中国濒危物种野生玫瑰的遗传多样性及其影响因子研究
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摘要
野生玫瑰为国家二级保护植物,是栽培玫瑰育种的重要种质资源。野生玫瑰主要分布在中国北部沙质和砾质海岸、河口沙丘,是当地灌丛群落的优势物种之一。在过去二十年中,由于生境退化和丧失,野生玫瑰种群的分布范围、种群数目和大小都在急剧缩减,玫瑰灌丛常以小斑块状存在。本文对牟平野生玫瑰进行了种群生态学调查,在每个斑块中详细记录了斑块的面积,乔木层郁闭度,玫瑰的平均高度、盖度,草本层植物的种类等指标,对玫瑰的繁殖特性也进行了调查。
玫瑰种群生态学调查表明:野生玫瑰多分布于海滨林缘地带,既喜光又需要乔木层对强光的遮蔽;在野生玫瑰群落中,玫瑰一般为单优种,灌木层伴生植物极少,而草本层种类相对较多,共有40余种。对玫瑰的繁殖特性进行的调查表明,尽管玫瑰每年都产生大量的果实和种子,但种子苗却很少见,土壤种子库也很贫乏,在自然环境中,野生玫瑰以行茎繁殖为主。在牟平种群中共观测到52种昆虫,其中访花昆虫9种,出现频率较高的黄胸木蜂(Xylocopa appendiculataSmith)、中华细腹食虫虻(Leptogaster sinensis Hsia)都起到了为玫瑰传播花粉的作用。
把群落数据用相关软件进行分析,得出玫瑰的高度与盖度与其它指标之间的关系。研究发现:玫瑰的平均高度与盖度具有非常显著的正相关(r=0.4744,p=0.0108),相关分析表明当玫瑰的高度增加时,玫瑰的盖度也相应增加;乔木层的郁闭度与玫瑰的盖度之间呈显著的负相关(r=-0.5474,p=0.0026),但与高度相关不显著。当乔木层郁闭度增加时,玫瑰通过向外扩散来获得更多的阳光,但其高度不会显著增加。乔木层郁闭度与斑块面积没有显著的相关性,乔木层的覆盖不会使玫瑰无限制地向外扩张,玫瑰灌丛都形成了明显的边界。相关分析表明,玫瑰的高度与盖度与草本层植物的种类都没有显著相关。这些结果表明:乔木层郁闭度是影响玫瑰生长的主要的非生物因素,而玫瑰的存在不会显著影响当地生态系统的物种多样性。
为了研究影响玫瑰遗传多样性的因素,我们对牟平玫瑰种群的遗传多样性与相关指标进行了相关分析。结果表明,玫瑰的遗传多样性与高度和盖度之间没有显著关系,与乔木层的郁闭度也没有显著关系,与克隆多样性的关系也不显著。但是当取样数较小(小于14)时,取样大小对遗传多样性有着显著影响,两者呈现显著的正相关(r=0.7279,p=0.00002)。当斑块面积较小时,斑块面积与遗传多样性呈显著正相关(r=0.6434,p=0.0012),这与在斑块面积大的样地,取样数目多是分不开的。当斑块面积大于500 m~2时,斑块面积与遗传多样性不再显著相关,表明斑块面积大的样地,遗传多样性不一定高。草本层植物的种类与玫瑰遗传多样性显著正相关(r=0.3927,p=0.0387),草本层的覆盖改善了斑块内的小环境,利于有性繁殖的成功。对玫瑰空间距离与遗传距离的相关分析表明,随着空间距离的增大,遗传距离显著升高,各斑块间的遗传分化明显,相邻斑块间的关系较近。
为了分析玫瑰的整体遗传多样性水平及遗传结构,本研究采用RAPD方法检测了该物种在中国仅存的四个较大种群的遗传多样性水平。遗传多样性研究结果显示,中国野生玫瑰种群具有较高的遗传多样性水平。四个种群的平均多态位点百分比(P%)为57.99%,平均Shannon多样性指数(H_0)为0.2826;总的遗传多样性水平为P%=75.30%,H_0=0.3513。中国野生玫瑰种群间具有较低的遗传分化(G_(ST)=0.1878),表明大部分的遗传多样性存在于种群内部。四个野生玫瑰种群中,图们江种群的遗传多样性最高(P%=66.27%,H_0=0.3117),而且在种群中检测到了两条种群特异性谱带(OPS20-1,OPX04-3);长山岛种群遗传多样性次之(P%=59.04%,H_0=0.3065)。图们江种群中包含了95.33%在总种群中频率大于0.05的带,长山岛种群包含了99.33%。这些结果说明在野生玫瑰的就地保护中,图们江种群和长山岛种群应给予优先保护;在迁地保护中,这两个种群也可以作为野生玫瑰种子或根茎繁殖体的来源地。在中国,四个种群均包含了95%以上在总居群中频率大于0.05的基因,野生玫瑰的濒危主要是人为影响而不是种群内因,因此要保护和恢复野生玫瑰种群首先要停止人类对其生境的破坏。本研究也可为其它与野生玫瑰伴生的海岸沙生濒危植物如单叶蔓荆、珊瑚菜、沙滩黄芩等的保护提供借鉴。
Rosa rugosa Thunb.is valuable germplasm for breeding cultivated roses,which had been ranked in the secondary protected plants.R.rugosa,is one of the dominant and important shrub species in estuary dunes and shingle beaches of northern China. However,its area of distribution and the number of populations as well as the size of each have decreased rapidly in the past two decades because of degradation and loss of habitats,and the habitats of R.rugosa have been fragmented.In the study, population ecology of R.rugosa in Muping was investigated.In every patch,patch area,arbor closure,mean height and coverage of R.rugosa,number of herbaceous species was measured.Propagating characteristic of R.rugosa was investigated in natural condition.
The results showed R.rugosa mainly distributed at the edge of the forest,which were inclined to sunlight but needed some shade of arbor.In R.rugosa communities, there were few of shrubs but many herbaceous species,more than 40 species.R. rugosa were usually dominant species in communities.R.rugosa produce a great deal of fruits and seeds every year,but seedlings is seldom and soil seed band is lack.In natural habitat,R.rugosa usually propagated by rhizomes.Fifty-two insects species was observed in population MP,nine of those visited flowers of R.rugosa.Xylocopa appendiculata Smith and Leptogaster sinensis Hsia wered observed most time,both of them were dispersers of pollen of R.rugosa.
Correlative software was used to analyze the data to get the relation between the mean height and coverage of R.rugosa and other indexes.The results showed:the mean height of R.rugosa had remarkable correlation with the coverage of R.rugosa (r=0.4744,p=0.0108).It suggested the coverage of R.rugosa increased with the increasing of mean height of R.rugosa.The arbor closure had remarkable negative correlation with the coverage of R.rugosa(r=-0.5474,p=0.0026),but not remarkable correlation with the mean height of R.rugosa.It suggested that R.rugosa grew dispersed to get more sunlight when arbor closure increased,but the height of R. rugosa did not increase remarkably.The arbor closure had not remarkable correlation with the patch area,which suggested shading coming from arbor did not make R. rugosa spread infinitely.The mean height and coverage of R.rugosa had not remarkable correlation with the number of herbaceous species.All results showed arbor closure was important abiotic factor to influence the growth of R.rugosa,and existing of R.rugosa did not influence biodiversity of local ecosystem in current circumstance.
To study the factors influencing genetic diversity of R.rugosa,the genetic diversity of R.rugosa in population MP and correlative indexes were analyzed.These results showed:the genetic diversity of R.rugosa had not remarkable correlation with the height(r=-0.0380,p=0.8479) and coverage(r=-0.1019,p=0.6060) of R.rugosa, the arbor closure(r=0.1609,p=0.4134) and the clonal diversity(r=0.3736,p=0.0502). But the sampling number had remarkable positive correlation with the genetic diversity when sampling number was not more than 14(r=0.7279,p=0.00002).The patch area had remarkable positive correlation with the genetic diversity when the patch area is small(r=0.6434,p=0.0012).When the patch area is more than 500 m~2, the patch area had not remarkable correlation with genetic diversity of R.rugosa, which suggested genetic diversity of R.rugosa might be low in big patch.The number of herbaceous species had remarkable correlation with genetic diversity of R.rugosa (r=0.3927,p=0.0387),which indicated herbaceous plants could improve little circumstance in patch,and it was of advantage to success of sexual propagating.The correlative analysis between spatial distance and genetic distance showed genetic distance increased remarkably with the increasing of spatial distance.It suggested there were close relation in neighboring patch,and there were obvious genetic differentiation between patches.
RAPD(random amplified polymorphic DNA) markers were applied to detect the genetic diversity of four remaining large natural populations and to discuss an effective conservation strategy of the endangered species R.rugosa in China.High genetic variations were detected in R.rugosa populations of China.The mean percentage of polymorphic loci(P%) within four local populations was 57.99%,and the percentage of polymorphic loci(P%) of the total population was 75.30%;the mean Shannon's information index(H_0) was 0.2826,and the total Shannon's information index(H_0) was 0.3513.The genetic differentiation among populations (G_(ST)) was 0.1878,which indicated most genetic diversity occurs within populations. Population Tumenjiang(TMJ) showed the highest genetic diversity(P%=66.27%, H_0=0.3117),and contained two exclusive bands;population Changshandao(CSD) showed higher genetic diversity(P%=59.04%,H_0=0.3065).Population TMJ contained 95.33%,and population CSD contained 99.33%of loci with moderate to high frequency(p＞0.05) of the total population.These results indicated that population TMJ and CSD should be given priority for in situ conservation and regarded as seed or propagule sources for ex situ conservation.The study also suggested that R.rugosa in China had become endangered as a result of human actions rather than genetic depression of populations;thus,human interference should be absolutely forbidden in R.rugosa habitats.This study can be referenced to establish conservation strategies for other coastal endangered species accompanied with R.rugosa,such as Vitex trifolia L.var.simplicifolia Cham.,Glehnia littoralis F. Schmidt ex Miq.,Scutellaria strigillosa Hemsl..
玫瑰种群生态学调查表明:野生玫瑰多分布于海滨林缘地带,既喜光又需要乔木层对强光的遮蔽;在野生玫瑰群落中,玫瑰一般为单优种,灌木层伴生植物极少,而草本层种类相对较多,共有40余种。对玫瑰的繁殖特性进行的调查表明,尽管玫瑰每年都产生大量的果实和种子,但种子苗却很少见,土壤种子库也很贫乏,在自然环境中,野生玫瑰以行茎繁殖为主。在牟平种群中共观测到52种昆虫,其中访花昆虫9种,出现频率较高的黄胸木蜂(Xylocopa appendiculataSmith)、中华细腹食虫虻(Leptogaster sinensis Hsia)都起到了为玫瑰传播花粉的作用。
把群落数据用相关软件进行分析,得出玫瑰的高度与盖度与其它指标之间的关系。研究发现:玫瑰的平均高度与盖度具有非常显著的正相关(r=0.4744,p=0.0108),相关分析表明当玫瑰的高度增加时,玫瑰的盖度也相应增加;乔木层的郁闭度与玫瑰的盖度之间呈显著的负相关(r=-0.5474,p=0.0026),但与高度相关不显著。当乔木层郁闭度增加时,玫瑰通过向外扩散来获得更多的阳光,但其高度不会显著增加。乔木层郁闭度与斑块面积没有显著的相关性,乔木层的覆盖不会使玫瑰无限制地向外扩张,玫瑰灌丛都形成了明显的边界。相关分析表明,玫瑰的高度与盖度与草本层植物的种类都没有显著相关。这些结果表明:乔木层郁闭度是影响玫瑰生长的主要的非生物因素,而玫瑰的存在不会显著影响当地生态系统的物种多样性。
为了研究影响玫瑰遗传多样性的因素,我们对牟平玫瑰种群的遗传多样性与相关指标进行了相关分析。结果表明,玫瑰的遗传多样性与高度和盖度之间没有显著关系,与乔木层的郁闭度也没有显著关系,与克隆多样性的关系也不显著。但是当取样数较小(小于14)时,取样大小对遗传多样性有着显著影响,两者呈现显著的正相关(r=0.7279,p=0.00002)。当斑块面积较小时,斑块面积与遗传多样性呈显著正相关(r=0.6434,p=0.0012),这与在斑块面积大的样地,取样数目多是分不开的。当斑块面积大于500 m~2时,斑块面积与遗传多样性不再显著相关,表明斑块面积大的样地,遗传多样性不一定高。草本层植物的种类与玫瑰遗传多样性显著正相关(r=0.3927,p=0.0387),草本层的覆盖改善了斑块内的小环境,利于有性繁殖的成功。对玫瑰空间距离与遗传距离的相关分析表明,随着空间距离的增大,遗传距离显著升高,各斑块间的遗传分化明显,相邻斑块间的关系较近。
为了分析玫瑰的整体遗传多样性水平及遗传结构,本研究采用RAPD方法检测了该物种在中国仅存的四个较大种群的遗传多样性水平。遗传多样性研究结果显示,中国野生玫瑰种群具有较高的遗传多样性水平。四个种群的平均多态位点百分比(P%)为57.99%,平均Shannon多样性指数(H_0)为0.2826;总的遗传多样性水平为P%=75.30%,H_0=0.3513。中国野生玫瑰种群间具有较低的遗传分化(G_(ST)=0.1878),表明大部分的遗传多样性存在于种群内部。四个野生玫瑰种群中,图们江种群的遗传多样性最高(P%=66.27%,H_0=0.3117),而且在种群中检测到了两条种群特异性谱带(OPS20-1,OPX04-3);长山岛种群遗传多样性次之(P%=59.04%,H_0=0.3065)。图们江种群中包含了95.33%在总种群中频率大于0.05的带,长山岛种群包含了99.33%。这些结果说明在野生玫瑰的就地保护中,图们江种群和长山岛种群应给予优先保护;在迁地保护中,这两个种群也可以作为野生玫瑰种子或根茎繁殖体的来源地。在中国,四个种群均包含了95%以上在总居群中频率大于0.05的基因,野生玫瑰的濒危主要是人为影响而不是种群内因,因此要保护和恢复野生玫瑰种群首先要停止人类对其生境的破坏。本研究也可为其它与野生玫瑰伴生的海岸沙生濒危植物如单叶蔓荆、珊瑚菜、沙滩黄芩等的保护提供借鉴。
Rosa rugosa Thunb.is valuable germplasm for breeding cultivated roses,which had been ranked in the secondary protected plants.R.rugosa,is one of the dominant and important shrub species in estuary dunes and shingle beaches of northern China. However,its area of distribution and the number of populations as well as the size of each have decreased rapidly in the past two decades because of degradation and loss of habitats,and the habitats of R.rugosa have been fragmented.In the study, population ecology of R.rugosa in Muping was investigated.In every patch,patch area,arbor closure,mean height and coverage of R.rugosa,number of herbaceous species was measured.Propagating characteristic of R.rugosa was investigated in natural condition.
The results showed R.rugosa mainly distributed at the edge of the forest,which were inclined to sunlight but needed some shade of arbor.In R.rugosa communities, there were few of shrubs but many herbaceous species,more than 40 species.R. rugosa were usually dominant species in communities.R.rugosa produce a great deal of fruits and seeds every year,but seedlings is seldom and soil seed band is lack.In natural habitat,R.rugosa usually propagated by rhizomes.Fifty-two insects species was observed in population MP,nine of those visited flowers of R.rugosa.Xylocopa appendiculata Smith and Leptogaster sinensis Hsia wered observed most time,both of them were dispersers of pollen of R.rugosa.
Correlative software was used to analyze the data to get the relation between the mean height and coverage of R.rugosa and other indexes.The results showed:the mean height of R.rugosa had remarkable correlation with the coverage of R.rugosa (r=0.4744,p=0.0108).It suggested the coverage of R.rugosa increased with the increasing of mean height of R.rugosa.The arbor closure had remarkable negative correlation with the coverage of R.rugosa(r=-0.5474,p=0.0026),but not remarkable correlation with the mean height of R.rugosa.It suggested that R.rugosa grew dispersed to get more sunlight when arbor closure increased,but the height of R. rugosa did not increase remarkably.The arbor closure had not remarkable correlation with the patch area,which suggested shading coming from arbor did not make R. rugosa spread infinitely.The mean height and coverage of R.rugosa had not remarkable correlation with the number of herbaceous species.All results showed arbor closure was important abiotic factor to influence the growth of R.rugosa,and existing of R.rugosa did not influence biodiversity of local ecosystem in current circumstance.
To study the factors influencing genetic diversity of R.rugosa,the genetic diversity of R.rugosa in population MP and correlative indexes were analyzed.These results showed:the genetic diversity of R.rugosa had not remarkable correlation with the height(r=-0.0380,p=0.8479) and coverage(r=-0.1019,p=0.6060) of R.rugosa, the arbor closure(r=0.1609,p=0.4134) and the clonal diversity(r=0.3736,p=0.0502). But the sampling number had remarkable positive correlation with the genetic diversity when sampling number was not more than 14(r=0.7279,p=0.00002).The patch area had remarkable positive correlation with the genetic diversity when the patch area is small(r=0.6434,p=0.0012).When the patch area is more than 500 m~2, the patch area had not remarkable correlation with genetic diversity of R.rugosa, which suggested genetic diversity of R.rugosa might be low in big patch.The number of herbaceous species had remarkable correlation with genetic diversity of R.rugosa (r=0.3927,p=0.0387),which indicated herbaceous plants could improve little circumstance in patch,and it was of advantage to success of sexual propagating.The correlative analysis between spatial distance and genetic distance showed genetic distance increased remarkably with the increasing of spatial distance.It suggested there were close relation in neighboring patch,and there were obvious genetic differentiation between patches.
RAPD(random amplified polymorphic DNA) markers were applied to detect the genetic diversity of four remaining large natural populations and to discuss an effective conservation strategy of the endangered species R.rugosa in China.High genetic variations were detected in R.rugosa populations of China.The mean percentage of polymorphic loci(P%) within four local populations was 57.99%,and the percentage of polymorphic loci(P%) of the total population was 75.30%;the mean Shannon's information index(H_0) was 0.2826,and the total Shannon's information index(H_0) was 0.3513.The genetic differentiation among populations (G_(ST)) was 0.1878,which indicated most genetic diversity occurs within populations. Population Tumenjiang(TMJ) showed the highest genetic diversity(P%=66.27%, H_0=0.3117),and contained two exclusive bands;population Changshandao(CSD) showed higher genetic diversity(P%=59.04%,H_0=0.3065).Population TMJ contained 95.33%,and population CSD contained 99.33%of loci with moderate to high frequency(p＞0.05) of the total population.These results indicated that population TMJ and CSD should be given priority for in situ conservation and regarded as seed or propagule sources for ex situ conservation.The study also suggested that R.rugosa in China had become endangered as a result of human actions rather than genetic depression of populations;thus,human interference should be absolutely forbidden in R.rugosa habitats.This study can be referenced to establish conservation strategies for other coastal endangered species accompanied with R.rugosa,such as Vitex trifolia L.var.simplicifolia Cham.,Glehnia littoralis F. Schmidt ex Miq.,Scutellaria strigillosa Hemsl..
引文
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