外来植物野茼蒿的入侵生物学及其综合管理研究
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摘要
野茼蒿(Crassocephalum crepidioides)是中国境内的一种外来入侵植物,为有效地综合管理该外来入侵种,我们在实验和野外自然条件下开展了该种的繁殖生物学、抗逆生理学、光合生理、入侵地种群生态位、化学他感作用的实验研究,并测定了其重金属元素含量,旨在预测该种在我国境内可能的生境和分布范围,为该种的有效治理和利用提供技术支撑。主要结论如下:
     1、野茼蒿的种子几乎没有休眠期,成熟后即可萌发,且萌发率极高,达到100.00%。在常温条件下储藏一年后,但种子仍然保持活力。这些特征可能使野茼蒿种子在不适宜时期萌发,导致大量幼苗死亡,从而限制了其种群的扩张;
     2、野茼蒿是一种典型的需光种子,这预示对土壤进行深耕,可遏制野茼蒿种子的萌发和出苗,从而达到防除该外来入侵植物的目的;
     3、野茼蒿种子对NaCl盐浓度的耐受性较差,当环境盐浓度高于0.05mol·L~(-1)时种子萌发就明显受到限制,当环境盐浓度高于0.15mol·L~(-1)时,其萌发率极低,种子几乎不萌发;
     4、野茼蒿种子对环境的酸碱性适应范围较宽,pH值为4.00~10.00时,其种子萌发率均高于82.67%;
     5、野茼蒿种子在15℃~30℃范围内,萌发率高达98.00%,在35℃高温或10℃低温的环境中,依然有少量萌发。相比较于低温环境,高温对野茼蒿种子萌发率的抑制作用更大;
     6、野茼蒿种子能够在相对干旱的环境中萌发,在土壤湿度5%时,最终萌发率仍然达到66.67%;
     7、野茼蒿等菊科杂草种子的风扩散试验表明,野茼蒿及其菊科伴生杂草的瘦果大多分布在母株4m以内的范围,但也有少部分扩散到4m以外的范围;
     8、野茼蒿具有较高的光合效率(19.98μmol CO_2·m~(-2)·s~(-1))、光补偿点(20.16μmol·m~(-2)·s~(-1))和饱和点(1311.88μmol·m~(-2)·s~(-1)),是一种比较典型的阳生杂草。比较发现,野茼蒿对光照的适应幅不如小飞蓬(Conyza canadensis)和一年蓬(Erigeron annuus)。
     9、试验表明,低于或等于0.14mol.L~(-1)浓度的NaCl范围内,野茼蒿植株叶片内MDA含量与对照组没有明显差异,说明野茼蒿植株能够适应低浓度NaCl(0.14mol.L~(-1))盐胁迫。在轻微的盐胁迫下,野茼蒿植株在纵向生长受到抑制的情况下,可能通过增加叶片数量,使自身产生更多枝芽。
     10、5℃低温或42℃高温处理野茼蒿48h后,与对照相比,野茼蒿叶片内的MDA、可溶性糖、可溶性蛋白及POD酶活性均没有明显差异,反映出野茼蒿对高温胁迫不敏感,也预示着该外来植物在我国有广阔的潜在分布范围。
     11、草甘膦和百草枯作用下,野茼蒿叶片的MDA、Pro含量和POD活性增高,而可溶性蛋白含量和SOD活力降低,表明草甘膦和百草枯均使野茼蒿细胞脂膜受损,蛋白质合成受阻,导致其生长严重受抑。应用这些生理指标,能够在实验室条件下快速地比较野茼蒿对不同除草剂的敏感程度;使用1/2倍推荐剂量的百草枯或草甘膦就可以达到控制野茼蒿的目的。
     12、在浙江金华北山地区的野茼蒿入侵地,野茼蒿的生态位宽度为0.4305,位于伴生杂草中的第7位,与野茼蒿生态位重叠最大主要杂草有铁苋菜(Acalypha australis)、马唐(Digitaria sanguinali)、一年蓬(Erigeronannuus)、狗尾草(Setaria viridis)、爵床(Rostellularia procumbens)、睫毛牛膝菊(Galinsoga parriflora)、龙葵(Solanum nigrum)、藜(Chenopodiumalbum)、鳢肠(Eclipta prostrata)、苋菜(Amaranthus tricolor)、稗草(Echinochloa crusgalli)、野艾蒿(Artemisia lavandulaefolia)和山莴苣(Lagedium sibiricu)等。就调查的样地而言,野茼蒿的出现频率和生态位宽度均比常见杂草一年蓬、狗尾草、马唐要低,说明其杂草性不强。
     13、野茼蒿茎和叶浸提液中含有化学他感作用物质,对其他植物有一定的抑制作用,但能促进野茼蒿幼苗的生长,说明野茼蒿种子形成了适应自身分泌的化感物质某种机制,这种机制可有效的保护其后代,为扩大种群奠定基础。
     14、对4种重金属元素分析结果表明,野茼蒿对某些有毒重金属原始具有一定的富集效应。但从总体上分析,在比较的24种野生蔬菜中,野茼蒿的食用风险处于第16位,具有一定的食用风险。
     综上所述,野茼蒿对温度、pH值、湿度等具有较宽的耐受幅;虽然模拟盐份胁迫逆境条件下的生理指标测定表明野茼蒿能够适应轻微的盐份胁迫环境,但是盐环境下野茼蒿种子萌发受到明显抑制,总体上,野茼蒿不适于盐碱地和海边滩涂环境。同时,野茼蒿对光照的适应幅不如小飞蓬、一年蓬,对环境资源的利用率(即生态位宽度)也不如加拿大一枝黄花(Solidago canadensis)、一年蓬、狗尾草和马唐等广阔。因此,与其入侵地其它常见的伴生杂草相比,野茼蒿属于入侵性不强的一类外来植物。在防除上,经常翻耕可以有效地遏制野茼蒿种子的萌发和出苗,选用百草枯可以快速、高效地触杀野茼蒿植株体。如果作为野生蔬菜,由于其对某些重金属原始具有一定的富集效应,因此食用时应注意其可能存在的安全问题。
Crassocephalum crepidioides is a new exotic plant in China.In order to effectively control and utilize the exotic species,predict its potential distribution range and habitats in China,we studied its reproductive biology,stress physiology,population niche,Allelopathy,and also determined the content of the heavy metal elements in the plants and its nutrient values.The main results are as follows:
     1.The seed s of C.crepidioides almost have no dormancy period,and are able to germinate immediately after their mature,with a germination rate as high as 100%.Their seeds,even saved for one year in dry,still kept alive.Compared with the seeds of other weeds with dormancy,those of C.crepidioides could germinate even and quickly in favorable environments,which will lead to death of a great number of seedlings if they meet unfavorable conditions later,and thus limit their populations expansion;
     2.Light is necessary for the seeds of C.crepidioides to germinate,which indicate that appropriate soil tillage,mainly by ploughing will reduce the germination of C.crepidioides and thus control this exotic invasive plant;
     3.The seeds of C.crepidioides are sensitive to NaCl,their germination was inhibited when treated with NaCl at a concentration of 0.05 mol·L~(-1).Almost no seeds of C.crepidioides were able to germinate when treated with NaCl at a concetration of 0.15 mol·L~(-1);
     4.Relative wider adaptation range for the seeds of C.crepidioides to environmental pH conditions was observed during our experiments.Their seeds could germinate with a percentage of 82.67%when treated in the solution with pH from 4.00~10.00;
     5.Treated under 15℃~30℃,the seeds of C.crepidioides had higher germination rate(≥98.00%),even treated under 35℃or 10℃,some seeds still germinated.Compared with that at lower temperature,the inhibitory effect on their seed germination at higher temperature is stronger;
     6.The seeds of C.crepidioides are able to germinate in drier conditions, with a germination rate as high as 66.67%in soils with 5%humidity;
     7.Wind spreading experiment showed that the seeds of C.crepidioides and other five Compositae species could be spread to a range of 4 m or more far from the mother plants;
     8.Crassocephalum crepidioides has higher net photosynthesic rate(19.98μmol CO_2·m~(-2)·s~(-1)),light compensate point(20.16μmol·m~(-2)·s~(-1)),light saturation point(1311.88μmol·m~(-2)·s~(-1)),showing its adaptation to sunny environments.
     9.Experiments showed that MAD content in the leaves of C.crepidioides treanted with 0.07 or 0.14 mol·L~(-1)NaCl concentration was not significantly different from that of control,indicating that C.crepidioides is able to grow the environment with NaCl at lower concentraion.When slightly stressed by NaCl, The growth of C.crepidioides in height was inhibited by NaCl at lower concentration,but could be compensated by the increase of leaf number and producing more branches.Such response of its stems and leaves to lower NaCl stress facilitates the species to grow in the environments with NaCl at lower concentration.
     10.Treated under 5℃or 42℃for 48h,the content of MDA,soluble sugar and soluble protein,as well as the activity of POD in the leaves of C. crepidioides are not significantly different from those of the control,indicating that C.crepidioides is not very sensitive to high temperature(42℃)or low temperature(5℃).Therefore,this exotic plant might have wider potential geographic distribution range in China.
     11.Treat with either paraquat or glyphosate,the content of MDA and Proline,as well as the POD activity increased,while the content of soluble protein and the SOD activity decreased,which indicated that both paraquat and glyphosate caused obvious damages to their cell membrane,inhibit the formation of relative protein,thus inhibit its growth.The application of paraquat or glyphosate with a concentration of 1/2 dose recommended could control the species effectively;
     12.In the areas(North Mountain,Jinhua,Zhejiang province)invaded by C. crepidioides,C.crepidioides has a niche wide of 0.4305,ranked as 7~(th)among the species in the surveyed sites.The accompanied weeds,according to their niche overlaps with C.crepidioides from high to low,are Acalypha australis, Digitaria sanguinali,Erigeron annuus,Setaria viridis,Rostellularia procumbens,Galinsoga parriflora,Solanum nigrum,Chenopodium album, Eclipta prostrata,Amaranthus tricolor,Echinochloa crusgalli,Artemisia lavandulaefolia,and Lagedium sibiricu,ect.Niche analyses showed that both of frequency and niche width of C.crepidioides are lower than those of Erigeron annuus,Setaria viridis,Digitaria sanguinali,which shows that the weediness of C.crepidioides is not very strong.
     13.The aqueous extracts from the leaves and stems of C.crepidioides contain some kinds of allelochemicals,which could inhibit the seed germination and seedling growth of other species,but had no inhibitory effects on or even facilitate its own growth in young stage,which might protect its own young generation from competing with other accompany species,thus facilitate to expand it population.
     14.Determination on the content of heavy metal elements in C. crepidioides and other 23 wild vegetables showed that the risk of damage to our health as a vegetable ranked 16 among the 24 wild vegetables.Attention should be paid to the harmful effects on our health when we utilize this plant as a vegetatbel.
     Together the above experimental results,we found,C.crepidioides is able to grow under wider range of temperatures and pH values in soils. Crassocephalum crepidioides has narrower range of illumination than comnyza Canadensis,Erigeron annuus,and the niche width of C.crepidioides is than those of Erigeron annuus,Setaria viridis,Digitaria sanguinali,Therefore,the invasiveness of C.crepidioides is not stronger than thoses of other common weeds and worst invasive exotic plants.For control purpose,appropriate soil tillage,mainly by ploughing will reduce the seed germination and the seedling emergence of C.crepidioides,and the treatment with paraquat will control the species quickly and effectively.Because of the accumulated effect of some toxic heavy metal elements,attention should be paid to the risk of C.crepidioides for our health if the plant is utilized as a wild vegetable.
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