 3. 3 Karyotype(<)= 2'·"'A••+213'" + 2C•• + 2"1>"' + 2"'t"' + 2F1m + 2F,"' + 2f',•• + 2G" + 211" B I I I I 3. 2 I I 4.0 3.4 3.5 (11 ) I 3.8 Length IG. 3 I5. 4 4.I 9 4. 2 4. 5 Karyotypc(<)~ 2'·"'A"'+2U•• + 2C"' + 2<D•• + 2~;m Fig. I. Somatic chromosomes and karyotype of Hatomugi (Coix Ma yuen Roman.) and Juzudama (Coix Lacryma-Jobi L.) A: Hatomugi, B: Juzudama 20 15 >. Q C "'::, <:r ~ 10 tt.. H 5 'I •I ' ' 1• ----, ' 0 ... - ' 100 150 200 250 Plant height (cm) --- C.M. - F2 (C.M. X C.L.) ··-• ···· C.L. ----- F2 (C.L. X C.M.) Fig. 2. Frequency distributions of plant height of Hatomugi, Juzudama and F2 populations C.M.: Hatomugi, C.L.: Juzudama 59 Fz (C. M. xC. L.) 0.214° Number of - 0.301·· F -0. 374 - Plant height, 0. 237 --Pollen fertility tiller's -0. 015 -0.009 ~ I 0. 763 .. 0. 725° '-0. 095. 0. 522 .. 0. 841 0.741 0.097 0. 532 0.211 0. 658 0. 130~ -0. ~18 I Heading date Leaf length rSeed fertility I 0.599.. 0.699° 0.630 0.831 0. 455 0.407 / I ~P-la_n_t _w_e_i_g_h_t I Leaf width F1 (C. L. x C. M.) 0.211•• Number of -0.162* -0. 199 - - Plant hei~ht 0. 236 ····- Pollen fertility tillers - 0.015 - 0.009 I I 0.752.. ~ 0. 8,11 .. ~ -0. 008 0.487•• 0.806 0. 843 - 0.015 0.494 /,,,......0.211 0.658 0. 1:10 -0.018 .------~ I I Heading date Leaf length Seed fertility I 0. 705 .. 0. 698 .. 0.736 0. 845 0.455 0.407 I Plant weight Leaf width Fig. 3. Phenotypic, genetic and environmental correlation coefficients between main characters of F2 population Each set of three figures indicates phenotypic, genetic, and environmental correlation coefficients in that order. * Significant at the 5% level ** Significant at the 1% level Table 1. Phenotypic, genetic and environmental correlation coefficients between plant height and some other characters of F3 lines Correlation Plant height-No. of tillers Plant height-Heading date Plant height-Seed fertility rp -o. 6421** 0.8799** o. 1688 •G - o. 8931 0.9477 o. 1845 'E - o. 4948 0.4374 0.2274 'P, 'G and 'E show phenotypic, genetic and environmental correlation coefficients respectively ** Significant at the 1% level. 60 JARQ Vol. 13, No. 1, 1979 ing date was very high, i.e. more than 0.8 Tsble 2.