松嫩草地羊草种子发育进程、休眠特性及与盐碱耐性关系的研究
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摘要
羊草（Leymus chinensis），是禾本科赖草属根茎型优质禾草，不但营养价值高、适口性好，而且具有较强的耐旱、耐寒和耐盐碱性。羊草既是东北松嫩平原的优势种，也是干旱与半干旱地区建植人工草地的优良草种。近年来，随着畜牧业的发展及生态环境治理力度的加大，人工羊草草地不断的建植，为此，人们对羊草种子的需求量与品质的要求也越来越高。本文通过研究羊草种子的发育进程、休眠特性及与盐碱耐性的关系，一方面明确羊草种子的最适宜收获时间，为农业生产上收获高品质羊草种子提供科学依据，另一方面挖掘羊草种子的发芽潜能、深入解析抗逆机理，为提高其利用率及抗逆新品种的选育提供理论基础。主要研究结果与结论如下：
（1）通过对羊草种子发育动态的研究结果表明，羊草种子在发育过程中，随着成熟度的不断提高，种子的颜色由绿色变为浅绿色，再变成黄色，最后变为棕黑色。种子的千粒重不断的增加，在盛花后33d达到最大值，之后趋于恒定。含水量与种子浸出液电导率则呈下降趋势，含水量在盛花后36d达到最小值，而两个试验年份电导率值有所差异，分别在盛花后27d和30d达到最小值。标准发芽试验结果显示，羊草种子在盛花后39d发芽率最高，此时种子的开始发芽时间、50%种子发芽时间、发芽势等指标均为最优。尽管加速老化试验的发芽指标与标准发芽试验略有差异，但是盛花后39d的种子同样具有最强的抗老化能力。上述试验结果表明，盛花后39d羊草种子活力最高，品质最佳，是种子最适宜的收获时间。
（2）不同成熟度的羊草种子对土壤埋深与盐碱胁迫同样具有不同的响应方式。羊草种子出苗与其后的幼苗生长能力随着种埋深度的增加而降低，1cm是最适宜的播种深度，此时的出苗率最高、出苗时间最短，并且叶片与根系长度与生物量最大；另外不同成熟度的羊草种子同样表现出不同的出土成苗能力，盛花后39d的羊草种子活力最大，其上述各项幼苗生长指标均为最优。种子的成熟度与盐、碱胁迫及其交互作用显著影响羊草种子的发芽率与发芽势，盛花后39d的羊草种子在胁迫下具有最高的发芽率与发芽势，特别是在高浓度（400mM）盐胁迫下，尤为明显。复萌试验结果显示，盛花后39d的羊草种子在盐碱胁迫（特别是高盐环境）解除后同样具有最高的发芽率。上述结果表明，尽管不同成熟度的羊草种子均具有发芽能力，但是盛花后39d的羊草种子出苗及抵御盐碱胁迫伤害的能力最强，这也进一步支撑了39d是羊草种子适宜收获时间这一结论。另外羊草适宜浅播，1cm是其最适宜的播种深度。
（3）通过人工手段处理可以明显打破羊草种子的休眠特性。研究结果表明，除了热水浸种处理外，其余方法如浓硫酸、冷层积、PEG、GA_3、KNO_3及清水浸种均能一定程度上提高羊草种子的发芽率，发芽速率、开始发芽时间及50%种子发芽时间。但是在生产实际中，既要考虑高效性也要考虑经济耗费，结合本试验的研究结果，我们推荐在生产中采用低温浸种20d的方法来打破羊草种子的休眠，提高其发芽率。
（4）稃是抑制羊草种子萌发的重要因素，但同时也一定程度提高了种子的抗盐性。通过测定稃对羊草种子吸水、脱水、不同温度条件下的发芽响应以及不同持续时间盐胁迫对种子发芽的影响，结果发现稃可以显著提高羊草种子的吸水量，并同时减缓种子在干旱环境下的脱水速率，使种子不会过度脱水而死亡。稃、不同温度处理及两者交互作用显著降低羊草种子的发芽率与发芽速率，表明稃对羊草种子萌发具有一定的抑制作用。在不同持续时间的盐胁迫处理下，未萌发的带稃种子复萌率均高于去稃种子，特别是在长时间及高盐胁迫下尤为明显，表明稃对羊草种子耐盐性起着重要的调节作用，一旦雨水、融雪等条件降低了土壤盐浓度，带稃种子就可以继续萌发出土。
（5）20-30°C是羊草种子最适宜的发芽温度，高温、低温均显著降低种子的发芽率与发芽速率，并且此温度可一定程度上减缓盐胁迫与碱胁迫对种子发芽的抑制效应。随着盐、碱胁迫浓度的增加，羊草种子发芽率与发芽速率均呈下降趋势，且在碱胁迫下的下降幅度更大。在盐胁迫下，当盐浓度＜200mM时，低温是影响种子萌发的主要因素，随着盐浓度的不断增加，高温则更加剧了盐胁迫对种子萌发的抑制作用；而在碱胁迫下，即使碱浓度较低，高温与其交互作用也大大加剧了对种子发芽的抑制。盐胁迫下未萌发的羊草种复萌率随盐浓度增加而增加，而在碱胁迫下则随着碱浓度增加呈先上升后下降的趋势，高浓度碱胁迫使羊草种子失去活力而死亡，并且碱胁迫下种子的复萌率低于盐胁迫，25-35°C同样不利于种子的复萌。幼苗生长对温度与盐碱胁迫交互作用的响应方式与发芽阶段相似，20-30°C同样是最适宜温度；另外，盐碱胁迫均对羊草幼苗根生长的抑制作用更强。基于以上研究结果，我们建议在初夏（七月上旬），高降雨过后，温度与土壤条件适宜的情况下进行播种，以提高羊草种子的发芽率，更好的达到恢复退化草地的效果。
（6）在混合盐碱胁迫下，羊草种子的发芽率与发芽速率均随着盐浓度的增加不断下降，且碱性盐比例越大下降越明显。在250mM盐浓度下，A组处理发芽率为6.5%，而其余5组处理发芽率均为0。羊草幼苗生长阶段同样受盐浓度、pH及两者交互作用影响，并且根系对胁迫伤害更敏感，所受抑制作用更强。逐级回归分析结果表明，在种子萌发阶段，盐浓度是羊草种子在混合盐碱胁迫下能否萌发的决定性因素，而一旦胚根突破种皮进入幼苗生长阶段，pH就转变为主导因素。上述研究表明，混合盐碱胁迫对羊草种子萌发与早期幼苗生长阶段的抑制机理有所不同，其中高盐浓度与高pH的交互作用对种子萌发与幼苗生长的抑制效应最强。
（7）盐胁迫与碱胁迫均显著降低羊草幼苗的长度、鲜重与含水量，且碱胁迫抑制作用更强。两种胁迫均造成羊草幼苗Na~+浓度与Na~+/K~+升高，并且K~+浓度下降，但是在碱胁迫下，Na~+浓度、Na~+/K~+上升幅度与K~+下降幅度均大于盐胁迫。另外，在盐胁迫下，羊草幼苗大量积累Cl~-，有机酸含量变化不大；而在碱胁迫下，Cl~-、NO_3~-与H_2PO_4~-均呈下降趋势，而有机酸则大量积累，其中苹果酸、柠檬酸是主要的有机酸组分，可溶性糖是羊草幼苗在两种胁迫下共同的渗透调节物质。上述结果表明，碱胁迫由于具有高pH，对羊草早期幼苗生长的抑制作用更强，Cl~-与有机酸积累特征的差异表明羊草早期幼苗在盐胁迫与碱胁迫下具有不同的适应策略。
Leymus chinensis (Trin.) Tzvel. is a perennial rhizomatous high quality species of thefamily Poaceae. It not only has high forage value and good palatability, but also has greattolerant to drought conditions, cold extremes and salt-alkaline conditions. This plant is adominant grass species in Songnen Plain of Northern China, and also an ideal grass forrangeland use in arid and semiarid regions. In recent years, with the development of animalhusbandry and the intensified efforts on ecological environment, artificial grasslands of L.chinensis were constantly built. Therefore, the request on the quantity and quality of L.chinensis seeds was also more and more high. In this paper, we investigated seed development,dormancy characters and salt-alkaline tolerance of Leymus chinensis. On the one hand, weclearly determined the optimum seed harvest time of this species, and provided scientificbasis for agricultural production and high quality seed harvest. On the other hand, weexcavated seed germinating potential, deeply analysed the stresses tolerance mechanism, andprovided theoretical basis for the utilization rate and breeding the new varieties of Leymuschinensis. The main results and conclusions from our experiments were as follows:
(1) The results of seed development of Leymus chinensis showed that, seed colorchanged from green to light green, yellow and heavy brown into the final along with seedmaturity.1000seeds weight increased constantly, and reached hightest at33days after peakanthesis. However, water content and electric conductivity of the seeds showed a decliningtrend, water content reached lowest at36days after peak anthesis, and the values of electricconductivity of two experimental years were different, which reached minimum at27and30days after peak anthesis, respectively. The results of standard germination test showed thatgermination percentage was highest at39days after peak anthesis. At this time, germinationstarting days,50%germination days and germination energy were all reached the optimalvalue. Although slightly difference was found between standard germination test andaccelerated aging test, seeds at39days after peak anthesis also had the strongest anti-agingcapability. Above results showed that seeds at39days after peak anthesis has the hightestvigor and best quality, and is the optimum seed harvest time of Leymus chinensis.
(2) Response to burial depth and salt-alkaline stress of Leymus chinensis seeds atdifferent maturation time were also greatly differed. The ability of seedling emergence andgrowth were decreased with increasing burial depth, and1cm was the most suitable plantingdepth. At this time, the seedling emergence rate was highest, time to seedling emergence wasshortest, and the length and biomass of the shoot and root were also highest. In addition, theability of seedling growth of the seeds at different maturation time was also different. Seeds at 39days after peak anthesis had the highest vigor, the above index of seedling growth wereoptimum. Germination percentage and energy were significantly affected by seed maturationtime, salt-alkaline stress and their interactions. Highest germination percentage andgermination energy were occurred at at39days after peak anthesis, especially at highest salinrstress (400mM). The recovery test showed that recovery percentage was also highest at39days after peak anthesis. Above results show that although seed of different maturation timehas germination ability, the ability of seedling emergence and resistance to salt-alkaline stressis highest at39days after peak anthesis, which further support the view that39days afterpeak anthesis is the optimum seed harvest time of Leymus chinensis. In addition, shallowsowing is suitable for Leymus chinensis, and1cm is the most optimum planting depth.
(3) Artificial treatments can obviously break the seed dormancy of L. chinensis. Theresults showed that many methods such as H2SO4, cold stratification, PEG, GA_3, KNO_3andsoaking in the water all enhanced germination percentage, germination rate, startinggermination time, and50%germination time. However, in production practice, bothefficiency and ecomomic cost should be considered, and combined with our results, werecommend the way of soaking seeds in water under lower temperature for20d in order tobreak seed dormancy and increase germination percentage of Leymus chinensis.
(4) Lemma is an important fator inhibiting seed germination of Leymus chinensis, but itcan also improve salt resistance of the seeds. We investigated the effects of lemmas on seedimbibition and dehydration, germination responses to various temperature regimes and theimpact of different duration salt stress on seed germination. The results showed that lemmascould significantly enhance water absorption, and slowed down the dehydration rate underdrought conditions. Lemmas, temperature regimes and their interactions significantlydecreased germination percentage and germination rate, indicating that lemmas could inhibitgermination process of Leymus chinensis. Under different duration of salt stress, recoverypercentages of non-germinated seeds with lemmas were higher than that without lemmas,especially under long duration of salt stress, indicating that lemmas could enhance the saltresistance of seed. Once the precipitation and melting snow decreased salinity concentrationin the soil, seeds with lemmas can geminate again.
(5) Germination percentages and rates were inhibited by either an increase or decreasetemperature from the optimal temperature20-30°C, and this temperature can alleviate theinhibitory effects of salt-alkaline stress on seed germination. With the increasing salinity andalkalinity, seed germination and germination rate were both decreased, and the reductionswere much greater under alkaline stress. Under salt stress, when salinity＜200mM, lowertemperature was the main factor inhibiting seed germination. With the increasing salinity,higher temperature aggravated the adverse effects. While under alkali stress, germinationpercentage and germination rate were both decreased markedly at25-35°C even though the alkalinity was very low. Recovery percentage of non-germinated seeds increased with theincreasing salinity, but increased at first and then declined under alkaline stress, and recoverypercentages were lowest in both stresses at25-35°C, especially under alkaline stress. Seedlinggrowth had similar response to the interactions of temperature and salt-alkaline stress,20-30°C was also the optimum temperature. In addition, both of the two stresses inhibitedroot growth much stronger. Due to the above results, early July sowings in field would berecommended, when temperature is appropriate and salinity-alkalinity concentrations arealways reduced by the high rainfall.
(6) Under mixed salt-alkaline stresses, germination percentage and germination ratedecreased with increasing salinity under all the stress treatments, and the reductions weregreater under treatment which the proportion of alkaline salts was greater. At250mM salinity,germination percentage of treatment A was6.5%, but the other five treatments were0.Seedling growth was also affacted by salinity, pH and their interactions. However, radiclelength decreased more markedly with increasing salinity and pH. Stepwise regression analysisresults showed that salinity was the dominant factor for seed germination under mixedsalt-alkaline stress conditions. However, once radicle break through the seed coat, and pHchanged into the dominant factor for seedling establishment. Above results indicate thatmixed salt-alkaline stresses has different impacts on germination and early seedling stages ofL.chinensis. The interactions of high salinity and high pH have the strongest inhibition onseed germination and seedling growth.
(7) Both saline stress and alkaline stress significantly decreasd seedling length, freshweight and water content, and the reductions were much greater under alkaline stress. TheNa~+concentration, Na~+/K~+ratio increased butK~+concentration decreased under both stresses,and the changes were greater under alkali stress. Under salt stress, shoots mainly accumulatedCl~-and little change was found in organic acids. While under alkali stress, the concentrationsof Cl~-, NO_3~-and H_2PO_4~-were all decreased, and organic acids were accumulated, especiallymalic acid and citric acid. In addition, soluble sugar was the same osmoregulation under thetwo stresses. Above results indicate that alkali stress inhibited early seedlings of Leymuschinensis much greater because of the high pH, different accumulation characters of Cl~-andorganic acids indicated that different adaptive mechanism to saline and alkaline stresses wereexist of Leymus chinensis during early seedling stage.
（1）通过对羊草种子发育动态的研究结果表明，羊草种子在发育过程中，随着成熟度的不断提高，种子的颜色由绿色变为浅绿色，再变成黄色，最后变为棕黑色。种子的千粒重不断的增加，在盛花后33d达到最大值，之后趋于恒定。含水量与种子浸出液电导率则呈下降趋势，含水量在盛花后36d达到最小值，而两个试验年份电导率值有所差异，分别在盛花后27d和30d达到最小值。标准发芽试验结果显示，羊草种子在盛花后39d发芽率最高，此时种子的开始发芽时间、50%种子发芽时间、发芽势等指标均为最优。尽管加速老化试验的发芽指标与标准发芽试验略有差异，但是盛花后39d的种子同样具有最强的抗老化能力。上述试验结果表明，盛花后39d羊草种子活力最高，品质最佳，是种子最适宜的收获时间。
（2）不同成熟度的羊草种子对土壤埋深与盐碱胁迫同样具有不同的响应方式。羊草种子出苗与其后的幼苗生长能力随着种埋深度的增加而降低，1cm是最适宜的播种深度，此时的出苗率最高、出苗时间最短，并且叶片与根系长度与生物量最大；另外不同成熟度的羊草种子同样表现出不同的出土成苗能力，盛花后39d的羊草种子活力最大，其上述各项幼苗生长指标均为最优。种子的成熟度与盐、碱胁迫及其交互作用显著影响羊草种子的发芽率与发芽势，盛花后39d的羊草种子在胁迫下具有最高的发芽率与发芽势，特别是在高浓度（400mM）盐胁迫下，尤为明显。复萌试验结果显示，盛花后39d的羊草种子在盐碱胁迫（特别是高盐环境）解除后同样具有最高的发芽率。上述结果表明，尽管不同成熟度的羊草种子均具有发芽能力，但是盛花后39d的羊草种子出苗及抵御盐碱胁迫伤害的能力最强，这也进一步支撑了39d是羊草种子适宜收获时间这一结论。另外羊草适宜浅播，1cm是其最适宜的播种深度。
（3）通过人工手段处理可以明显打破羊草种子的休眠特性。研究结果表明，除了热水浸种处理外，其余方法如浓硫酸、冷层积、PEG、GA_3、KNO_3及清水浸种均能一定程度上提高羊草种子的发芽率，发芽速率、开始发芽时间及50%种子发芽时间。但是在生产实际中，既要考虑高效性也要考虑经济耗费，结合本试验的研究结果，我们推荐在生产中采用低温浸种20d的方法来打破羊草种子的休眠，提高其发芽率。
（4）稃是抑制羊草种子萌发的重要因素，但同时也一定程度提高了种子的抗盐性。通过测定稃对羊草种子吸水、脱水、不同温度条件下的发芽响应以及不同持续时间盐胁迫对种子发芽的影响，结果发现稃可以显著提高羊草种子的吸水量，并同时减缓种子在干旱环境下的脱水速率，使种子不会过度脱水而死亡。稃、不同温度处理及两者交互作用显著降低羊草种子的发芽率与发芽速率，表明稃对羊草种子萌发具有一定的抑制作用。在不同持续时间的盐胁迫处理下，未萌发的带稃种子复萌率均高于去稃种子，特别是在长时间及高盐胁迫下尤为明显，表明稃对羊草种子耐盐性起着重要的调节作用，一旦雨水、融雪等条件降低了土壤盐浓度，带稃种子就可以继续萌发出土。
（5）20-30°C是羊草种子最适宜的发芽温度，高温、低温均显著降低种子的发芽率与发芽速率，并且此温度可一定程度上减缓盐胁迫与碱胁迫对种子发芽的抑制效应。随着盐、碱胁迫浓度的增加，羊草种子发芽率与发芽速率均呈下降趋势，且在碱胁迫下的下降幅度更大。在盐胁迫下，当盐浓度＜200mM时，低温是影响种子萌发的主要因素，随着盐浓度的不断增加，高温则更加剧了盐胁迫对种子萌发的抑制作用；而在碱胁迫下，即使碱浓度较低，高温与其交互作用也大大加剧了对种子发芽的抑制。盐胁迫下未萌发的羊草种复萌率随盐浓度增加而增加，而在碱胁迫下则随着碱浓度增加呈先上升后下降的趋势，高浓度碱胁迫使羊草种子失去活力而死亡，并且碱胁迫下种子的复萌率低于盐胁迫，25-35°C同样不利于种子的复萌。幼苗生长对温度与盐碱胁迫交互作用的响应方式与发芽阶段相似，20-30°C同样是最适宜温度；另外，盐碱胁迫均对羊草幼苗根生长的抑制作用更强。基于以上研究结果，我们建议在初夏（七月上旬），高降雨过后，温度与土壤条件适宜的情况下进行播种，以提高羊草种子的发芽率，更好的达到恢复退化草地的效果。
（6）在混合盐碱胁迫下，羊草种子的发芽率与发芽速率均随着盐浓度的增加不断下降，且碱性盐比例越大下降越明显。在250mM盐浓度下，A组处理发芽率为6.5%，而其余5组处理发芽率均为0。羊草幼苗生长阶段同样受盐浓度、pH及两者交互作用影响，并且根系对胁迫伤害更敏感，所受抑制作用更强。逐级回归分析结果表明，在种子萌发阶段，盐浓度是羊草种子在混合盐碱胁迫下能否萌发的决定性因素，而一旦胚根突破种皮进入幼苗生长阶段，pH就转变为主导因素。上述研究表明，混合盐碱胁迫对羊草种子萌发与早期幼苗生长阶段的抑制机理有所不同，其中高盐浓度与高pH的交互作用对种子萌发与幼苗生长的抑制效应最强。
（7）盐胁迫与碱胁迫均显著降低羊草幼苗的长度、鲜重与含水量，且碱胁迫抑制作用更强。两种胁迫均造成羊草幼苗Na~+浓度与Na~+/K~+升高，并且K~+浓度下降，但是在碱胁迫下，Na~+浓度、Na~+/K~+上升幅度与K~+下降幅度均大于盐胁迫。另外，在盐胁迫下，羊草幼苗大量积累Cl~-，有机酸含量变化不大；而在碱胁迫下，Cl~-、NO_3~-与H_2PO_4~-均呈下降趋势，而有机酸则大量积累，其中苹果酸、柠檬酸是主要的有机酸组分，可溶性糖是羊草幼苗在两种胁迫下共同的渗透调节物质。上述结果表明，碱胁迫由于具有高pH，对羊草早期幼苗生长的抑制作用更强，Cl~-与有机酸积累特征的差异表明羊草早期幼苗在盐胁迫与碱胁迫下具有不同的适应策略。
Leymus chinensis (Trin.) Tzvel. is a perennial rhizomatous high quality species of thefamily Poaceae. It not only has high forage value and good palatability, but also has greattolerant to drought conditions, cold extremes and salt-alkaline conditions. This plant is adominant grass species in Songnen Plain of Northern China, and also an ideal grass forrangeland use in arid and semiarid regions. In recent years, with the development of animalhusbandry and the intensified efforts on ecological environment, artificial grasslands of L.chinensis were constantly built. Therefore, the request on the quantity and quality of L.chinensis seeds was also more and more high. In this paper, we investigated seed development,dormancy characters and salt-alkaline tolerance of Leymus chinensis. On the one hand, weclearly determined the optimum seed harvest time of this species, and provided scientificbasis for agricultural production and high quality seed harvest. On the other hand, weexcavated seed germinating potential, deeply analysed the stresses tolerance mechanism, andprovided theoretical basis for the utilization rate and breeding the new varieties of Leymuschinensis. The main results and conclusions from our experiments were as follows:
(1) The results of seed development of Leymus chinensis showed that, seed colorchanged from green to light green, yellow and heavy brown into the final along with seedmaturity.1000seeds weight increased constantly, and reached hightest at33days after peakanthesis. However, water content and electric conductivity of the seeds showed a decliningtrend, water content reached lowest at36days after peak anthesis, and the values of electricconductivity of two experimental years were different, which reached minimum at27and30days after peak anthesis, respectively. The results of standard germination test showed thatgermination percentage was highest at39days after peak anthesis. At this time, germinationstarting days,50%germination days and germination energy were all reached the optimalvalue. Although slightly difference was found between standard germination test andaccelerated aging test, seeds at39days after peak anthesis also had the strongest anti-agingcapability. Above results showed that seeds at39days after peak anthesis has the hightestvigor and best quality, and is the optimum seed harvest time of Leymus chinensis.
(2) Response to burial depth and salt-alkaline stress of Leymus chinensis seeds atdifferent maturation time were also greatly differed. The ability of seedling emergence andgrowth were decreased with increasing burial depth, and1cm was the most suitable plantingdepth. At this time, the seedling emergence rate was highest, time to seedling emergence wasshortest, and the length and biomass of the shoot and root were also highest. In addition, theability of seedling growth of the seeds at different maturation time was also different. Seeds at 39days after peak anthesis had the highest vigor, the above index of seedling growth wereoptimum. Germination percentage and energy were significantly affected by seed maturationtime, salt-alkaline stress and their interactions. Highest germination percentage andgermination energy were occurred at at39days after peak anthesis, especially at highest salinrstress (400mM). The recovery test showed that recovery percentage was also highest at39days after peak anthesis. Above results show that although seed of different maturation timehas germination ability, the ability of seedling emergence and resistance to salt-alkaline stressis highest at39days after peak anthesis, which further support the view that39days afterpeak anthesis is the optimum seed harvest time of Leymus chinensis. In addition, shallowsowing is suitable for Leymus chinensis, and1cm is the most optimum planting depth.
(3) Artificial treatments can obviously break the seed dormancy of L. chinensis. Theresults showed that many methods such as H2SO4, cold stratification, PEG, GA_3, KNO_3andsoaking in the water all enhanced germination percentage, germination rate, startinggermination time, and50%germination time. However, in production practice, bothefficiency and ecomomic cost should be considered, and combined with our results, werecommend the way of soaking seeds in water under lower temperature for20d in order tobreak seed dormancy and increase germination percentage of Leymus chinensis.
(4) Lemma is an important fator inhibiting seed germination of Leymus chinensis, but itcan also improve salt resistance of the seeds. We investigated the effects of lemmas on seedimbibition and dehydration, germination responses to various temperature regimes and theimpact of different duration salt stress on seed germination. The results showed that lemmascould significantly enhance water absorption, and slowed down the dehydration rate underdrought conditions. Lemmas, temperature regimes and their interactions significantlydecreased germination percentage and germination rate, indicating that lemmas could inhibitgermination process of Leymus chinensis. Under different duration of salt stress, recoverypercentages of non-germinated seeds with lemmas were higher than that without lemmas,especially under long duration of salt stress, indicating that lemmas could enhance the saltresistance of seed. Once the precipitation and melting snow decreased salinity concentrationin the soil, seeds with lemmas can geminate again.
(5) Germination percentages and rates were inhibited by either an increase or decreasetemperature from the optimal temperature20-30°C, and this temperature can alleviate theinhibitory effects of salt-alkaline stress on seed germination. With the increasing salinity andalkalinity, seed germination and germination rate were both decreased, and the reductionswere much greater under alkaline stress. Under salt stress, when salinity＜200mM, lowertemperature was the main factor inhibiting seed germination. With the increasing salinity,higher temperature aggravated the adverse effects. While under alkali stress, germinationpercentage and germination rate were both decreased markedly at25-35°C even though the alkalinity was very low. Recovery percentage of non-germinated seeds increased with theincreasing salinity, but increased at first and then declined under alkaline stress, and recoverypercentages were lowest in both stresses at25-35°C, especially under alkaline stress. Seedlinggrowth had similar response to the interactions of temperature and salt-alkaline stress,20-30°C was also the optimum temperature. In addition, both of the two stresses inhibitedroot growth much stronger. Due to the above results, early July sowings in field would berecommended, when temperature is appropriate and salinity-alkalinity concentrations arealways reduced by the high rainfall.
(6) Under mixed salt-alkaline stresses, germination percentage and germination ratedecreased with increasing salinity under all the stress treatments, and the reductions weregreater under treatment which the proportion of alkaline salts was greater. At250mM salinity,germination percentage of treatment A was6.5%, but the other five treatments were0.Seedling growth was also affacted by salinity, pH and their interactions. However, radiclelength decreased more markedly with increasing salinity and pH. Stepwise regression analysisresults showed that salinity was the dominant factor for seed germination under mixedsalt-alkaline stress conditions. However, once radicle break through the seed coat, and pHchanged into the dominant factor for seedling establishment. Above results indicate thatmixed salt-alkaline stresses has different impacts on germination and early seedling stages ofL.chinensis. The interactions of high salinity and high pH have the strongest inhibition onseed germination and seedling growth.
(7) Both saline stress and alkaline stress significantly decreasd seedling length, freshweight and water content, and the reductions were much greater under alkaline stress. TheNa~+concentration, Na~+/K~+ratio increased butK~+concentration decreased under both stresses,and the changes were greater under alkali stress. Under salt stress, shoots mainly accumulatedCl~-and little change was found in organic acids. While under alkali stress, the concentrationsof Cl~-, NO_3~-and H_2PO_4~-were all decreased, and organic acids were accumulated, especiallymalic acid and citric acid. In addition, soluble sugar was the same osmoregulation under thetwo stresses. Above results indicate that alkali stress inhibited early seedlings of Leymuschinensis much greater because of the high pH, different accumulation characters of Cl~-andorganic acids indicated that different adaptive mechanism to saline and alkaline stresses wereexist of Leymus chinensis during early seedling stage.
引文
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