摘要
土荆芥(Chenopodium ambrosioides L.)为藜科藜属一年生或多年生草本植物。有强烈芳香味,通常生长在河岸、公路边、荒地及农田边,是一种入侵植物,常与一年蓬、黄花蒿等植物混生在一起形成杂草群落。目前全国各地均有分布。但各地其情况又有不同,有些地方(如长江流域一带)土荆芥已入侵成为优势种,阻碍了本土植物的生长繁殖,并给当地农业造成了一定危害。彻底根除土荆芥,控制其危害和扩散蔓延已成为这些地方的农业要求。目前对入侵植物土荆芥的研究仅限于其化学成分、药用价值、种子贮藏与萌发特性等方面；对其入侵机理方面的研究基本还是一片空白。因此,以同属本土近缘种藜作对比,研究其在逆境条件下的生理适应机制对于了解入侵机理、提出有效的防治对策等具有重要的理论与实践价值,也能为其它入侵植物的研究和防除提供参考。
藜(Chenopodium album L.)是跟土荆芥亲缘关系较近的一种难以除掉的本土杂草,为藜科藜属一年生草本植物,常见于路旁、荒地及田间。本文以藜作为实验对照,在盐、人工模拟酸雨、重金属镉等胁迫因子以及模拟不同程度光照强度的条件下,从形态、生物量分配、抗氧化酶活性、渗透调节物质(脯氨酸、可溶性糖)等方面研究了土荆芥的生长和生理适应机制,以期为入侵植物入侵机制的认识、防治对策的制定等提供一定的理论与实践依据。研究结果表明：
1)人工模拟盐胁迫和碱胁迫下,土荆芥的生长受到了抑制,且处理浓度越高抑制作用越明显。随NaCl盐胁迫程度增加,土荆芥各形态指标数据呈明显下降趋势,根长表现出先增后降的趋势,根重比、根冠比在各处理水平均高于对照；丙二醛含量在低浓度NaCl处理时变化较为平缓,高浓度处理时呈上升趋势,抗氧化酶除SOD活性变化较小,CAT、POD的活性均呈现出先增后降的趋势。Na2C03碱胁迫时,土荆芥各形态指标的变化趋势和抗氧化酶的活性变化趋势与NaCl处理时相似,但根重比和根冠比低于对照,丙二醛含量呈上升趋势,SOD、CAT和POD均具有不同的酶活最高点。两种盐处理对比表明Na2C03胁迫对土荆芥造成的危害更大,土荆芥耐受NaCl胁迫能力强于Na2C03胁迫。近缘植物藜的生长也同样受到了抑制,且SOD、CAT、POD分别具有与土荆芥不同的活性变化以及酶活最高点,但其根重比、根冠比、抗氧化酶活性的变化程度均低于土荆芥,说明土荆芥可能比藜更能耐受盐、碱胁迫。另外,土荆芥脯氨酸在NaCl处理时得到了积累,而藜脯氨酸在低浓度Na2C03处理时有所积累,且仅低浓度的盐、碱胁迫能提高土荆芥和藜可溶性糖的含量,表明脯氨酸和可溶性糖的渗透调节作用具有一定的局限性。
2)人工模拟酸雨胁迫下,土荆芥的形态变化较小,各酸雨处理水平除根长、叶片数、分枝数存在显著性差异,其它形态指标变化不大。土荆芥根受到的影响较为明显,pH3.5-4.59水平酸雨均促进了土荆芥根的生长,但根重比和根冠比随模拟酸雨pH值的减小显著下降,叶重比有上升趋势。随模拟酸雨胁迫程度的加深,土荆芥丙二醛含量增加,SOD活性呈小幅度上升趋势,CAT活性呈下降趋势,POD活性则表现为先升后降呈单峰曲线型变化。酸雨对藜的形态变化影响也较小,除pH2.0酸雨外,其它水平酸雨均促进了藜根的生长。与土荆芥不同的是,pH2.0处理组藜的根重比和根冠比高于对照,其原因不明。除SOD活性变化与土荆芥一致,藜CAT和POD活性分别具有与土荆芥不同的变化趋势,且在酸雨pH4.59时,三种酶的活性均高于对照,表明藜对当地pH4.59的酸雨已具有一定的适应性,但强酸雨条件下土荆芥SOD、CAT、POD活性变化仍高于藜,说明土荆芥比藜具有更高的抗逆能力。酸雨处理下土荆芥脯氨酸含量变化不大,酸度较小的模拟酸雨能有效促进藜脯氨酸的积累,但二者可溶性糖的合成均受到了抑制。
3)人工模拟重金属镉胁迫下,土荆芥和藜生长受到了严重的抑制,形态和生物量的分配发生了较大变化。土荆芥的茎高、叶片数、分枝数存在明显下降趋势,根长、根重比、根冠比表现出先增后降的趋势。藜除叶长外,各形态指标均存在显著性差异,其根长也有先升后降的趋势,但其根重比、根冠比变化不大。随镉胁迫程度的增加,土荆芥和藜MDA含量呈现先增后降的趋势,高浓度镉处理下MDA含量降低说明镉胁迫已超出细胞抵御的最大能力。土荆芥和藜SOD、POD活性变化不稳定,CAT活性呈下降趋势,表明酶系统的功能已失调或无法再激活。低浓度镉胁迫能有效促进二者脯氨酸的积累,土荆芥更为明显：二者可溶性糖含量随镉浓度增加显著升高,此时藜可溶性糖的积累更为明显。镉浓度为10mg·mL-1时,土荆芥的根长、根重比、根冠比、脯氨酸增加幅度达到最大,对镉胁迫表现出了一定的抗性。
4)在人工模拟不同光照强度(遮阴)条件下,土荆芥和藜生长受到了严重的影响,三层遮阴处理的植株其生长已完全受到抑制。二者各形态指标数据随光照强度的降低均呈下降趋势。土荆芥根重比和根冠比有先增后降的趋势,在一层遮阴处其根重比和根冠比达到最大,二层遮阴处即低于对照。藜根重比、根冠比随光照强度降低而减小。二层遮阴时,土荆芥和藜MDA含量开始下降,表明二者受到严重伤害,已超出细胞抵御的最大能力。除对SOD活性影响较小,弱光抑制了土荆芥和藜CAT、POD的活性。光照强度的降低促进了土荆芥脯氨酸和可溶性糖的积累,但藜渗透调节物质积累具有一定的局限性,藜脯氨酸含量呈现出先降后升的趋势,可溶性糖含量呈现出先增后降的趋势,前者在光照强度极低的情况下才大量积累有可能是脯氨酸生成失调。一层遮阴处理时土荆芥根重比和根冠比达到最大,对光照强度降低仍作出了一些反应,表现出了一定的抗性。
综上结果表明,土荆芥具有一定的表型可塑性,会改变自身的形态和生理特征来响应环境的变化；对逆境具有较强的抵抗能力,比藜具有更高的生理调节水平,证明了入侵植物的优势在于其本身具有较强的适应性和忍耐性,使其能快速适应新环境。在逆境条件下,土荆芥各指标的变化并不同步,其抗性很难用单一的指标来衡量,在某个时期某个指标占了主导地位。鉴于土荆芥具有抗低浓度的盐、镉能力,将其用作盐化草甸建群草种以及镉污染土壤的修复具有重要的现实意义。
Chenopodium ambrosioides L. is the annual or perennial plant of Chenopod. It is an invasive plant, having strong aroma and usually growing on the riverside, road, badlands or cropland, and always forms ruderal community with Erigeron annuus L.and Artemisia annua L.It can be seen all the country, but in some places (like drainage area of the Changjiang River), it has become preponderant species, disturbing growth and propagation of the native plants, and bringing harms to native agriculture. Eradicating C.ambrosioides L.thoroughly and controlling its damage and spread has turned into these places'agricultural requests. At present, the research of C.ambrosioides L. is still nothingness, only having researches in its chemic components and officinal value, and starting pilot study on the seed storage and germination. So for managing this invasive plant effectively, it's meaning to investigate the adaptability of growth and physiology in adversity, and the result also can provide new theoretical method and practical experience for mechanism of biological invasion.
Chenopodium album L. is a sort of weed which is difficult to dispose of, and is the relatives plant to C.ambrosioides L..It is the annual plant of Chenopod, always growing on the roadside, badlands or field. The experiment is taken to research adaptability of growth and physiology of C. ambrosioides L. in comparison with C.album L.n the way of morphological indicators, biomass allocation, antioxidant enzymes activity, and osmotic adjustment substances (proline and soluble sugar), under simulating the salt and alkali stress, acid rain, heavy metal-Cadmium stress, and different light intensity (overshadowing). The results of researches showed that:
Under the simulated salt and alkali stress, the growth of C.ambrosioides L.was inhibited, and this restraining effection was obvious along with extent of intimidation. Under NaCl stress, morphology indicators datas of C.ambrosioides L.decreased, root length increased at first and then decreased, root weight ratio and crown-root ratio were higher than comparison; malondialdehyde content changed gently in low concentrations and gained in high concentrations; SOD activity had changed little, POD and CAT activity had increased firstly and then decreased. Under Na2CO3 stress, trend of morphology indicators and antioxidant enzymes activity of C.ambrosioides L. was similar to that under NaCl stress, but root weight ratio and crown-root ratio were lower than comparison, malondialdehyde content increased all through, and SOD、CAT and POD activity also had different changes. A result between the two stresses showed Na2CO3 stress did more harm to C. ambrosioides L. than NaCl stress. The growth of C.album L.was also inhibited, and its'SOD、CAT and POD activity had different diversifications with C. ambrosioides L. The root weight ratio、crown-root ratio and antioxidant enzymes activity of C.album L. was lower than that of C.ambrosioides L., indicating that C.ambrosioides L. had stronger tolerance to the salt and alkaline stress than C.album L..In addition, the proline of C. ambrosioides L. accumulated under NaCl stress, proline of C.album L.got together in low concentrations of Na2CO3 stress, and soluble sugar of these two species all gained in low concentrations of salt and alkali stress, indicating that the osmotic adjustment function of proline and soluble sugar had definite limitation.
Under simulated acid rain, it had small influence on morphology of C. ambrosioides L.,except root length and the number of laminae and ramification had markedly otherness, other morphology indicators changed little. It seriously affected C. ambrosioides L.'s root, promoted its growth from pH 3.5 to 4.59,but root weight ratio and crown-root ratio decreased clearly as simulated acid rain's pH values declined, besides leaves ratio increased slightly. As stress of acid rain increased, malondialdehyde content increased, SOD activity gained a little, CAT activity declined, and POD activity increased at first and then decreased. Acid rain also had little effect on morphology of C.album L..Except acidity of pH 2.0, other acidity promoted its root growth. Root weight ratio and crown-root ratio of C.album L. were higher than comparison at pH 2.0, but the reason was unaware. In addition, activity of CAT and POD had different trends with C. ambrosioides L., but SOD activity was accordant with it. At pH 4.59, antioxidant enzymes activity was higher than comparison, indicating that C.album L. had been adapted to the local acid rain, whereas, under strong acid rain, the antioxidant enzymes activity of C. ambrosioides L. was higher than that of C. album L., showing C. ambrosioides L. was more tolerant than C.album L. under adversity. Proline of C. ambrosioides L. was influenced a little, but that of C. album L. was gained under weak acid rain, and soluble sugar synthesization of these two species was all restrained.
Under simulated heavy metal-Cadmium stress, the growth of C. ambrosioides L. and C.album L.were inhibited seriously, and their morphology and biomass allocation had obvious changes. Stem height and the number of laminae and ramification of C. ambrosioides L. had dropped, and its root length, root weight ratio and crown-root ratio had increased at first and then decreased. Except leaf length, other morphology indicators of C.album L. also had markedly otherness, the root length also had the trend like C. ambrosioides L., but root weight ratio and crown-root ratio changed little. Their malondialdehyde content still increased firstly and then decreased, reducement of malondialdehyde content under high concentrations of Cd treatment showing Cd stress had exceeded cells'resist capability farthest. Their SOD and POD activity was unstable, CAT activity was degressive, indicating the functiong of enzyme system was maladjusted or unactivated. Low concentrations of Cd can effectively promoted accumulation of proline, specially for C. ambrosioides L., and also availably added soluble sugar, yet specially for C.album L.. At the 10 mg-L'concentration of Cd, root length, root weight ratio, crown-root ratio and proline of C. ambrosioides L. had come to the most, figuring that C. ambrosioides L.had put up some resistance for Cd stress.
Under the simulated different light intensity (overshadowing), the growth of C. ambrosioides L. and C.album L. were inhibited seriously, at the overshadowing of three layers stress, their growth had been completely repressive. Their morphology indicators datas decreased. Root weight ratio and crown-root ratio of C. ambrosioides L. were increased firstly and then decreased, higher than comparison at one layer's overshadowing, but lower than comparison at two layers'overshadowing. By contraries, root weight ratio and crown-root ratio of C.album L. dropped all through. At the overshadowing of two layers stress, their malondialdehyde content declined, showing stress had exceeded cells'resist capability farthest. Low light restrained CAT and POD activity, except SOD activity. Low light intensity promoted proline and soluble sugar's accumulation of C. ambrosioides L., but accumulation of osmotic adjustment substance of C.album L. had some limitation, its proline content had decreased at first and then increased, soluble sugar content had the opposite trend, guessing synthesization of proline had been maladjusted. At the overshadowing of one layer, root weight ratio and crown-root ratio of C. ambrosioides L. had come to the most, showing certain resistance to the decrease of light intensity.
According to these results above, C. ambrosioides L. has some phenotypic plasticity,and can change configuration and physiology in answer to variety of environment; has stronger resistance to adversity and higher physiological adjustment level than C.album L., testifying that invasive plant can quickly adapt to new environment due to its predominances of adaptability and endurance. In adversity, indexes of C. ambrosioides L. are nonsynchronous, the resistance can't measure with single index, and some index is more principal than others in some phase. Since C. ambrosioides L. has the ability of resisting salt and Cd in low concentrations, it's meaning that making use of it as grass of saline and restoration of Cd-contaminative soil.
引文
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