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Karyological Studies on Iris Japonica Thunb. and Its Allies1) 2) by K

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Karyological Studies on Iris Japonica Thunb. and Its Allies1) 2) by K 180 Cytologia 10 Karyological Studies on Iris japonica Thunb. and Its Allies1) 2) By K. Yasui Tokyo Imperial University (With 8 text figures) Receic ed MTV 28, 1939 Introduction There are several karyological investigations on Iris japonica. KAZAO (1928) considered the sporophyte of this species as an auto triploid having 54 chromosomes. But SIMONET (1932, 1934) re ported I. japonica as a diploid plant with 34 chromosomes in its root-tip cells, and considered I. japonica var aphrodite as a hyper triploid and interpreted the chromosome constitution as 2n=51+3, the latter 3 chromosomes being considered as derived from 3 of 51 chromosomes in I. japonica by fragmentation which occurred att their median constrictions. The karyological investigation in I. japonica and the other 2 allied species, which were kindly put at my deposal by Dr. S. IMA MURA of Kyoto Imp. University, gave me some results different from those of the previous authors as is described below. Material and Method In the fixing of the root-tip cells both NAVASHIN's and LEWITSKY's solutions were used. When fixed in the latter, the chromosomes were found longer and slender than when fixed in the former solution, but there was no essential difference in the distribu tion of chromosomes in the equatorial plate. NEWTON'S gentian violet staining was generally adopted. Materials for the study of meiosis were fixed mostly with NAVASHIN'S fluid and stained with HEIDENHAIN'S iron-alum haematoxylin. Acetocarmine smears were also tried in the study of the pollen mother cells. Drawings were made with a camera lucida. 1) Contributions from the Divisions of Genetics and of Plant-Morphology, Botanical Institute, Faculty of Science, Tokyo Imperial University, No. 231. 2) This work was made possible by grants from The Japan Society for the Advancement of Cytology. 1939 Karyological studies on Iris japonica Thunb . and its allies 181 Observations A. Iris japonica Thunb. I. Meiosis in. PMC. There were observed 2-5 trivalents, mostly 14 bivalents, and mostly 17, sometimes nearly 17 univalents in a PMC with 54 chromosomes (Figs. 1 and 2). Three constituent chromo somes of one of the trivalents had a long slender constriction region between the short and long arms always closely associated wth their short arms (Figs. 1, a, and 2 A, a) in the diaphase, while the rest of the trivalents showed the end to end association (Fig. 2 A, b). Several chromosomes showed intimate connections with nucleoli (Fig. 2 D). These connections remained until the first meiotic metaphase, and very often the relic nucleolus was seen in the spindle and also in the cytoplasm. Figs. 1-5. Iris japonica. 1, diaphase nucleus of PMC. a, trivalent chromosome; n, nucleolus. 2, some chromosomes in diaphase of a PMC; n, nucleolus; 2 A, 2 trivalents; 2 B, different configurations of 11 bivalents; 2 C, 2 univalents; 2 D, a vacuolated nucleolus (n) and 6 attached chromosomes. 3, a PMC in which triva lents and bivalents (in black) are going to separate at the equator of the spindle, and the univalents (in outline) are scattering around them; some univalents behind the tri and bi-valents are not shown. 4, anaphase of 1st meiotic division in PMC, Drosera type. 5, chromosomes in a diaphase nucleus of a MMC; 5 A, 2 trivalents, a, of the same shape with that of 2 A, a; 2 B, 14 bivalents; 5 C, 20 univalents. ca. •~1300. 182 K. YASUI Cytolegia 10 In the metaphase these trivalents, as well as the bivalents, formed the equatorial plate earlier than the univalents which were found scattered around them (Fig: 3). In the anaphase these trivalents generally separated into two sets, one with one chromo some and another with 2 chromosomes, and each set proceeded to the pole, together with those of the separated halves of bivalents. Generally the univalents, which came later into the equator, migrated at random into both poles without separation of their chromatids (Fig. 4). Retarded chromosomes forming miniature nuclei or ex tra-nuclear chromosomes were observed. Cytokinesis did not occur after the 1st meiotic division. Second meiotic division in the PMC is rather regular, though some few retarded chromosomes were found also. The miniature nuclei of the 1st division generally divided into two. After the 2nd division a simultaneous cytokinesis occurred. Table 1. Karyotype of Iris japonica Thunb. Denotations: L, very long; 1. long; M, median: m, short median: S. short: s. very short. 2. Meiosis in the megaspore mother cell (MMC). The behavi our of the chromosomes in the meiosis of the MMC resembled generally that of the PMC; few trivalents and the compensating number of the bivalents and the univalents (Fig. 5), miniature nuclei, extra-nuclear chromosomes in the 1st and 2nd divisions were observed.. But in one case in the 1st division almost all of the univalents divided at the 1st metaphase and followed the separated trivalent and bivalent chromosomes into both poles. This may be due 1939 Karyological studies on Iris japonica Thunb. and its allies 183 to the different cellular conditions in the PMC and the MMC, which influence the behaviour of the univalent chromosomes. 3. Chromosomes in the root-tip cells. The chromosome number in the root tip cells of I. japonica is 54 as KAZAO (1928) reported (Fig. 6). We can recognize in them 3 component sets of chromo somes. Two of them resembled each other in their size and struc ture, but the third was of a different type, though a few chromosomes of the third group resemble those in the other 2 sets. The determina tion of the karyotypes in such a species as I. japonica, which has rather numerous and various kinds of chromosomes, is not easy. Moreover, due to the small number of the observations the karyotype shown in Table 1 may not be conclusive, but it shows the general features. Either three nucleoli, of which 2 were larger and one smaller, or one large nucleolus were found generaly in the resting or interkinetic nuclei of the root-tip cells. B. Iris sp. (Chinese origin) This species has 36 chromosomes in its root-tip cells (Fig. 7). Two nucleoli of equal size or one large nucleolus were found in the resting or interkinetic nucleus of the root-tip cells. Karyotype structure of the chromosomes are shown in Table 2. Table 2. The karyotype of Iris sp. (Chinese origin) Denotations same as In Table 1. 184 K. YASUI Cytolcgia 10 C. Iris formosana Ohi This species has 28 chromosomes in the root-tip cells. Compared with the other two species, however, it has a very different chromo some constitution (Fig. 8). Two smaller nucleoli of equal size or Figs. 6-8. Equatorial plates in root-tip nuclei. 6, Iris japonica; 7, Iris sp. (Chinese origin); 8, Iris formosana. ca. •~1270. one larger nucleolus were found in the resting and interkinetic nuclei in the root-tip cells. The karyotype diagnosis is shown in Table 3. Table 3. Karyotype of Iris formosana Ohi Denotations same as in Table 1. 1939 Karyological studies on Iris japonica Thunb. and its allies 185 Discussion 1. Derivation of I. japonica Thunb. "Syaga" (Japanese name) which was described by Thunberg (1794) as I. japonica is a wild indigenous plant in this country, being distributed from Kyusyu to northern Homo and Sikoku, but not found in Hokkaido, Ryuky(i, Taiwan, and Tyosen. An allied plant, "Kotyokwa", was already known in China, which does not grow wild in Japan. I. japonica growing in Japan has 54 somatic chromosomes irr its root-tip cells. My observation in the meiosis of the PMC a: well as of the MMC show that 1) there are very few trivalents, 2) about one third of the chromosomes remain as univalents, and 3) the remaining, mostly 14 pairs of the chromosomes, are asso ciated as bivalents. From these data and the chromosome be haviour in the meiosis, it seems that the species in question is not an autotriploid as was considered by KAZAO at that time, but an allotriploid, which was derived from a hybrid between 2 specie; having different karyotypes, and in which the chromosome set of one parent has been doubled. The chromosome studies in the root tip cells gave sufficient evidence; namely the chromosomes can be divided into 3 sets, 2 of them much resembling each other, while the third has only a few chromosomes resembling those in the other 2 sets. Consequently SIMONET's I. japonica can not be identified with I. japonica ("Syaga"), because the former is a diploid plant having 17 haploid chromosomes. I. japonica var. aphrodite having 54 somatic chromosomes (SIMONET, 1934) might be a variegated I. japonica ("Syaga") probably being mutated vegetatively but is not the direct derivative of SIMONET's I. japonica. My Chinese material is a diploid plant having 36 somatic chro mosomes in its root-tip cells. This number may suggest an intimate relation of this species to I. japonica, but its karyotype does not exactly coincide with either of the karyotypes of I. japonica. Another allied species, I. formosana, which is known to grow only in a limited area in the southern part of Formosa, is also a diploid plant having only 28 chromosomes. Not only the karyotype but also the chromosome number is different from those of other species. Thus we cannot find out the parent plant of I. japonica among those diploid species. I. gracilipes is a diploid plant (KAZAO, 1928) which has 36 somatic chromosomes, but its morphological characters are so different from those of I. japonica and other allied diploid species as to permit the suggestion that it has direct relationship with I.
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