巴山木竹种群生物学研究
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摘要
巴山木竹隶属于禾本科(Poaceae)竹亚科(Bambusoideae)青篱竹属(Arundinaria),是一中高山分布的中大型复轴混生型竹种;分布区以大巴山和秦岭两大山脉为主,涉及川、甘、陕、渝、鄂、豫等六省市和中亚热带、北亚热带、暖温带等三个气候带。巴山木竹是大熊猫的主食竹种之一,优质的造纸原料,被广泛用于搭菜架、编织等多种用途,其笋有较高的营养价值。本文对巴山木竹种群的一些生物学特性和遗传多样性进行了研究,结果如下:
随着海拔的升高,巴山木竹出笋期渐迟,海拔1900m处于4 月底5 月初出笋,退笋率31.50%,主要是营养不良的自然退笋;前期不同时间出土的笋,其高生长停止的时间基本相同,需时约70 天,白天的高生长量(58.13%)大于夜晚的,回归模型能很好地预测笋生长过程;巴山木竹种群高生长和土壤温度呈显著正相关关系。
巴山木竹地下茎的梢端可常年保持分生能力,受顶端优势的影响,很少产生岔鞭,所以对于带有梢端的巴山木竹地下茎的年龄,可根据节间长短的规律性变化进行鉴别;在较好的立地条件下,每年约产生13节,延伸60cm左右。巴山木竹地下茎和根的生命力维持时间可长达20余年,
巴山木竹无性系种群的空间分布格局为集群分布,且集群程度随时间进程而加强;格局规模即斑块大小都是92.16m2, 格局强度(RDD)达到3.89,说明巴山木竹种群聚块(密距段)分株数是间隙(疏距段)分株数的3.89倍。格局纹理为38.17cm,即从一个聚块(密距段)到另一个聚块(密距段)的聚块平均间距是38.17cm。
巴山木竹种群各构件单位的生物量配置为:秆(55.56%)>蔸(20.77%)>枝(9.51%)>鞭(8.70%)>叶(3.40%)>根(2.06%)。地上部生物量57.2%集中于6 龄及其以上竹中,种群年龄结构老化。7 龄以上立竹的秆重开始呈现下降趋势。
通过竹笋-幼竹数量动态生命表和生存曲线分析,巴山木竹无性系分株集中死亡的时期是幼竹刚进入迅速高生长的时期;巴山木竹无性系种群具有较大的更新潜力;静态生命表在巴山木竹种群数量统计中的应用有一定的局限性
施肥可显著增加出笋数和活笋数,短期内对笋地径、新竹及其胸径无显著影响;巴山木竹无性系分株之间生理整合的收益和耗费都比较明显;局部施肥可以提高施肥效率。早春的密度调节对当年各个密度处理下的出笋数、活笋数均未出现显著差异;对当年新竹的高度和胸径,也得到类似结果。
佛坪种群(51 个样品)的遗传多样性略高于镇巴种群(72 个样品);就此两种群而言,遗传多样性主要存在于种群内,占92.41%。从保护遗传多样性意义上讲,对巴山木竹无性系种群的取样原则应该是:取相距较远的种群,每个种群取样数量50 个即可。
对17 个种群进行遗传分析后得知,巴山木竹种内多样性丰富(Gst=0.5324)。
Arundinaria fargesii E.G.Camus belongs to Bambusoideae, Poaceae.It is a middle-mountain distributed middle-large amphipodial bamboo. It distributes mainly in the mountainous area of Dabashan and Qinling mountains, extending to Sichuan, Gansu, Shanxi, Chongqing, Hubei, Henan,six provinces. A. fargesii is one of the main food bamboos of Panda. It is fine material for paper making, and is widely used in weaving, as far as frame material in greenhouse. The bamboo shoots has high value of nutrition.
The more higher the altitude,the later the emergence of bamboo shoots. At the altitude of 1900m, A. fargesii shoots emerge at about the end of April or the early of May. Mortality of the shoots is about 31.50%. The main reason is shortage of nutrition. Shoots which emerged asynchronously at the early of shooting time, end their growth almost synchronously. The process lasts about 70 days. Regression equation could forecast the growing process well.The amount of growing during the day(58.13%) is larger than that of during the night. The heigh-growth of A. fargesii clonal population presents a positive correlativity with the 5cm-depth soil temperature.
The meristem at the rhizomatous tip of A. fargesii could keep its fissility for 20 years at least. Therefore the branching rhizome is hardly found. To such rhizome, however, it becomes easier to identify its ages because the length of the internodes varies regularly with aging in a year. In the favorable environment, the rhizome could produce about 13 nodes and stretch approximately 60cm one year.
The contiguous grid quadrats have been applied to sampling for field data. The spatial pattern of A. fargesii clone population was the clumped distribution and the pattern scale was at 92.16m2, the relative intensity difference(RDD) was 3.89, which means the culms in the clumps are 3.89 times density of those in the interspace between clumps. The pattern grain was 38.17cm, which implies the mean distance between clumps is 38.17cm. The sympodial ramets hold the dominant status in the clone growth of the A. fargesii population. The strengthening trend of clumped distribution with ageing implies that the population was under a relatively stronge environmental pressure.
The biomass allocations among culm, branch, leaf, root, rhizome, bamboo stump of A. fargesii were 55.6%, 9.51%, 3.40%, 2.06%, 8.70%, 20.77% respectively, while the aboveground biomass allocations of ramets from 1 to 5, 6 and 7, 8 and 9, 10 and more than 10 years old were 19.0%, 5.2%, 5.4%, 4.3%, 8.9%, 14.6%, 18.2%, 24.4% respectively. The underground biomass allocations among 0~20cm, 20~40cm and more deeper in depth of soil were 13.4%, 42.8%, 43.8% respectively. The stem weight of bamboos which were less than 3 years and over 8 age were lower than the standard stem weight, while which between 4 and 7 years were higher than the standard stem weight; With ageing: young----mature----old, the weight of leaf per ramet and the ratio of it to the weight of stem present the regularity: increasing----steady----declining.
Analyzing based on the dynamic life table and survivorship curve, the results show that the clone individuals of A. fargesii died mainly at the time when the young bamboos just enter the rapid-growing period; A. fargesii population has considerable potencial for renovating. The using of static life table in the demography of A. fargesii population is not perfect, for the longevity of individuals and the difficulty to identify the exact ages of the old ones. Fertilization could increase the amount of the bamboo shoots and their survivors distinctly. However there is no distinct effect on the basic diameter of bamboo shoots and the height and diameter at breast height(DBH) of the new bamboos. The integration among ramets is evident, including the benefits and the costs of physiological integration. Local fertilization would improve fertilizing effects in the cultivation of A. fargesii forest. The density regulation at early spring has no evident effect on the the number of daughter ramets and their survivors in the same year, and to the height and DBH of the new bamboos, has the similar result. More amplification products and polymorphic loci was obtained from Foping population with 51 samples than Zhenba population with 72 samples. Foping population has more genetic diversity than Zhenba population. As far as these two populations, the genetic diversity exist mainly within populations(92.41%). Sampling from far apart each other and representative populations, 50 samples is enough for one population, could obtain the most plenty genetic diversity of A. fargesii, with the fewest samples. The results from 17 populations show that there are plenty genetic diversity within A. fargesii (Gst=0.5324).
随着海拔的升高,巴山木竹出笋期渐迟,海拔1900m处于4 月底5 月初出笋,退笋率31.50%,主要是营养不良的自然退笋;前期不同时间出土的笋,其高生长停止的时间基本相同,需时约70 天,白天的高生长量(58.13%)大于夜晚的,回归模型能很好地预测笋生长过程;巴山木竹种群高生长和土壤温度呈显著正相关关系。
巴山木竹地下茎的梢端可常年保持分生能力,受顶端优势的影响,很少产生岔鞭,所以对于带有梢端的巴山木竹地下茎的年龄,可根据节间长短的规律性变化进行鉴别;在较好的立地条件下,每年约产生13节,延伸60cm左右。巴山木竹地下茎和根的生命力维持时间可长达20余年,
巴山木竹无性系种群的空间分布格局为集群分布,且集群程度随时间进程而加强;格局规模即斑块大小都是92.16m2, 格局强度(RDD)达到3.89,说明巴山木竹种群聚块(密距段)分株数是间隙(疏距段)分株数的3.89倍。格局纹理为38.17cm,即从一个聚块(密距段)到另一个聚块(密距段)的聚块平均间距是38.17cm。
巴山木竹种群各构件单位的生物量配置为:秆(55.56%)>蔸(20.77%)>枝(9.51%)>鞭(8.70%)>叶(3.40%)>根(2.06%)。地上部生物量57.2%集中于6 龄及其以上竹中,种群年龄结构老化。7 龄以上立竹的秆重开始呈现下降趋势。
通过竹笋-幼竹数量动态生命表和生存曲线分析,巴山木竹无性系分株集中死亡的时期是幼竹刚进入迅速高生长的时期;巴山木竹无性系种群具有较大的更新潜力;静态生命表在巴山木竹种群数量统计中的应用有一定的局限性
施肥可显著增加出笋数和活笋数,短期内对笋地径、新竹及其胸径无显著影响;巴山木竹无性系分株之间生理整合的收益和耗费都比较明显;局部施肥可以提高施肥效率。早春的密度调节对当年各个密度处理下的出笋数、活笋数均未出现显著差异;对当年新竹的高度和胸径,也得到类似结果。
佛坪种群(51 个样品)的遗传多样性略高于镇巴种群(72 个样品);就此两种群而言,遗传多样性主要存在于种群内,占92.41%。从保护遗传多样性意义上讲,对巴山木竹无性系种群的取样原则应该是:取相距较远的种群,每个种群取样数量50 个即可。
对17 个种群进行遗传分析后得知,巴山木竹种内多样性丰富(Gst=0.5324)。
Arundinaria fargesii E.G.Camus belongs to Bambusoideae, Poaceae.It is a middle-mountain distributed middle-large amphipodial bamboo. It distributes mainly in the mountainous area of Dabashan and Qinling mountains, extending to Sichuan, Gansu, Shanxi, Chongqing, Hubei, Henan,six provinces. A. fargesii is one of the main food bamboos of Panda. It is fine material for paper making, and is widely used in weaving, as far as frame material in greenhouse. The bamboo shoots has high value of nutrition.
The more higher the altitude,the later the emergence of bamboo shoots. At the altitude of 1900m, A. fargesii shoots emerge at about the end of April or the early of May. Mortality of the shoots is about 31.50%. The main reason is shortage of nutrition. Shoots which emerged asynchronously at the early of shooting time, end their growth almost synchronously. The process lasts about 70 days. Regression equation could forecast the growing process well.The amount of growing during the day(58.13%) is larger than that of during the night. The heigh-growth of A. fargesii clonal population presents a positive correlativity with the 5cm-depth soil temperature.
The meristem at the rhizomatous tip of A. fargesii could keep its fissility for 20 years at least. Therefore the branching rhizome is hardly found. To such rhizome, however, it becomes easier to identify its ages because the length of the internodes varies regularly with aging in a year. In the favorable environment, the rhizome could produce about 13 nodes and stretch approximately 60cm one year.
The contiguous grid quadrats have been applied to sampling for field data. The spatial pattern of A. fargesii clone population was the clumped distribution and the pattern scale was at 92.16m2, the relative intensity difference(RDD) was 3.89, which means the culms in the clumps are 3.89 times density of those in the interspace between clumps. The pattern grain was 38.17cm, which implies the mean distance between clumps is 38.17cm. The sympodial ramets hold the dominant status in the clone growth of the A. fargesii population. The strengthening trend of clumped distribution with ageing implies that the population was under a relatively stronge environmental pressure.
The biomass allocations among culm, branch, leaf, root, rhizome, bamboo stump of A. fargesii were 55.6%, 9.51%, 3.40%, 2.06%, 8.70%, 20.77% respectively, while the aboveground biomass allocations of ramets from 1 to 5, 6 and 7, 8 and 9, 10 and more than 10 years old were 19.0%, 5.2%, 5.4%, 4.3%, 8.9%, 14.6%, 18.2%, 24.4% respectively. The underground biomass allocations among 0~20cm, 20~40cm and more deeper in depth of soil were 13.4%, 42.8%, 43.8% respectively. The stem weight of bamboos which were less than 3 years and over 8 age were lower than the standard stem weight, while which between 4 and 7 years were higher than the standard stem weight; With ageing: young----mature----old, the weight of leaf per ramet and the ratio of it to the weight of stem present the regularity: increasing----steady----declining.
Analyzing based on the dynamic life table and survivorship curve, the results show that the clone individuals of A. fargesii died mainly at the time when the young bamboos just enter the rapid-growing period; A. fargesii population has considerable potencial for renovating. The using of static life table in the demography of A. fargesii population is not perfect, for the longevity of individuals and the difficulty to identify the exact ages of the old ones. Fertilization could increase the amount of the bamboo shoots and their survivors distinctly. However there is no distinct effect on the basic diameter of bamboo shoots and the height and diameter at breast height(DBH) of the new bamboos. The integration among ramets is evident, including the benefits and the costs of physiological integration. Local fertilization would improve fertilizing effects in the cultivation of A. fargesii forest. The density regulation at early spring has no evident effect on the the number of daughter ramets and their survivors in the same year, and to the height and DBH of the new bamboos, has the similar result. More amplification products and polymorphic loci was obtained from Foping population with 51 samples than Zhenba population with 72 samples. Foping population has more genetic diversity than Zhenba population. As far as these two populations, the genetic diversity exist mainly within populations(92.41%). Sampling from far apart each other and representative populations, 50 samples is enough for one population, could obtain the most plenty genetic diversity of A. fargesii, with the fewest samples. The results from 17 populations show that there are plenty genetic diversity within A. fargesii (Gst=0.5324).
引文
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