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题名: 天仙子族和钩毛草属的细胞学特征及其在生物地理学上的意义
作者: 涂铁要
学位类别: 硕士
答辩日期: 2005
授予单位: 中国科学院昆明植物研究所
授予地点: 中国科学院昆明植物研究所
导师: 孙航
关键词: 横断山区 ; 天仙子族 ; 山莨菪属 ; 天蓬子属 ; 天仙子属 ; 马尿泡属 ; 茄参属 ; 钩毛草属 ; 多倍化 ; 染色体进化
学位专业: 植物学
中文摘要: 对横断山区的特征类群茄科天仙子族(Hyoscyalneae)和横断山(东亚西部)-北美西部间断类群茜草科钩毛草属(Kelloggia)进行了细胞地理学研究。共报道了5属10种14个居群的染色体数目或核型,第一次报道了山蓑若属(Abusidys)、天蓬子属(Atropanthe)和钩毛草属所有种的染色体数目和核型以及茄参(Mandragoracalescense)的染色体数目和马尿泡属(Przewalskia)的核型。通过分析这两个类群的细胞地理特征并且综合其它相关的细胞学资料以及横断山区的地质历史和气候环境特点,讨论了这两个类群在古地中海退却和喜马拉雅一横断山隆起后的染色体进化模式和机制。对天仙子族的细胞地理学研究结果包括以下几个方面:1)有丝分裂前期染色体形态在山蓑若属(Anisodus)、天蓬子属(Atropanthe)、茄参属(Mandragora)、马尿泡属(Przewalskia)和天仙子属(Hyoscyamus)中均为近基型。2)山蓑若属、天蓬子属和茄参属的间期核属于球状前染色体型,马尿泡属的间期核属于棒状前染色体型,而天仙子属的间期核属于复杂染色中心型。前4属植物的间期核属于前染色体亚类型,不同于天仙子属的复杂染色中心型,这表明前4属有较近的亲缘关系,这一结果也得到了形态解剖学证据的支持。3)分布于青藏高原的单型属马尿泡属的系统位置以前存在很大争论。一部分学者认为该属是由山蓑若属适应高寒环境后分化出来的年轻的属,而Hoare&Knapp(1 997)认为该属比山蓑若属、赛蓑若属(Scopolia)等更为原始·我们从核型不对称性的角度对这一群植物进行分析表明马尿泡属的核型不对称性类型为ZA型,而山蓑若属和天蓬子属为ZB型,从核型不对称性的一般发展趋势来看,马尿泡属显然更为原始,因此它更可能是青藏高原的一个古老残遗。4)茄参属因具浆果而被许多植物学家认为是一个孤立的属,不应包括在天仙子族中,本研究表明茄参属具有和山蓑若属及天蓬子属一致的染色体基数和前期染色体形态以及间期核形态,细胞学结果表明本属应置于天仙子族之中。5)我们所研究的本族5属7种植物的核型公式分别为:铃铛子(Anisodus luridus Link:2,,=48=32m+16sm(Zsat);三分三(A .acutal;gulus C:Wu&c.chen):Zn=48=4om+ssm(lsat);山蓖若州.ta叮uticus(Maxim.)Pascher):Zn=48=40m+ssm(Zsat);赛蓑著例.ca,·,,iolicoides(C.Y.Wu&C.chen)D,Aicy&2.Y. Zhang):Zn:48=4om+ssm(3sat);天蓬子(At].oranthe sine,;515(Hemsley)Pascher):2,?=48=46m+Zsm;马尿泡(P,ze飞性,alskia tangutica Maxim.):2,2=44=36m+ssm(lsat);天仙子(场)05卿a刀,us,?igerL.):21;=34=24m+10sm(2 sat)。不难看出山蓑若属4种植物的核型极其相似,从而支持包括铃挡子、三分三、山蓖若和赛蓑著4种植物在内的山莫著属是一个自然的类群。天蓬子属的核型也和山蓑著属非常相近,它们可能是后来才发生分化的较为年轻的类群。6)综合分析该族植物的染色体倍性和地理分布特征我们发现分布于横断山区,的山蓑若属、天蓬子属和茄参属的种均为x=6的八倍体,而分布于地中海地区的茄参属和颠茄属既有较高的倍性(16x),也存在较低的倍性(4x),这说明多倍性普遍存于该族中,天仙子族中的多倍化可能在喜马拉雅一横断山隆升以前就已经发生。天仙子属中的古多倍体(x:14,17)也说明了这一点—多倍体在早期就己经形成,到后来又经过二倍化以后才形成了现在的次生二倍体。钩毛草属(Kell口ggia)是茜草科中的一个寡种属,含2种,一种(云南钩毛草Kelloggia cllinensis Franch.)分布于东亚西部的横断山区(海拔3000一4300米),另一种(K. galioides Torr.)见于北美西部山区(海拔1000一3000米)。我们第一次对全部2种共4个居群进行了细胞学研究。2种植物的间期核形态和前期染色体形态一致,分别为稀淡分散型和近基型。它们的核型公式分别为:云南钩毛草:加:22== 16m+6sm;尤.alioi山s:Zn=22;16m+6sm(Zsat)。两种植物的核型的高度相似性说明该属是一个自然的类群。云南钩毛草和K.gazioides的核型不对称类型分别为IA和ZA。北美的类群表现出了较为不对称的核型特征,故推测,北美的类群可能起源于欧亚大陆。从染色体倍性来看,2种植物均为x=11的二倍体句可见,喜马拉雅一横断山的隆起对其染色体倍性没有影响。本研究中的两个类群都是古地中海起源的,由于第三纪以后喜马拉雅隆升、欧亚大陆环境变迁和中亚干旱,才形成了今天这样的分布格局。细胞学资料揭示了这两个古地中海发生的类群其细胞进化主要体现在染色体内部结构的变化上(如随体数目、核型不对称性),而不是在倍性的变化上。高原隆升对此二类群的染色体倍性似乎没有明显的影响。
英文摘要: Cytological studies were carried out on the tribe Hyoscyameae, which is the only tribe of Solanaceae distributed in Eurasia and is centred in the Himalayan-Hengduan Mountains and in the adjacent areas of SW China, and a small genus Kelloggia with two species, which intercontinently disjuncts between eastern Asia and western North America. Chromosome numbers and/or karyotypes of 14 populations from 10 species in 5 genera are reported in this study. The chromosome numbers of Anisodus, Atropanthe, Kelloggia, Mandragora caulescense and the karyotypes of Anisodus, Atropanthe, Kelloggia, and Przewalskia are reported for the first time. Combining the cytological characters of these two taxa with the geological history and environment condition, the model and mechanism of chromosome evolution for these two taxa in the Hengduan Mountains were discussed. The results and conclusions of Hyoscyameae include: 1) All species studied in this tribe showed the proximal type of mitotic prophase chromosome condensation pattern. 2) Three types of interphase nuclei were recognized: the round prochromosome type for Anisodus, Atropanthe, and Mandragora, the rod prochromosome type for Przewalskia, and the complex chromocentre type for Hyoscyamus. The interphase nuclei of the former four genera belong to the sub-prochromosome type, which is different from that of Hyoscyamus (complex chromocentre type), and indicate a close relationship of the former four genera, which was also supported by previous studies. 3) The systematic position of the Tibet endemic Przewalskia has long been controversial. Some botanists thought it a very young genus differentiated from Anisodus after adapted to the extreme climate on the Plateau. But Hoare & Knapp (1997) considered it to be basal. The basal position of Przewalskia can also be supported by the karyotype asymmetry in this study because type 2A of Przewalskia is often considered a more primitive character than type 2B of Anisodus and Atropanthe. Thus it might be an old relic on the Plateau. 4) Some earlier classifications excluded Mandragora out of the tribe Hyoscyameae because they bear bemes and not capsules. This view point can not be supported by our cytological data because the base number, prophase chromosome pattern, and interphase nuclei of Mandragora are consistent with that of Anisodus and Atropanthe. 5) The karyotypes of seven species studied are: Anisodus luridus Link: 2n = 48 = 32m + 16sm (2sat); A. acutangulus C. Y. Wu & C. Chen: In - 48 = 40m + 8sm(lsat); A. tanguticus (Maxim.) Pascher: In = 48 = 40m + 8sm(2sat); A. carniolicoides (C. Y. Wu & C. Chen) D'Arcy & Z. Y. Zhang: 2n = 48 = 40m + 8sm(3sat); Atropanthe sinensis (Hemsley) Pascher: In = 48 = 46m + 2sm; Przewalskia tangutica Maxim..- In = 44 = 36m + 8sm(lsat); Hyoscyamus niger L.: In - 34 = 24m + 10sm(2sat)o The karyotypes of four species of Anisodus are very similar. This result supports that Anisodus, which includes Anisodus luridus, A. acutangulus, A. tanguticus and A. carniolicoides, is a very nature genus. The karyotype of Atropanthe is also similar to that of Anisodus. These two genera might have very short evolutionary history. 6). Polyploids are found in most species of the tribe in the Himalayan-Hengduan Mountains, as well as in the Mediterranean region, WC Asia, and E Asia. It seems that polyploidy probably happened very early in Hyoscyameae, before the uplift of the Himalayan-Hengduan Mountains. The paleopolyploids in Hyoscyamus (x = 14, 17) also show an early occurrence of its polyploidization, which are followed by diploidization to produce hypo-diploids with x = 14 and x = 17. The Himalayan orogeny might have played a minor role in the polyploid evolution of plants in this tribe. Another cytological study was carried out for the first time on four populations of only two species of Kelloggia (Rubiaceae), which occurs disjunctly between eastern Asia (K. chinensis Franch.) and western North America (K. galioides Torr.). The consistent mitotic prophase chromosome condensation pattern as proximal type and interphase nuclei as the sparsely diffuse type were determined for both species. The karyotypes of In = 22 = 2x = 16m + 6m and In = 22 = 2x = 16m + 6m (2sat) and the asymmetry of 1A and 2A were found for K. chinensis and AT. galioides respectively. Although the two species exhibit a high level of morphological divergence and are widely disjunctive, little difference was found in their karyomorphological features. Their main differences in cytology are the karyotype asymmetry and number of satellites. The chromosome base number of this genus is safely suggested to be x = 11. Our results support the close relationship of these two species and the Old World origin of this genus. The distributional patterns of these two Tethyan flora originated taxa were attributed to geological changing and M Asian aridity corresponding to the uplift of Himalayan-Hengduan Mountains. Our cytological data indicate that the intrachromosomal variation in the structure of these two taxa represents a major evolutionary line on the cytological evolutionary level in the Hengduan Mountains and the uplift of the Plateau didn't play a great role on their polyploidization.
语种: 中文
内容类型: 学位论文
URI标识: http://ir.kib.ac.cn/handle/151853/814
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天仙子族和钩毛草属的细胞学特征及其在生物地理学上的意义.涂铁要[d].中国科学院昆明植物研究所,2005.20-25
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