利用拟南芥Na~+/H~+逆向转运蛋白基因（AtNHXl）提高杨树耐盐性的研究
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摘要
盐胁迫是抑制植物生长,导致减产的主要因素之一,并对世界农业的发展构成较大威胁。为适应盐渍环境,植物可以通过把吸收过多的Na+排到细胞外,或者将其区隔化在液泡膜内来降低胞质内的Na+。其中,Na+在液泡内的区域化是通过植物液泡膜Na+/H+逆向运转蛋白来实现的。过量表达液泡膜Na+/H+逆向运转蛋白基因显著提高转基因植株的耐盐性已在多种植物中得到证实,说明过量表达该基因在植物的耐盐性中起着重要的作用。
杨树是我国最重要的用材树种之一,大多数杨树均属非耐盐品种,提高优质杨树耐盐性对我国农业的发展具有重要的现实意义。过量表达Na+/H+逆向运转蛋白基因提高转基因植株耐盐性的研究主要集中在草本和灌木中,而对乔木树种的研究甚少。为提高优质用材林树种欧美107杨的耐盐性,本课题组采用根癌农杆菌介导法,已经成功将拟南芥Na+/H+逆向转运蛋白基因(AtNHX1)转入其中,并证实已获得转AtNHX1基因植株。本研究以欧美107杨非转基因(WT)和转AtNHX1基因植株(TR)为材料,用NaCl、海水和土培加盐对其进行处理,分析盐胁迫对2个株系生长和生理的影响,试图阐明转AtNHX1基因杨树的耐盐机制,揭示AtNHX1基因在杨树耐盐中的作用；通过海水处理,探索转基因植株的耐海水能力,阐明AtNHX1基因与树木耐海水胁迫的关系,拓宽对转基因植物耐盐机理的了解。主要结果如下：
1、经PCR和RT-PCR对转AtNHX1基因欧美107杨的检测,证实AtNHX1基因已整合到欧美107杨基因组中,并进行了表达。
2、用不同浓度NaCl(0、75和150mmol·L-1)浇灌欧美107杨WT和TR幼苗30d,2个株系植株的生长受到不同程度抑制。WT和TR植株生长和生理指标差异显著,TR植株耐盐性显著高于WT植株。在75和150mmol·L-1NaCl处理下,TR比WT植株的苗高相对生长率(RGRH)分别高46%和44%,直径相对生长率(RGRD)高36%的61%；TR比WT维持了更大的叶面积,积累了更多的干物质。
NaCl处理下,TR植株比WT植株具有更高的生长速率和积累更多的干物质主要是由于维持了更高的光合能力。在75和150mmol·L-1NaCl处理5d,TR比WT植株的Pn分别高22%和9%；而在75和150mmol·L-1NaCl处理30d,TR比WT植株的Pn分别高13%和48%。气孔限制是盐胁迫下欧美107杨光合速率下降的主要因素。盐处理导致2个株系的PS Ⅱ最大光化学效率(Fv/Fm)、PS Ⅱ实际光化学量子产量(ΦPSⅡ)、光化学猝灭系数(qP)和电子传递速率(ETR)均下降,而非光化学猝灭系数(NPQ)则有不同程度的提高。但是,在同等NaCl处理下,TR植株比WT植株维持了更高Fv/Fm、ΦPSⅡ、qP、ETR和较低的NPQ。课件,TR植株在盐胁迫下能够更好的保护光系统,使其具有较高的光系统活性和光能转化效率,能将所吸收的光能有效地转化为化学能,提高光合电子传递速率,从而提高生物量,这在一定程度上提高了其抵御盐胁迫的能力。
NaCl胁迫下,活性氧清除剂SOD、POD、CAT活性在TR植株叶片中受盐诱导上调的幅度较大。NaCl处理30d,WT植株中的这些参数则显著低于对照。说明盐胁迫下,TR植株比WT植株能更有效地清除H202。
NaCl胁迫下,2个株系植株体内离子含量存在较大差异。总的说来,在75和150mmol·L-1NaCl处理下,TR植株的根、茎和叶均比WT植株积累了更多的Na+。在同等NaCl处理下,TR植株对K+的吸收明显大于WT植株。例如,在150mmol·L-1NaCl处理下,TR植株根、茎和叶的K+含量比WT植株分别高32%、38%和35%。尽管TR植株较WT植株吸收了更多的Na+,但维持了更高的K+和K+/Na+比率,说明TR植株具有更高的耐盐性,可能是由于对K+具有更好的吸收。
3、用不同浓度的海水(0%、10%、20%和30%)浇灌WT和TR幼苗30d,比较了二者的生长指标、光合参数和离子含量。2个株系植株的生长受到不同程度抑制,但二者的植株生长和生理指标存在显著的差异。在同等海水处理下,TR植株比WT植株具有更高的生长速率、总叶面积、光合速率和Fv/Fm,并积累了更多的干物质。TR植株的组织中比WT植株的积累了更多的Na+和K+,TR植株耐海水能力显著高于WT植株。在10%的海水处理下,TR植株的干重与对照相比并无显著差异,在20%和30%的海水处理下,其干重分别下降了8%和22%。可见,10%～20%海水用于浇灌TR欧美107杨是可行的。尽管如此,海水胁迫对TR植株个体发育的影响尚待进一步的田间耐盐试验研究。
4、以不同浓度NaCl(0、75和150mmol·L-1)浇灌土培盆栽的欧美107杨WT和TR幼苗30d。低盐处理下,WT植株生长显著受到抑制,随盐分强度加大,抑制作用增大,高盐处理下其干重只有对照的50%；而TR植株在低盐处理下干重与对照差异不显著,高盐处理时其干重为对照的74%。同等盐度处理下,TR的干重显著高于WT,且随着盐度升高,2个株系间植株干重差异增大。盐处理后,TR植株叶片叶绿素和类胡萝卜素的含量均显著高于WT,并能维持较高的Pn和Fv/Fm；虽然TR叶片和根系均较WT积累了更多的Na+,但同时也维持了更高的K+和K+/Na+比率,而且叶片对K+选择性的运输明显高于WT；同时,盐胁迫下TR叶片MDA含量和相对电导率显著低于WT。盐对TR叶片叶绿体超微结构的影响较轻,在高盐下仍保持了较好的内部结构。
5、AtNHX1的导入能够显著提高转基因欧美107杨的耐盐性。耐盐转基因杨树的获得也为提高树木耐盐性提供了一种可行的研究方法。本研究结果也验证了转单一的Na+/H+逆向转运蛋白基因是能够明显提高植物耐盐性的。
Salt stress is one of the major stresses that depress plant growth and limit crop production, and has been a great threat to agricultural development all over the word. To cope with salt stress, some plants have the ability of either extruding Na+out of cells or sequestering Na+into the vacuole in order to decrease excessive Na+in the cytoplasm. The compartmentation of Na+into vacuoles is implemented by Na+/H+antiporter located in tonoplast. The importance of vacuolar sequestration to plant salinity resistance has been underlined by experiments in which constitutive overexpression of different vacuolar Na+/H+antiporter genes can greatly increase salinity resistance in a wide range of plant species.
Poplar is one of the most important timber tree species planted in China. Therefore, understanding of the mechanism of its response to salt stress is of paramount importance to agriculture application and plant improvement. Overexpression of Na+/H+antiporters have been reported to increase the salt resistance of various herbaceous or shrub species, but very little is known about the role of these antiporters in the salt resistance of trees. Successful Agrobacterium-mediated transformation of AtNHXl to the shoot stem explants of Populus×euramericana 'Neva' and regeneration of transgenic plants has been previously performed. In order to elucidate mechanism of overexpressing the AtNHXl gene response to salt-tolerance in woody plants, the wild-type plants (WT)(Populus×euramericana'Neva') and its transgenic varieties (TR) that overexpress the AtNHX1gene were investigated to identify their growth and physiological character responses to salt stress. Moreover, the WT and TR were also exposed to different concentrations of seawater to clarify the correlation between overexpressing the AtNHXl gene and seawater-tolerance in poplars. All these would help to develop strategies for improving the understanding of transgenic plants response to salt stress. The main results obtained were shown as follows:
1. All tests using PCR and RT-PCR indicated that the AtNHX1gene was integrated into the genome of Populus×euramericana 'Neva' and transcribed in transgenic poplar plants.
2. Seedlings of WT and TR were subjected to different concentrations of NaCl (from0to150mmol·L-1) stress for30days. Plant growth was inhibited by salinity in both WT and TR but in different degrees, and salt-resistance of TR was much higher than that of WT. The relative growth rate of stem diameter (RGRH) values of TR were46%and44%and the relative growth rate of shoot height (RGRD) values were36%and61%more than that of WT at75and150mM NaCl treatments, respectively. Moreover, TR produced much more total leaf area and dry weight than WT.
Under the same NaCl level treatments, TR maintained higher RGR, total leaf area, and dry weight than WT because TR had a higher Pn. The Pn values in TR were22%and9%higher than that of WT when grown at75and150mmol·L-1NaCl for5days, and13%and48%higher for30days. Stomatal limitation was the primary factor limiting Pn in Populus×euramericana 'Neva' under salt stress. At the same time, maximum photochemical efficiency of PS Ⅱ (Fv/Fm), actual quantum yield of PS Ⅱ (Φps Ⅱ), photochemical quenching coefficient (qP) and relative electron transport rate (ETR) in both WT and TR decreased while non-photochemical quenching coefficient (NPQ) increased in various degrees. However, under the same level of NaCl treatment, TR maintained higher Fv/Fm, ΦpsⅡ qP, ETR and lower NPQ than those of WT. These suggest that TR could better protect PSⅡ from injury induced by the salt stress. So it had higher PSⅡ activity and light energy transform efficiency, which could turn the absorbed light energy into chemical energy effectively, and then increased the transmitting rate of photosythetic electrons, thereby enhancing biomass. To some extent, the propertites of TR decribed above improved salt resistance.
The activity of antioxidant enzymes such as SOD, POD and CAT were induced to increase greatly in the leaf of TR. To WT. salt stress for30days induced obvious decline on the activity of these enzymes. These indicate that TR was likely to be more effective in removing H2O2than WT under salt stress.
Differences in the pattern of K+accumulation between WT and TR were apparent under NaCl stress. In general, TR accumulated more Na+in the root, stem, and leaf than WT at both75and150mmol·L-1NaCl concentrations. The K+uptake in TR was much more than that in WT under the same NaCl level. For example, the K+contents of TR in the root, stem, and leaf were32%,38%, and35%more than those of WT under the150mmol·L-1NaCl treatment. Despite a higher accumulation of Na+in TR, it maintained relatively higher K+and K+/Na+ratio compared to WT. Results show that plants that TR was markedly more resistant to NaCl, possibly because of intracellular K+content regulation.
3. Seedlings of WT and TR were exposed to different seawater concentrations (from0to30%) for30days to determine the effects of seawater on seedling growth, photosynthetic productivity, and ion content. Plant growth was inhibited by seawater in both WT and TR but in different degrees, and seawater-resistance of TR was much higher than that of WT. TR could maintain higher RGR, total leaf area, Pn, and Fv/Fm, so that it is able to accumulate more dry mass than WT at the same seawater level. TR accumulated more Na" and K+contents in plant tissue than WT. At10%seawater treatment, the dry weight of TR was unaffected, but decreased by8%and22%at20%and30%seawater treatments, respectively. These findings indicate that10%-20%seawater could be used to irrigate TR plants, thus implying the viable utilization of seawater. However, further investigations of the effects of stresses relative to transgenic poplar ontogeny are necessary to assess realistic stress pressures that occur naturally in the field.
4. Seedlings of WT and TR which cultivated in pot with soil were subjected to different concentrations of NaCl (from0to150mmol·L-1) stress for30days. We investigated the salt-resistance of the WT and TR plants by measuring growth parameters, chlorophyll (Chl) and carotenoid (Car) content, Pn, ion content, leaf malondialdehyde (MDA) and electrolytic leakage. Compared with the control, the growth of WT was restrained significantly in the presence of both low salt and high salt. The dry weight of WT was significantly lower under salt stress than that of control, the dry weight decreased gradually with increasing NaCl concentration and their dry weight under high salt was just50%of the control. However, the dry weight of TR in low salt was similar to the control, up to the high salt treatment, where their dry weight was74%of the control. Moreover, the dry weight of TR was significant higher than that of WT in both NaCl treatments, and the discrepancy of dry weight was increased with increasing NaCl concentration. In the presence of NaCl, Chl and Car content of TR were significantly higher than that of WT, and the TR maintained a remarkably high Pn and Fv/Fm. Although TR accumulated more Na+in their roots and leaf tissues under salinity conditions compared with the WT, they absorbed more K+and maintained a higher K+/Na+ratio. Moreover, these TR plants kept a lower MDA and electrolytic leakage level than that of wild-type. Under salt stress, the chloroplasts of TR under low salt stress were not different from the control, and the structure of chloroplasts under high salt was better than that of WT. Under the same salinity level, the TR can maintain a relative integrated chloroplast, high chlorophyll contents and preferable net photo synthetic rate, so its salt tolerance were improved.
5. These findings indicate that transformation of AtNHXl gene into Populus×euramericana 'Neva' can confer plants more tolerance to salinity than its wild-type plants. These transgenic poplars provided a feasible way for improving important trees to adapt to salt stress conditions that are common in saline and arid soils. Our result also proved that transformation of single AtNHXl gene into plants can confer plants more resistance to salinity.
杨树是我国最重要的用材树种之一,大多数杨树均属非耐盐品种,提高优质杨树耐盐性对我国农业的发展具有重要的现实意义。过量表达Na+/H+逆向运转蛋白基因提高转基因植株耐盐性的研究主要集中在草本和灌木中,而对乔木树种的研究甚少。为提高优质用材林树种欧美107杨的耐盐性,本课题组采用根癌农杆菌介导法,已经成功将拟南芥Na+/H+逆向转运蛋白基因(AtNHX1)转入其中,并证实已获得转AtNHX1基因植株。本研究以欧美107杨非转基因(WT)和转AtNHX1基因植株(TR)为材料,用NaCl、海水和土培加盐对其进行处理,分析盐胁迫对2个株系生长和生理的影响,试图阐明转AtNHX1基因杨树的耐盐机制,揭示AtNHX1基因在杨树耐盐中的作用；通过海水处理,探索转基因植株的耐海水能力,阐明AtNHX1基因与树木耐海水胁迫的关系,拓宽对转基因植物耐盐机理的了解。主要结果如下：
1、经PCR和RT-PCR对转AtNHX1基因欧美107杨的检测,证实AtNHX1基因已整合到欧美107杨基因组中,并进行了表达。
2、用不同浓度NaCl(0、75和150mmol·L-1)浇灌欧美107杨WT和TR幼苗30d,2个株系植株的生长受到不同程度抑制。WT和TR植株生长和生理指标差异显著,TR植株耐盐性显著高于WT植株。在75和150mmol·L-1NaCl处理下,TR比WT植株的苗高相对生长率(RGRH)分别高46%和44%,直径相对生长率(RGRD)高36%的61%；TR比WT维持了更大的叶面积,积累了更多的干物质。
NaCl处理下,TR植株比WT植株具有更高的生长速率和积累更多的干物质主要是由于维持了更高的光合能力。在75和150mmol·L-1NaCl处理5d,TR比WT植株的Pn分别高22%和9%；而在75和150mmol·L-1NaCl处理30d,TR比WT植株的Pn分别高13%和48%。气孔限制是盐胁迫下欧美107杨光合速率下降的主要因素。盐处理导致2个株系的PS Ⅱ最大光化学效率(Fv/Fm)、PS Ⅱ实际光化学量子产量(ΦPSⅡ)、光化学猝灭系数(qP)和电子传递速率(ETR)均下降,而非光化学猝灭系数(NPQ)则有不同程度的提高。但是,在同等NaCl处理下,TR植株比WT植株维持了更高Fv/Fm、ΦPSⅡ、qP、ETR和较低的NPQ。课件,TR植株在盐胁迫下能够更好的保护光系统,使其具有较高的光系统活性和光能转化效率,能将所吸收的光能有效地转化为化学能,提高光合电子传递速率,从而提高生物量,这在一定程度上提高了其抵御盐胁迫的能力。
NaCl胁迫下,活性氧清除剂SOD、POD、CAT活性在TR植株叶片中受盐诱导上调的幅度较大。NaCl处理30d,WT植株中的这些参数则显著低于对照。说明盐胁迫下,TR植株比WT植株能更有效地清除H202。
NaCl胁迫下,2个株系植株体内离子含量存在较大差异。总的说来,在75和150mmol·L-1NaCl处理下,TR植株的根、茎和叶均比WT植株积累了更多的Na+。在同等NaCl处理下,TR植株对K+的吸收明显大于WT植株。例如,在150mmol·L-1NaCl处理下,TR植株根、茎和叶的K+含量比WT植株分别高32%、38%和35%。尽管TR植株较WT植株吸收了更多的Na+,但维持了更高的K+和K+/Na+比率,说明TR植株具有更高的耐盐性,可能是由于对K+具有更好的吸收。
3、用不同浓度的海水(0%、10%、20%和30%)浇灌WT和TR幼苗30d,比较了二者的生长指标、光合参数和离子含量。2个株系植株的生长受到不同程度抑制,但二者的植株生长和生理指标存在显著的差异。在同等海水处理下,TR植株比WT植株具有更高的生长速率、总叶面积、光合速率和Fv/Fm,并积累了更多的干物质。TR植株的组织中比WT植株的积累了更多的Na+和K+,TR植株耐海水能力显著高于WT植株。在10%的海水处理下,TR植株的干重与对照相比并无显著差异,在20%和30%的海水处理下,其干重分别下降了8%和22%。可见,10%～20%海水用于浇灌TR欧美107杨是可行的。尽管如此,海水胁迫对TR植株个体发育的影响尚待进一步的田间耐盐试验研究。
4、以不同浓度NaCl(0、75和150mmol·L-1)浇灌土培盆栽的欧美107杨WT和TR幼苗30d。低盐处理下,WT植株生长显著受到抑制,随盐分强度加大,抑制作用增大,高盐处理下其干重只有对照的50%；而TR植株在低盐处理下干重与对照差异不显著,高盐处理时其干重为对照的74%。同等盐度处理下,TR的干重显著高于WT,且随着盐度升高,2个株系间植株干重差异增大。盐处理后,TR植株叶片叶绿素和类胡萝卜素的含量均显著高于WT,并能维持较高的Pn和Fv/Fm；虽然TR叶片和根系均较WT积累了更多的Na+,但同时也维持了更高的K+和K+/Na+比率,而且叶片对K+选择性的运输明显高于WT；同时,盐胁迫下TR叶片MDA含量和相对电导率显著低于WT。盐对TR叶片叶绿体超微结构的影响较轻,在高盐下仍保持了较好的内部结构。
5、AtNHX1的导入能够显著提高转基因欧美107杨的耐盐性。耐盐转基因杨树的获得也为提高树木耐盐性提供了一种可行的研究方法。本研究结果也验证了转单一的Na+/H+逆向转运蛋白基因是能够明显提高植物耐盐性的。
Salt stress is one of the major stresses that depress plant growth and limit crop production, and has been a great threat to agricultural development all over the word. To cope with salt stress, some plants have the ability of either extruding Na+out of cells or sequestering Na+into the vacuole in order to decrease excessive Na+in the cytoplasm. The compartmentation of Na+into vacuoles is implemented by Na+/H+antiporter located in tonoplast. The importance of vacuolar sequestration to plant salinity resistance has been underlined by experiments in which constitutive overexpression of different vacuolar Na+/H+antiporter genes can greatly increase salinity resistance in a wide range of plant species.
Poplar is one of the most important timber tree species planted in China. Therefore, understanding of the mechanism of its response to salt stress is of paramount importance to agriculture application and plant improvement. Overexpression of Na+/H+antiporters have been reported to increase the salt resistance of various herbaceous or shrub species, but very little is known about the role of these antiporters in the salt resistance of trees. Successful Agrobacterium-mediated transformation of AtNHXl to the shoot stem explants of Populus×euramericana 'Neva' and regeneration of transgenic plants has been previously performed. In order to elucidate mechanism of overexpressing the AtNHXl gene response to salt-tolerance in woody plants, the wild-type plants (WT)(Populus×euramericana'Neva') and its transgenic varieties (TR) that overexpress the AtNHX1gene were investigated to identify their growth and physiological character responses to salt stress. Moreover, the WT and TR were also exposed to different concentrations of seawater to clarify the correlation between overexpressing the AtNHXl gene and seawater-tolerance in poplars. All these would help to develop strategies for improving the understanding of transgenic plants response to salt stress. The main results obtained were shown as follows:
1. All tests using PCR and RT-PCR indicated that the AtNHX1gene was integrated into the genome of Populus×euramericana 'Neva' and transcribed in transgenic poplar plants.
2. Seedlings of WT and TR were subjected to different concentrations of NaCl (from0to150mmol·L-1) stress for30days. Plant growth was inhibited by salinity in both WT and TR but in different degrees, and salt-resistance of TR was much higher than that of WT. The relative growth rate of stem diameter (RGRH) values of TR were46%and44%and the relative growth rate of shoot height (RGRD) values were36%and61%more than that of WT at75and150mM NaCl treatments, respectively. Moreover, TR produced much more total leaf area and dry weight than WT.
Under the same NaCl level treatments, TR maintained higher RGR, total leaf area, and dry weight than WT because TR had a higher Pn. The Pn values in TR were22%and9%higher than that of WT when grown at75and150mmol·L-1NaCl for5days, and13%and48%higher for30days. Stomatal limitation was the primary factor limiting Pn in Populus×euramericana 'Neva' under salt stress. At the same time, maximum photochemical efficiency of PS Ⅱ (Fv/Fm), actual quantum yield of PS Ⅱ (Φps Ⅱ), photochemical quenching coefficient (qP) and relative electron transport rate (ETR) in both WT and TR decreased while non-photochemical quenching coefficient (NPQ) increased in various degrees. However, under the same level of NaCl treatment, TR maintained higher Fv/Fm, ΦpsⅡ qP, ETR and lower NPQ than those of WT. These suggest that TR could better protect PSⅡ from injury induced by the salt stress. So it had higher PSⅡ activity and light energy transform efficiency, which could turn the absorbed light energy into chemical energy effectively, and then increased the transmitting rate of photosythetic electrons, thereby enhancing biomass. To some extent, the propertites of TR decribed above improved salt resistance.
The activity of antioxidant enzymes such as SOD, POD and CAT were induced to increase greatly in the leaf of TR. To WT. salt stress for30days induced obvious decline on the activity of these enzymes. These indicate that TR was likely to be more effective in removing H2O2than WT under salt stress.
Differences in the pattern of K+accumulation between WT and TR were apparent under NaCl stress. In general, TR accumulated more Na+in the root, stem, and leaf than WT at both75and150mmol·L-1NaCl concentrations. The K+uptake in TR was much more than that in WT under the same NaCl level. For example, the K+contents of TR in the root, stem, and leaf were32%,38%, and35%more than those of WT under the150mmol·L-1NaCl treatment. Despite a higher accumulation of Na+in TR, it maintained relatively higher K+and K+/Na+ratio compared to WT. Results show that plants that TR was markedly more resistant to NaCl, possibly because of intracellular K+content regulation.
3. Seedlings of WT and TR were exposed to different seawater concentrations (from0to30%) for30days to determine the effects of seawater on seedling growth, photosynthetic productivity, and ion content. Plant growth was inhibited by seawater in both WT and TR but in different degrees, and seawater-resistance of TR was much higher than that of WT. TR could maintain higher RGR, total leaf area, Pn, and Fv/Fm, so that it is able to accumulate more dry mass than WT at the same seawater level. TR accumulated more Na" and K+contents in plant tissue than WT. At10%seawater treatment, the dry weight of TR was unaffected, but decreased by8%and22%at20%and30%seawater treatments, respectively. These findings indicate that10%-20%seawater could be used to irrigate TR plants, thus implying the viable utilization of seawater. However, further investigations of the effects of stresses relative to transgenic poplar ontogeny are necessary to assess realistic stress pressures that occur naturally in the field.
4. Seedlings of WT and TR which cultivated in pot with soil were subjected to different concentrations of NaCl (from0to150mmol·L-1) stress for30days. We investigated the salt-resistance of the WT and TR plants by measuring growth parameters, chlorophyll (Chl) and carotenoid (Car) content, Pn, ion content, leaf malondialdehyde (MDA) and electrolytic leakage. Compared with the control, the growth of WT was restrained significantly in the presence of both low salt and high salt. The dry weight of WT was significantly lower under salt stress than that of control, the dry weight decreased gradually with increasing NaCl concentration and their dry weight under high salt was just50%of the control. However, the dry weight of TR in low salt was similar to the control, up to the high salt treatment, where their dry weight was74%of the control. Moreover, the dry weight of TR was significant higher than that of WT in both NaCl treatments, and the discrepancy of dry weight was increased with increasing NaCl concentration. In the presence of NaCl, Chl and Car content of TR were significantly higher than that of WT, and the TR maintained a remarkably high Pn and Fv/Fm. Although TR accumulated more Na+in their roots and leaf tissues under salinity conditions compared with the WT, they absorbed more K+and maintained a higher K+/Na+ratio. Moreover, these TR plants kept a lower MDA and electrolytic leakage level than that of wild-type. Under salt stress, the chloroplasts of TR under low salt stress were not different from the control, and the structure of chloroplasts under high salt was better than that of WT. Under the same salinity level, the TR can maintain a relative integrated chloroplast, high chlorophyll contents and preferable net photo synthetic rate, so its salt tolerance were improved.
5. These findings indicate that transformation of AtNHXl gene into Populus×euramericana 'Neva' can confer plants more tolerance to salinity than its wild-type plants. These transgenic poplars provided a feasible way for improving important trees to adapt to salt stress conditions that are common in saline and arid soils. Our result also proved that transformation of single AtNHXl gene into plants can confer plants more resistance to salinity.
引文
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