中国苹果属植物染色体研究

本文报道了中国苹果属Malus Mill．21种36类型的核型和6个多倍体类型的减数分裂联会构 型。 (木+多)(木+衣)海棠组Sect．Docyniopsis和花楸苹果组Sect．Sorbomalus中的滇池海棠系Ser． Yunnanenses  属2A型，陇东海棠系Ser．Kansuenses中的变叶海棠M．toringoides和花叶海棠M． transitoria属2A、2B则，三叶海棠系Ser．Sieboldianae属2B型，苹果组Sect．Malus中的苹果系 Ser．Pumilae和山荆子系Ser．Baccatae除小金海棠M．xiaojinensis 2B型外，均属3B型。减数分 裂观察结果有同源四倍体，节段异源三倍体和异源三倍体类型。本文还讨论了种类间在核型上的差异、栽培种的起源，核型的演化趋势以及无融合生殖类型的分类学处理。

Chromosome Studies of Chinese Species of Malus Mill.

The genus Malus Mill. of the Maloideae of the Rosaceae consists of about 35 recognized species, which are distributed in temperate region of the Northen Hemisphere including Asia,  Europe and North America.  Redher's classification divided the genus Malus into 6 series, 5 sections, of which 5 series, 3 section, including 27 species, occur in China. The chromosomal counts on 22 Chinese species were reported previously (Chen et al.,  1986;   Liang,  1987). However, no comprehensive karyotype analyses have been made for most species reported. Analyzed in this paper are karyotypes of 36 forms, 22 species, and synapsis configuration of 6 polyploid forms in Chinese Malus to explore the relationships among species and provide cytological evidence for delimitation of species in Malus. Chromosome classification follows Li and Chen (1985). The materials used are listed in Table 1 and the vouchers are deposited in our depart- ment. The cytological results are shown as follows (see the data in Table 1, Plates 1-- 8 and Fig. 1).      1  Sect. Malus      (1) Ser. 1 Baccata  The series consists of nine species endemic to China, 7 of which were cytologically observed: i.e. Malus baccata Borkh. 2n=34;  Sooo1-3 of M. baccata Borkh. 2n= 34;  M . mandshurica Kom. 2n= 34;  M. rockii Rehd. 2n=34;   Sooo 3-2 of M.  rockii Rehd.  2n=51;   M.  sikkimensis Koehne. 2n=34;  M. hupehensis Rehd.  2n=51;   Sooo4-4 of M.  hupehensis Rehd. 2n=51;  M. halliana Koehne.  2n=34 and M.  xiaojinensis Cheng et Jiang 2n=68. The karyotypes consist of sm (8- 9  homologous chromosomes) and m (6- 7  homologous chromosomes)  as well  as st (1-2 homologous chromosomes). The chromosomes range in relative length from 9.84 to 3.74, with the ratio of the longest to the shortest from 2.28 to 2.03 and the mean ratio arms from 2.08 to 1.86. All the karyotypes belong to Stebbins's 3B except 2B of M.  Xiaojinensis.  Pollen  mother  cell (PMC)  of M.  hupehensis  contains 17I+17II, and the average synapsis configuration per cell is 2n=3x=51=17. 70I+ 15. 87II+ 0.52III, while PMC meiosis of M. xiaojinensis shows the synapsis configufations of 2n=4x= 68=4II+ 15IV. The latter has a remarkable and con- stant deleted chromosome, which is also the smallest in the karyotype.     (2) Ser. 2 Pumilae.  The series consists of six species endemic to China (Yu, 1979), all of which were cytologically examined, with the results: Sooo 9-8 of M. pumila Mill. 2n=34;  Sooo 9-12 of M. pumila Mill. 2n=34;  Sooo 9-7 of M.  sp.  2n=34;   M.  asiatica Nakai.  2n=68;   M.  prunifolia Borkh. 2n=34;  Soo 12-2 of M. prunifolia Borkh. 2n=34;  M. spectabilis Borkh. 2n=51;  M. micromalus Makino. 2n=34 and M. sieversii Roem. 2n=34. The karyotypes consist of sm(8-9 homologous chromosomes) and m (6- 7 homologous chromosomes) as well as st (1-2 homologous chromosomes). The chromosomes range in relative length from 10.77 to 3.42, with the ratios of the longest to the shortest from 2.95 to 2.04, and the mean arm ratios from 2.10 to 1.93. Therefore, all the karyotypes belong to 3B. Among more than 20 differ- ent forms of M. sieversii, there is only one ploidy level (2n=2x=34). PMC meiosis of M. spectabilis shows the synapsis configurations 2n=3x=51 =5I + 5II+ 12III.     2  Sect. II Sorbomalus     (3) Ser. 3 Sieboldianae.  This series has only one species endemic to China, M.  sieboldii  Rehd.:  One  triploid  from  Zunyi,  Guizhou  Province  has 2n=3x=51=24m+24sm+3st.  The  chromosomes  range  in  relative  length 8.74-- 4.17, with the ratio of the longest to the shortest 2.08, and the mean arm ratio 1.80. The karyotype, therfore, belongs to 2B. PMC meiosis of this form shows the synapsis conriguration 2n = 3x = 51 = 6I+ 9II+ 9III.     (4) Ser. 4 Kansuensis  consisting of four species endemic to China, 2 of which were cytologically studied, with the results: M.  toringoides Hughes. 2n=34, 51, 68 and M. transitoria Schneid. 2n=34, 51. The karyotypes consist of m (7—9  homologous  chromosomes)  and  sm (7—9  homologous chromosomes) as well as a few st chromosomes.  The chromosomes range in relative length from 9.44 to 4.01, with the ratios of the longest to the shortest from 2.26 to 1.83, and the mean arm ratios from 1.87 to 1.72. In M. toringoides four forms, 2x, 3x, 4x from Aba, and 3x from Barkang, in Sichuan Province, are of 2B karyotypes while 2x from Year zhu in Sichuan Province is of 2A. A mixploid with 3x, 4x in M. toringoides is found from Aba, with one smallest de- leted chromosome at both levels much like that in M.  xiaojinensis.  PMC meiosis  of  triploid  M.  toringoides  from  Barkang  showed  the  synapsis configurations of 10I+ 10II+ 7III. In M. transitoria 2x, 3x forms from Barkang, Sichuan Province and Lüda, Liaoning Province belong to 2A, while 2x, 3x forms from Aba, Sichuan Province both belong to 2B. PMC meiosis of triploid form from Lüda showed 9I+ 9II+ 8III synapsis configurations.     (5) Ser. 5 Yunnanensis  This series consists of four species endemic to China, 3  of which  were  cytologically  studied  with  the  results:  M.  ombrophila Hand-Mazz.  2n=34;   M.  honanensis Rehd.  2n=34,  and M.  yunnanensis Schneid. 2n= 34. The karyotypes consist of nine pairs of m, seven pairs of sm and one pair of st chromosomes. The chromosomes range in relative length from 8.43 to 4.37, with the ratios of the longest to the shortest from 1.88 to 1.86, and the mean arm ratios from 1.75 to 1.72. Therefore, the karyotypes belong to 2A. Two pairs of heterozygous chromosomes, 1st and 10th, are found carrying a small satellite.      3 Sect. III Docyniopsis.  This section consists of three endemic species to China, 2 of which were studied with the results:  M. formosana Kawak et Koidz. 2n= 34 and M. melliana Rehd. 2n= 34. Both karyotypes consist of nine pairs of m, seven pairs of sm and one pair of st chromosomes. The chromosomes range in relative length from 8.90 to 4.48, with the ratios of the longest to the shortest 1.85 and 1.83, and the mean arm ratios 1.67 and 1.66. Both karyotyoes brlong to 2A.      As mentioned above, the karyotyoes of all the species examined are relatively uniform, i.e. chromosomes small;  chromosomes gradually decreasing in size from the longest to the shortest;  chromosomes mostly m and sm;  and 1 —2 pairs of st chromosomes always present. The karyotypes vary in the ratio of the longeat to the shortest, the ratio of arms and the number of different chromosome types and satellites. From cytological investigations and previous reports (Chen et al., 1986;  Liang, 1987), the cytology of the genus varies mainly in euploidy variation and chromosome structure. Intraspecific polyploids and polyploid species make up 58. 1% in 24 species endemic to China reported. The karyotype analyse has revealed the presence of three major types in Malus: 2A, 2B and 3B. The dele- tion in some polyploid forms shows another structural variation. It is clear that the three types are generally consistent with the separation of three sections of Chinese Malus.  According to Stebbins's (1971),  the evolutionary trend  of karyotypes is from 2A to 2B and then 3B. The karyotypic data are in accordance with the evidence from morphology (Langenfeld,  1971;   Huckins,  1972) and biochemistry (Williams, 1982). They both show that the section Docyniopsis and series Yunnanenses are the most preimitive, while the series Pumilae, Baccatae of the section Malus are the most advanced or derived. The most typical external characters in Docyniopsis and the series Yunnanenses e.g. grit-cells in fruit, leaves folded in bud, and the inflorescence according to Ponomarenko’s (1986) classifica- tion, might be very primitive. The 2A karyotypes in the genus correlated with many typically premitive characters. From the data mentioned above, we may suppose that the series Sieboldianae and Kansuenses in the section Sorbomalus are intermediate in evolution. In general, Southwest China is regarded as the probable area  of the  origin  of Malus,  and  the  section  Docyniopsis  and  the  series Yunnanenses as the relict primitive forms. This area is probably also the center of differentiation.     A good deal of attention has been paid by horticulturists. A comparison of the karyotypes in the series Pumilae shows that the diploid M. sieversii has 14 m chromosomes, while the other diploids M. pumila, M. sp.  and an alien species  M.  sylvestris  Mill.  from  Europe ( Liang,  unpub.)  have  12  m chromosomes. Thus M. sieversii may be relatively primitive, although the ratio of the longest to the shortest and the ratio of mean arm ratio are similar. The present investigation confirms Ponomarenko’s view that the diploid M. sieversii might be an ancestor of cultivated apple (M. pumila).     The wild polyploids of Malus were usually considered as allopolyploid (Brown, 1976). Studies on the meiotic pairing of five triploids and one tetraploid have revealed diverse types of polyploidy. M. hupehensis is a typical allotriploid species, M. spectabilis, M. sieboldii, M. toringoides and M.  transitoria are segmental allotriploids, and M. xiaojinensis is almost an autotetraploid.     Apomixis in all known polyploid species of Malus is facultative. The princi- pal adaptive advantage of facultative apomixis is to make sterile individuals restore fertility, and also to maintain evolutionary potential. Therefore, it plays an important role in evolution of Malus, but taxonomy of apomictical groups re- mains highly controversial. Usually, the apomictical taxa are not treated as species level (Stace, 1980;  Hong, 1990). In Rehder’s (1940) system, only M. hupehensis was recognized as species in apomictical taxa of Malus. We think this treatment to be reasonable because M. hupehensis is an allotripoid and so far its diplois form has not been found.  New species M. xiaojinensis described by Cheng and Jiang (1985) is nearly an autotetraploid, and its karyotype consti- tution is extremely similar to the tetraploid form of M.  toringoides  from Aba , Sichuan Province, especially the remarkable similarities of the smallest deleted chromosome, morphology and biochemistry (Li, 1990). We think, therefor, that the triploid form of M. toringoides from Aba, Sichuan with 3x, 4x mixploidy might be the maternal plant of M. xiaojinensis. For this reason, M. xiaojinensis should be placed in apomictical polyploid complex of M. toringoides of the seriesKansuensis, and this is also in accordance with Rehder’s classfication.