摘要
金属型植物的抗性种群与同物种的非抗性种群相比,一个突出的特点是生物量较低。虽然总生物量不同,但是生物量在各个器官的分配比例是否在种群间也存在差异目前还不清楚。根据文献检索的结果,我们提出假设：抗性植物具有不同于非抗性植物的生物量分配方式,即在重金属胁迫下抗性植物能将更多的生物量分配到对其生存和繁衍具有重要作用的器官中(比如根与种子)。为了对假设进行验证并对其可能的生理与分子机制进行分析,我们进行了下列研究：①营养生长时期抗性与非抗性种群植物根/苗生物量分配对比研究；②生殖生长时期抗性与非抗性种群植物生长/生殖生物量分配对比研究；③种群间根/苗、生长/生殖生物量分配差异与库器官中蔗糖代谢的关系；④种群间酸性转化酶蛋白活性差异是否可能与酸性转化酶氨基酸序列变化有关；⑤种群间酸性转化酶蛋白活性差异是否与酸性转化酶基因转录表达量不同有关。本研究选用兼性金属型植物齿果酸模的抗性和非抗性种群作为实验材料。抗性种群的种子采自已有3000多年历史的大冶铜绿山古铜矿遗址,非抗性种群的种子采自武汉大学校园。本文通过水培与土培的方式分别对齿果酸模营养生长时期和生殖生长时期进行研究。从实验结果中可以得出如下结论：
(1)在铜胁迫下抗性与非抗性种群齿果酸模具有不同的根/苗生物量分配方式。铜处理使非抗性种群齿果酸模的根冠比降低,而使抗性种群齿果酸模的根冠比增加。非抗性种群齿果酸模的主根长、根尖数、根表面积和平均根直径均被铜处理显著抑制。抗性种群齿果酸模的主根长、根尖数、根表面积与对照相比稍有下降,而平均根直径稍有增加。
(2)在铜胁迫下抗性与非抗性种群齿果酸模具有不同的生长/生殖生物量分配方式。在铜胁迫下,抗性种群齿果酸模的地上部分营养器官生物量下降,而生殖器官生物量仍然维持不变,收获指数也因此增加。在铜胁迫下,非抗性种群齿果酸模的地上部分营养器官生物量、生殖器官生物量和收获指数均下降。在铜胁迫下,抗性种群齿果酸模的物候期提前,而非抗性种群齿果酸模的物候期推迟。在铜胁迫下,抗性种群齿果酸模的千粒重、花数、种子数、分支数和结实率均高于非抗性种群。
(3)在铜胁迫下抗性与非抗性种群齿果酸模不同的生物量分配方式与它们库器官中不同的酸性转化酶活性有关。在铜胁迫下,抗性种群齿果酸模根与种子中细胞壁转化酶活性和液泡转化酶活性均显著高于非抗性种群。细胞质转化酶活性和蔗糖合酶分解方向活性在种群间没有显著性差异。
(4)在铜胁迫下抗性与非抗性种群齿果酸模不同的酸性转化酶活性可能与它们不同的氨基酸序列有关。抗性与非抗性种群齿果酸模细胞壁转化酶基因氨基酸序列之间的一致性为95%；抗性与非抗性种群齿果酸模液泡转化酶基因氨基酸序列之间的一致性为99%。氨基酸序列的改变可能是影响酸性转化酶活性的一个因素。
(5)在铜胁迫下抗性与非抗性种群齿果酸模酸性转化酶蛋白活性差异与酸性转化酶基因转录表达量不同有关。在铜胁迫下,抗性种群齿果酸模根与种子中细胞壁转化酶基因和液泡转化酶基因的转录表达量均显著高于非抗性种群。
The biomass of tolerant plant is generally lower than that of non-tolerant plant. Although the total biomass is different, whether the biomass allocation among organs is also different between the two populations is still unknown. According to the retrieval of literatures, we suggested hypothesis that tolerant plant would differ from non-tolerant plant in the allocation of biomass. Under stress situation, tolerant plant would allocate more biomass to organs which are crucial to survival and reproduction, such as root and seed. In order to test this hypothesis, we did the following researches:(1) the comparative study of root/shoot biomass allocation between tolerant plant and non-tolerant plant;(2) the comparative study of growth/reproduction biomass allocation between tolerant plant and non-tolerant plant;(3) the relationship between biomass allocation and sucrose metabolism;(4) the relationship between acid invertase enzyme-activity and the alteration of amino acid sequence;(5) the relationship between acid invertase enzyme-activity and the expression of acid invertase gene. The plant material of this study was a metallophyte of Rumex dentatus L., one from a copper mine at Tonglushan hill (Cu-tolerant population), and the other from campus of Wuhan university (non-tolerant population). Hydroponic and soil cultures were respectively conducted to investigate the differences of biomass allocation at vegetative growth period and reproductive growth period. Following conclusions could be obtained from this study:
(1) There were differences in root/shoot biomass allocation between tolerant plant and non-tolerant plant. The root/shoot ratio of non-tolerant plant was reduced by Cu treatment, while the root/shoot ratio of tolerant plant was increased by Cu treatment. The root length, number of tips, root surface area and average root diameter of non-tolerant plant were significantly inhibited by Cu treatment. The root length, number of tips and root surface area of tolerant plant were a little lower than control, while average root diameter of tolerant plant was a little higher than control.
(2) There were differences in growth/reproduction biomass allocation between tolerant plant and non-tolerant plant. Under Cu stress, the aerial vegetative biomass decreased, while the reproductive biomass remained constant, and harvest index thus increased in tolerant plant. The aerial vegetative biomass, reproductive biomass and harvest index of non-tolerant plant were all decreased by Cu treatment. The time to first flowering was decreased in tolerant plant, while it was delayed in non-tolerant plant. The1000-seed weight, number of flowers, number of seeds, number of branches and ratio of seed set in tolerant plant were higher than those in non-tolerant plant.
(3) The different biomass allocation between the two populations was related to differential acid invertase enzyme-activity. The cell wall invertase activity and vacuole invertase activity in tolerant plant were higher than those in non-tolerant plant. There were no significantly differences in neutral/alkaline invertase activity and sucrose synthase activity between the two populations.
(4) The different acid invertase enzyme-activity between the two populations might be related to the alteration of amino acid sequence. The homology of cell wall invertase amino acid sequence between the two populations was95%. The homology of vacuole invertase amino acid sequence between the two populations was99%.
(5) The different acid invertase-enzyme activity between the two populations was related to differential expression of acid invertase gene. Under Cu stress, the expression of acid invertase gene in tolerant plant was higher than that in non-tolerant plant.
引文
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