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盐胁迫下雷竹的离子响应及生理生态变化
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摘要
土壤盐化对农业的威胁是一个全球性问题,在世界范围内,灌溉土地的1/3受到盐分胁迫,且土壤盐化面积逐年增加,从而给农业生产造成重大损失。因此,选育耐盐植物品种、提高植物耐盐性,合理进行土壤化学改良和生物治理,促进农业发展,已成为迫切需要解决的重大课题。雷竹(Phyllostachys praecox cv. prevernalis)是中国特有的优良栽培笋用竹种,主要栽培区域集中在浙江省,由于投资回收周期短、效益好、适应性强,已在浙江省及其他省份广泛栽培,且发展到浙江省的上虞、慈溪、三门、乐清等县市的海涂种植,但由于海涂的土壤盐份含量较高,若不抗盐措施,则绝大部分雷竹林生长缓慢,甚至衰败死亡。为了解决这一难题,很有必要研究雷竹在盐胁迫下的生理生态变化和耐盐机制,以探索解决之途径。
     本论文结合前人研究成果,以雷竹作为研究主体,通过盆栽和人工盐胁迫处理,对其在盐胁迫条件下的离子响应和生理生态变化进行了系统研究,主要测定了雷竹在盐胁迫条件下对阳离子的选择性吸收和运输能力以及在体内的积累水平、根系活力和膜透性变化、渗透调节物质变化及光合作用的影响。
     研究结果表明:
     (1)离子选择性吸收能力随着盐胁迫浓度的升高而升高,在0.3%NaCl溶液处理时达到高峰,随后又开始下降,在处理后的第6天,离子选择性吸收能力较强,但随着处理时间的延长,其选择性吸收能力迅速下降,并且适当施用钙肥能缓解其受盐胁迫的伤害程度。
     (2)离子选择性运输能力ST(根/鞭)随着土壤盐分含量的升高而上升,在0.3%NaCl处理到达最大后又下降。除了0.5%处理外,ST(鞭/枝)极弱。随着盐胁迫时间的增加,ST(根/鞭)和ST(鞭/枝)的变化规律基本一致,即呈低-高-低-高的变化规律;在0.3%处理下,前期有较高的ST,另外0.1%和0.3%处理下,在前期ST基本都比0.5%处理的高。
     (3)盐胁迫处理下,不同器官Na~+,K~+的含量是不同的,K~+的绝对含量远高于Na~+的含量;在0.1%和0.5%处理时,Na~+、K~+含量的变化刚好呈现拮抗作用;从竹叶的含量分析,Na~+、K~+含量的变化规律比较明显,所有的处理K~+含量在第12天时达到最高,随后逐渐下降;竹鞭和竹枝的Na~+、K~+含量变化规律没有竹根和竹叶明显,在最后一次取样分析时,Na~+含量都在0.5%胁迫的处理最高。
     (4)不同盐胁迫下雷竹都保持了较高的超氧化物歧化酶(SOD)活性成分,且在胁迫后期SOD活性成分高于对照或与对照持平。0.5%处理的后期叶片中过氧化物酶(POD)活性成分和对照相比略有下降;0.3%处理的POD活性成分在中期高于对照,后期和对照持平。脯氨酸(Pro)含量在低浓度盐胁迫下最高,然后随着处理浓度的增加而下降,其中0.3%处理的Pro含量与对照基本一致。丙二醛(MDA)的含量与对照差异不显著,其中在后期0.1%胁迫下MDA的含量较对照低。不同处理条件下,各项生理生化指标间的相关性不显著。综合来看,0.3%处理下SOD、Pro和POD活性均保持较高水平,能抑制膜脂质过氧化作用,并使MDA含量保持在较低水平。
     (5)盐胁迫下根系活力对照为最高,其次是0.3%处理,0.5%处理为最低。在盐胁迫的前期雷竹对盐胁迫有一定的应激能力,根系能够保持较高的活力,但随着盐胁迫的加重和时间的推移,根系受到损伤,活力下降。
     (6)细胞膜透性随盐胁迫浓度的变化而变化,对照处理的细胞膜透性最小,0.1%和0.3%处理的细胞膜透性基本一致,而0.5%处理的细胞膜透性明显上升。在盐胁迫前期细胞膜透性的变化比较剧烈,而后期就比较平稳。
     (7)在0.1%处理下叶绿素含量略有增加,但随着浓度的提高,叶绿素含量呈下降的趋势;另外随着盐胁迫处理时间的延长,不同浓度处理的叶绿素含量基本呈高-低-高的变化趋势,但高浓度处理的后期,叶绿素含量升高不明显,但用0.3%浓度处理升高值尤为明显。
     (8)随着盐胁迫浓度的上升,净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)呈下降趋势,胞间CO_2浓度(Ci)与气孔限制值(Ls)的变化规律不明显,但两者呈相反的变化趋势;综合分析后认为雷竹在中低浓度盐胁迫下导致Pn下降主要是气孔限制,而在高浓度盐胁迫下主要是非气孔限制,在盐胁迫的前期导致Pn下降主要是气孔限制,而中后期主要是非气孔限制。
     (9)雷竹的Pn日变化呈“单峰”曲线,并都在11∶0 0达到最高值,但在11∶00以后对照处理的Pn都比其他3种浓度盐胁迫处理的要高。在7∶00到15∶00之间,3种浓度盐胁迫处理雷竹的Pn与Ls日变化相一致,呈先上升后降低的趋势,而Ci日变化刚好相反。水分利用效率(WUE)日变化基本一致,也呈“单峰”变化。
Soil salinization,threating the development of agriculture, is a global problem. One third of the irrigated land worldwide undergoes salt stress and the soil salinity area increases year by year,which causes heavy losses to agricultural production. Therefore,to select and cultivate salt-tolerant plants species,improve salt tolerance and soil physiochemical properties by chemical modification and biological methods,becomes a major issue fpr agricultural development and needs to be urgently resolved.
     Phyllostachys praecox cv. prevernalis(Ph. praecox)is an endemic bamboo species for shoot in China. The major cultivation region focuses in Zhejiang province and extends to other provinces owing to the short payback period,good benefits and strong adaptability. It is even widely cultivated in the coastal land of Shangyu,Cixi,Sanmen and Yueqing County in Zhejiang province. However,the salt content in the soil of coastal land is too high to make them grow regularly and some of them have tended to decline or even death. Therefore, the physiological ecological changes under salt stress and mechanism of salt resistance of Ph. praecox were studied in order to resolve this challenge.
     Ion response and physiological ecological changes of Ph. praecox under salt stress were systematically studied combined with the achievements before. The ionic selective absorption and transportation,accumulation,root activity,membrane permeability,osmotic adjustment and the photosynthesis of Ph. praecox under salt stress conditions are determined.
     The results showed as follows:
     (1)The ionic selective absorption capacity increaseed with salt stress concentration,peaked at 0.3% NaCl,and then declined. 6days after treatment,the ionic selective absorption was strong and then declined rapidly with the extension of the treated time. Moreover, appropriate calcium fertilizer could alleviate the damage of salt stress.
     (2)The ionic selective transportation of ST(root/rhizome)increaseed with salt in soil,and top at 0.3% NaCl treatment,then declined. That of ST(rhizome/branches)was very weak except 0.5% NaCl treatment. With the treatment days of salt stress,the change trend of ST(root/rhizome)and ST(rhizome/branches)was similar,that was,low- high-low-high. High ST was found at early time under the 0.3% NaCl treatment. Besides,ST at earlier time under 0.1% and 0.3% treatment was higher than that under 0.5%.
     (3)Under salt stress,the distribution of Na~+ and K~+ in different parts of the plant was different. The content of K~+ was far more higher than that of Na~+. The changes of Na~+ and K~+ content were just rendering antagonistic effects under 0.1% and 0.5% NaCl treatment. Each Na~+ and K~+ content in bamboo leaves showed obvious change law. K~+ content of treatment reaches the highest level by the 12th day,and then declined gradually. The change law of Na~+ and K~+ content in rhizome and branch was less regular than that in root and leaf. Na~+ content under 0.5% salt stress was the highest in the last sampling analysis.
     (4)Ph. praecox under different salt stress maintained high superoxide dismutase(SOD)activity,even higher or similar than the control by the later stage of treatment. The peroxidase(POD)activity under 0.5% NaCl treatment in leaf at later stage decreased slightly compared to the control,while higher and similar than the control at the medium stage and later stage under 0.3% treatment. Proline(Pro)content showed the highest level under low concentrations of salt stress,then declined with increase of concentration. Pro content under 0.3% treatment was similiar with the control and as well as malondialdehyde(MDA)content. MDA content under 0.1% treatment at the later properties stage was lower than control. The correlation between physiological and biochemical indexes was not significant under different treatments. Activity of SOD and POD and content of Pro under 0.3% treatment could maintain a high level thus inhibited membrane lipid peroxidation and kept the content of MDA at lower level.
     (5)Under salt stress,root system activity of the control was the highest,followed by 0.3% and 0.5%. Ph. praecox showed a certain degree of stress ability at the early stage of salt stress with high root system activity. However,the root system had been damaged and its vigor decreased with the increase of salt concentration and days of treatment.
     (6)The permeability of cell membrane varied with the concentration of salt stress. That of the control showed the minimal,and of 0.1% and 0.3% treatment was similar,and of 0.5% treatment increased significantly. The change of cell membrane permeability at early stage of salt stress treatment was evident,and stable at the later stage.
     (7)Chlorophyll content slightly increased under 0.1% treatment,and declined with the increase of salt concentration. Besides,chlorophyll content of different treatments showed trend of high-low-high with the days of treatment. However,the increase of chlorophyll content was not significantly increased at later stage of high concentrations treatment while increased obviously under 0.3% treatment.
     (8)Net photosynthesis rate(Pn),stomatal conductance(Gs)and transpiration rate(Tr)declined with the increase of salt concentration. The change law of intercellular CO2 concentration(Ci)and stomatal limit value(Ls)was not obvious,but in contrary change trend. Analysis demonstrated that the main cause leading to decline of Pn of Ph. praecox was stomatal limit under low concentration of salt stress at early stage,while non-stomatal limit under high concentration at later stage.
     (9)Pn diural changes of Ph. praecox showed sigl peak curve and reached maximum at 11:00. However,Pn of the control was higher than that of other three salt stress treatments after 11:00. The diurnal change of Pn and Ls of Ph. praecox under three treatments from 7:00 to 15:00 was similar,showing the trend of rising firstly and declined later,while the diurnal change of Ci showed an opposite trend. Diurnal variation of water use efficiency(WUE)was sigle peak curve.
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