Effects of Sand Burial on the Survival, Growth, and Biomass Allocation in Semi-shrub Hedysarum laeve Seedlings

Hedysarum laeve Maxim. (Leguminosae) is one of the major species used frequently in revegetation of dune_field in the sandlands of the northern part of China by means of aerial sowing. Seedlings of the species after emergence above the sand surface may be buried in sand to various depths during its establishment in late spring and early summer. A study was made to examine the effects of sand burial at different levels of 0 (control), 33%, 67%, 100% and 133% of their shoot height, on the survivorship, growth, and biomass allocation pattern of H. laeve seedlings (one and two weeks old after emergence). When burial depth was up to 100% of their shoot height, about 70% seedlings died; and the burial at depth of 133% of their shoot height led to death of all seedlings. When seedlings was buried at depth of 33% and 67% of their shoot height, respectively, after six_week growth, their biomass of whole plant, blade, and root and relative growth rate were higher than the unburied counterparts. The seedlings in both 33% and 67% sand burial treatments did not significantly change their biomass allocation pattern comparing with the unburied ones. Furthermore, the number of leaves and shoot height of the seedlings in both 33% and 67% sand burial treatments were not significantly different from those of unburied individuals, respectively. The newly born leaves of the surviving seedlings, in 33%, 67%, and 100% burial treatments, during the period of experiment, were significantly more than those in control.

Growth and physiological responses of Agriophyllum squarrosum to sand burial stress

Agriophyllum squarrosum is an annual desert plant widely distributed on mobile and semi-mobile dunes in all the sandy deserts of China. We studied the growth and physiological properties of A. squarrosum seedlings under different sand burial depths in 2010 and 2011 at Horqin Sandy Land, Inner Mongolia to understand the ability and mechanism that A. squarrosum withstands sand burial. The results showed that A. squarrosum had a strong ability to withstand sand burial. Its survival rate, plant height and biomass increased significantly at a burial depth 25% of seedling height and decreased significantly only when the burial depth exceeded the height of the seedlings; some plants still survived even if the burial depth reached 266% of a seedling height. The malondialdehyde （MDA） content and membrane permeability of the plant did not change significantly as long as the burial depth was not greater than the seedling height; lipid peroxidation increased and cell membranes were damaged if the burial depth was increased further. When subjected to sand burial stress, superoxide dismutase （SOD） and peroxidase （POD） activities and free proline content increased in the seedlings, while the catalase （CAT） activity and soluble sugar content decreased. Sand burial did not lead to water stress. Reductions in photosynthetic area and cell membrane damage caused by sand burial may be the major mechanisms increasing mortality and inhibiting growth of the seedling. But the increases in SOD and POD activities and proline content must play a certain role in reducing sand burial damage.

Impact of sand burial on maize(Zea mays L.)productivity and soil quality in Horqin sandy cropland,Inner Mongolia,China

Croplands are often suffering from sand burial in dry regions of northern China. For studying this phenomenon, we carried out a case study of field experiment including four sand burial levels, i.e. shallow （1-3 cm）, moderate （8-12 cm） and deep （15-20 cm） sand burials, and no sand burial （control, CK）, in a typical agro-pastoral transitional zone in Naiman Banner of eastern Inner Mongolia. The aim of this study was to assess the impacts of sand burial on maize （Zea rnays L.） productivity and the soil quality along a gradient of burial depths. Results showed that there was a strong negative effect of sand burial on maize productivity and soil quality, which significantly declined （P〈0.05） under moderate and deep sand burial treatments. In comparison with the CK, the maize yield and above-ground biomass reduced by 47.41% and 39.47%, respectively. The soil silt and clay, soil water, soil organic carbon and total nitrogen contents under deep sand burial decreased by 67.85%, 40.32%, 86.52% and 82.11%, respectively, while microbial biomass carbon, microbial abundance and enzyme activity decreased by 89.78%, 42.28%-79.66% and 69.51%-97.71%, respectively. There was no significant effect on crop productivity and soil quality with shallow sand burial treatment. The correlations analysis showed that there was significant positive correlations of both maize yield and above-ground biomass with soil silt and clay, soil organic carbon and total nitrogen contents, pH, electrical conductivity, soil water content, microbial abundance and biomass and all tested soil enzyme activities. Stepwise regression analysis indicated that soil water and total nitrogen contents, urease, cellobiohydrolase and peroxidase activities were key determining factors for maize productivity. This combination of factors explains reason of the decreased maize productivity with deep sand burial. We found that degradation of cropland as a result of sand burial changed soil physical-chemical properties and soil enzyme activities in the plow layer, and decreased overall maize productivity. Furthermore, decreased soil enzyme activity was a better indicator to predict sandy cropland degradation.

Effects of sand burial on dune plants: a review

Burial of different growth stages of plants （e.g., adult plants, seedlings and seeds） is frequent in dune ecosystems. The soil micro-environment, which differs from surface conditions, influences the survival and growth of dune plants. To sum up knowledge about the survival mechanisms of plants under sand burial and to promote practical rehabilitation of dune vegetation, we reviewed relevant published literature and concluded that： （1） Focus in recent years has been on impacts of sand burial on seed germination and seedling emergence. Generally, shallow burial increased seed germination and seedling emergence, but deeper burial was negative. Buried at the same depth, large seeds showed higher germination and seedling emergence rates, attributed to larger energy reserves. （2） Survival, growth and reproduction rates of dune plants show plasticity in response to sand burial. Long-term deep burial is fatal because it creates a physical barrier which overcomes the vertical growth of plants, reduces photosynthetic leaf area, and limits oxygen availability to roots. Modest burial, on the other hand, is advantageous for growth and reproduction of many dune plants, due to protection from excessive temperature and drought. （3） There are few reports concerning effects of sand burial on plant physiology, but a limited number of studies indicate that partial burial increases water use efficiency, chlorophyll content, transpiration rate and net photosynthetic rates. The antioxidant protective enzyme system and osmolyte balance were reported to be involved in the mechanisms of dune plant resistance to burial.

Effects of sand burial on growth and antioxidant enzyme activities of wheat(Triticum aestivum L.) in northern China

Growth and physiological responses of wheat to sand burial were studied in Horqin Sandy Land, to determine the impact on productivity and survival as well as antioxidant enzymes responses. This study consisted of one control （no sand） and four sand burial treatments： 25%, 50%, 75% and 100% of seedling height, respectively. Minor burial （25%） had no effect on wheat growth and survival; deep burial （100%） was fatal, and the others had an intermediate effect. Thus, the survival limit to sand burial was equal to seedling height. Sand burial mainly decreased shoot biomass and crop yield, but had small effects on belowground biomass. Superoxide dismutase （SOD） activity increased with time after burial in all treatments with surviving plants. Peroxidase （POD） activity increased after six days under burial, and catalase （CAT） activity de- creased after burial, but recovered after 12 days. The concentration of malondialdehyde （MDA）, a marker for oxidative stress, was low on the sixth day, but increased thereafter with burial depth. Thus, sand burial 〉25% should be avoided due to growth rate reduction leading to reduced crop yield, and even 25% burial showed physiological indicators of stress.

Effects of sand burial on survival and growth of Artemisia halodendron and its physiological response

There is a great deal of literature on the effects of sand burial upon the survival and growth of desert plants, but the physiological adaption mechanisms of desert plants to sand burial have as yet rarely been studied. Artemisia halodendron is widely distributed in the semi-arid deserts of China and is a dominant species in semi-moving dune vegetation. The growth and physiological properties ofA. halodendron seedlings under different sand burial depths were studied in 2010 and 2011 in the Horqin Sand Land, Inner Mongolia, to better understand the ability and physiological mechanism by which desert plants withstand sand burial. The results showed that A. halodendron as a prammophyte species had a stronger ability to withstand sand burial compared to non-prammophytes, with some plants still surviving even if buried to a depth reaching 225% of seedling height. Although seedling growth was inhibited significantly once the depth of sand burial reached 50% of the seedling height, seedling survival did not decrease significantly until the burial depth exceeded 100% of the seedling height. Sand burial did not result in significant water stress or MDA （Malondialdehyde） accumulation in the seedlings, but membrane permeability increased significantly when the burial depth exceeded 100% of the seedling height. After being subjected to sand burial stress, POD （Peroxidase） activity and proline content increased significantly, but SOD （Superoxide Dismutase） and POD activities and soluble sugar content did not. The primary mechanism resulting in in- creased mortality and growth inhibition were that cell membranes were damaged and photosynthetic area decreased when subjected to the severe stress of sand burial, while proline and POD played key roles in osmotic adjustment and protecting cell membranes from damage, respectively.

Responses of Hedysarum Laeve,a guerrilla clonal semi-shrub in the Mu Us Sandland,to local sand burial

In arid and semi-arid inland deserts,one of the environmental stresses for plants is recurrent sand burial,which can influence the physical and biotic microenviron-ments of the plants and soil.Previous studies have shown that different levels of sand burial have different effects on plants.Slight sand burial could increase the height increment,leaf biomass and the number of new ramets of the plants while heavy sand burial could impair the growth of the plants and even decrease their chances of survival.In other words,below a certain threshold level of burial,the growth of plants is stimulated probably because of multiple factors.However,as the level of burial increases,the positive response starts to decline until it becomes a negative value.Arid and semi-arid inland deserts are frequently colonized and stabilized by many rhizomatous clonal plants.Clonal physiological integration often helps clonal plants buffer local environmen-tal stress encountered by ramets.A rhizomatous clonal semi-shrub,Hedysarum laeve(H.laeve),is the dominant plant species and important for vegetation restoration in the Mu Us sandland.To investigate whether clonal integration can increase the threshold of sand burial and help rhizomatous H.laeve tolerate heavy sand burial,we conducted a field experiment.The results showed that slight sand burial could accelerate ramet growth and enhance leaf biomass,stem bio-mass and shoot biomass,while heavy sand burial reducesed the biomass of the plant and impairs survival and growth of the ramets.Clonal integration increased the threshold of sand burial.Under heavy sand burial,ramets connected to other ramets not buried in sand were more in terms of height increment,stem biomass,leaf biomass and shoot biomass compared to the ramets encountering sand burial but disconnected from other ramets.It suggested that clonal physiological integration could help H.laeve ramets tolerate relatively heavy sand burial.We also discussed that clonal integration plays a role in H.laeve presence in the Mu Us sandland.

Effects of sand burial on biomass, chlorophyll fluores-cence and extracellular polysaccharides of man-made cyanobacterial crusts under experimental conditions

Soil cyanobacterial crusts occur throughout the world, especially in the semiarid and arid regions. It always encounters sand burial, which is an important feature of mobile sand dunes. A greenhouse study was conducted to determine the effects of sand burial on biomass, chlorophyll fluorescence and extracellular polysaccharides of man-made cyanobacterial crusts in six periods of time (0, 5, 10, 15, 20 and 30 d after burying) and at five depths (0, 0.2, 0.5, 1 and 2cm). The results indicated that with the increase of the burial time and burial depth extracellular polysaccharides content and Fv/Fm decreased correspondingly and there were no significant differences between 20 and 30 burial days under dif-ferent burial depths. The degradation of chlorophyll a content appeared only at 20 and 30 burial days and there was also no significant difference between them under different burial depths. It was also observed a simultaneous decrease of the values of the Fv/Fm and the content of extracellular poly-saccharides happened in the crusted cyanobacterium Microcoleus vaginatus Gom. It may suggest that there exists a relationship between extracellular polysaccharides and recovery of the activity of pho-tosystem II (PS II) after rehydration.

Adaptation of Nitraria sphaerocarpa to wind-blown sand environments at the edge of a desert oasis

This study addresses the adaptation of Nitraria sphaerocarpa to blown sand at the edge of a desert oasis with regard to the aspects of soil seed banks, seedlings, and population. Horizontally, the total number of seeds per unit area decreased from the shrub canopy center to intershrub areas, and most seeds were found under shrub canopies. Vertically, the highest proportion of seeds was found at depths of 5-10 cm. The emergence percentage, seedling mass, and seedling height, which were significantly affected by both burial depth and seed size, were highest at the optimal burial depth of 2 cm, and decreased with increasing burial depth in each seed size-class. Although seedling mass was usually greatest for large seeds and least for small seeds at each burial depth, little difference was observed in seedling height at shallow burial depths of 0-3 cm. The population shows a patchy and discontinuous distribution pattern. Population height increases with increasing sand depth. Also the density increases with increasing depth of sand in the desert; however, there is a steady decrease when the depth of sand is more than 100 cm. This result indicates that the depth of sand that is most suitable for the growth of Nitraria sphaerocarpa is 100 cm. The size of the population is significantly correlated with the sand depth, which increases with increasing depth in the desert.

裸露与沙埋对极端干旱区凋落物分解和养分释放的影响

为了评估沙漠公路防护林林带裸露与沙埋处理下的凋落物分解和养分动态特征,以梭梭(Haloxylon ammodendron)和乔木状沙拐枣(Calligonum arborescens)同化枝、多枝柽柳(Tamarix ramosissima)叶凋落物为研究对象,用凋落物分解袋法进行分解试验。结果表明:(1)3种凋落物的质量损失在2种处理下均存在显著差异,沙埋处理下凋落物质量损失大于裸露处理,至分解试验结束,裸露处理下梭梭、乔木状沙拐枣和多枝柽柳的失重率分别为7%、6.8%、18.1%;沙埋处理下梭梭、乔木状沙拐枣和多枝柽柳的失重率分别为23.7%、9.7%、21.9%;(2)分解过程中,2种处理下凋落物C、N、P含量变化不太一致,梭梭与乔木状沙拐枣同化枝的N与P含量均呈净富集状态,而C含量则呈现净释放状态;多枝柽柳叶的N和P含量分别呈净富集、净释放状态,C含量为富集-释放状态;(3)分解速率k值与相关因素分析表明,凋落物初始N、P、C:N与C:P含量对分解速率影响显著(P<0.01)。这些结果说明,在极端干旱区,沙埋对凋落物的分解过程有显著的积极影响。

甘肃省某能源项目工程地质灾害危险性评估及防治措施研究

评估区位于河西走廊西段北部山前冲洪积平原,以砂砾石戈壁为主,风蚀沙埋、洪水冲蚀地质灾害发育。文章对地质灾害现状进行了评估,对工程建设可能引发的地质灾害进行预测,并提出了相应的防治措施。工程施工中应尽量减少扰动面积、及时回填,采取增加防尘网、洒水降尘等措施,防治风蚀沙埋地质灾害;在洪水沟修建导洪渠或挡水墙,截引、导排山区洪水,避免冲蚀建筑物,防治发生地质灾害。

砂卵石地层中浅埋暗挖马头门施工技术分析

砂卵石地层因其特殊的地质条件,给浅埋暗挖出入口施工带来了诸多挑战。文中以某城市高压线入地电力隧道工程为背景,分析了砂卵石地层中浅埋暗挖出入口施工的技术难点,并从施工工艺、支护措施、施工组织与管理等方面提出了相应的施工技术措施,为类似工程提供了有益的参考。

沙埋和种子大小对柠条锦鸡儿种子萌发、出苗和幼苗生长的影响

研究了沙埋深度和种子大小对内蒙古毛乌素沙地植被群落中占优势的柠条锦鸡儿(Caragana korshinskii Kom.)种子萌发、出苗、幼苗存活和生长的影响。结果表明,沙埋深度显著影响柠条锦鸡儿的种子萌发率、休眠率、出苗率、幼苗存活率及生物量。在0.5—2 cm的浅层沙埋下,种子萌发率、出苗率、幼苗存活率及生物量最高,休眠率最低;沙埋深度≥4 cm时,柠条锦鸡儿的种子萌发率、出苗率、幼苗存活率及生物量随着沙埋深度增加显著降低,而休眠率却显著升高;沙埋深度≥12 cm时,柠条锦鸡儿种子不能够出苗,幼苗也不能够存活。种子大小对柠条锦鸡儿种子萌发率没有显著影响,但对出苗率、幼苗存活率及生物量影响显著。在各个沙埋深度下,不同大小的柠条锦鸡儿种子间的萌发率没有显著差异。当沙埋深度≤6 cm时,不同大小的柠条锦鸡儿种子在同一沙埋深度下的出苗率间没有显著差异;但当沙埋深度≥8 cm时,在同一沙埋深度下,大种子的出苗率显著高于中种子和小种子的出苗率,而中种子和小种子出苗率间没有显著差异。0.5—10 cm的沙埋深度中,除6 cm和8 cm深度下中种子和小种子萌发幼苗的生物量间没有显著差异外,其余深度下,大种子萌发的幼苗的存活率及生物量显著高于同一沙埋深度下中种子萌发的幼苗的存活率及生物量,后者又显著高于小种子萌发的幼苗的存活率及生物量。可能正是种子萌发对沙埋环境的忍耐或响应能力以及种子的多态性提高了柠条锦鸡儿在毛乌素沙地的适合度,为其在流动或半流动沙丘环境中成功定居并形成优势群落奠定了基础。

海滨沙地砂引草对沙埋的生长和生理适应对策

在2009和2010年春夏季通过对烟台海滨沙地自然生长的耐沙埋植物砂引草(Messerschmidia sibirica Linn)进行不同厚度(1/3株高-轻度沙埋、2/3株高-中度沙埋、3/3株高-重度沙埋)沙埋试验,并测定沙埋过程中土壤温度、土壤含水量、叶片鲜重(FW)、干重(DW)、植株相对高度(%)、叶片细胞膜透性、相对含水量、(丙二醛)MDA、抗氧化酶活力(过氧化物岐化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT))、脯氨酸、可溶性糖含量的变化以探讨砂引草抗沙埋的生长和生理调节策略,为未来砂引草的科学管理和应用提供理论指导。结果表明,在沙埋第5天、第10天,随着沙埋厚度的增加土壤温度下降,土壤含水量增加,植株相对高度和整株砂引草叶片FW和DW呈上升趋势,同一植株沙上叶片FW、DW不断增加,沙下叶片FW、DW则不断减少。在沙埋第5天,不同厚度沙埋处理的植株叶片细胞膜透性和MDA含量均较对照低,而整株植物叶片SOD、CAT活力和脯氨酸含量均较对照高。同一植株沙下叶片细胞膜透性、MDA、脯氨酸含量及SOD和CAT活力均低于沙上叶片。研究表明,轻度和中度沙埋使沙上叶片受到地面热辐射、干旱、高温胁迫影响,导致叶片内含水量下降,膜脂过氧化加强,膜受损。但同时叶片中快速激活的保护酶和积累的脯氨酸抑制膜脂过氧化可能是保护沙埋后沙上叶片能快速生长的重要生理调控机理。在重度全埋枝叶处于沙下时,沙下缺氧和黑暗抑制了叶片呼吸作用和光合作用,同时缺氧也抑制了叶片对营养物质的消耗使叶片成为茎尖生长的物质和能量供体,促使暗中茎顶端不断的延伸生长冲出沙土再生。砂引草耐全埋的再生能力进一步表明,砂引草在海岸沙丘固沙、保滩、护岸和植被恢复上将具有重要开发和应用前景。

四种沙埋深度对羊草种子出苗和幼苗生长的影响

模拟羊草种子自然埋藏深度状况,设计了0、1、2和3cm共4种沙埋深度实验,研究了其出苗率及幼苗生长对沙埋深度的响应。结果表明,羊草种子出苗率随埋深的增加呈下降趋势,0cm时出苗率最高为89.3%,1和2cm时分别为81.5%和78.5%,3cm时最低(73.5%),与前三者具有显著差异。埋深对地上株高生长动态的影响主要表现在,播种后25d,0～2cm处理的羊草幼苗地上株高均高于3cm处理;播种后36d,4种处理的地上株高没有显著差异。此外,随着埋深的增加,羊草幼苗的根长、根数、根冠比呈先上升后下降趋势,叶片数呈下降趋势,绝对株高则呈上升趋势。表明1～2cm沙埋深度更适宜于羊草种子出苗及幼苗生长,而0或3cm埋深不利于其出苗和幼苗生长。

碟果虫实种子萌发对策及生态适应性

以荒漠地区1年生植物碟果虫实为材料,通过温度、光照、GA3和沙埋对其种子萌发和幼苗出土影响的研究,旨在探索其种子萌发对策及生态适应性。结果表明:(1)碟果虫实种子在0h光照的条件下,萌发的最佳恒温是20℃,萌发率为(18.00±1.41)%,最佳的变温是15℃/25℃,萌发率为(28.00±2.31)%。(2)除了5℃/25℃,碟果虫实在变温条件下的萌发率极显著高于恒温下的萌发率(P<0.01)。(3)碟果虫实种子萌发时属于需暗型,而且存在休眠现象。(4)在野外沙埋的条件下,碟果虫实的出苗率随着沙埋深度的增加极显著下降(P<0.05),适宜的出苗深度为0.00—2.00cm,最佳的出苗深度是0.50cm,出苗率为(16.80±2.85)%。(5)碟果虫实种子萌发具有迅速萌发和推迟萌发两种对策;根据种子萌发率和种子活力可以推断,碟果虫实具有持久性土壤种子库的特点。结合碟果虫实萌发时的自然生存环境和本研究的结果,显示出其种子萌发对策与栖息地的生境具有高度的生态适应性。

沙埋和种子大小对固沙禾草沙鞭的种子萌发与幼苗出土的影响

该文研究了野外条件下不同深度的沙埋对沙鞭(Psammochloavillosa)种子萌发和幼苗出土的影响,以及温室条件下种子大小对不同深度沙埋后的种子萌发和幼苗出土的影响。结果表明,沙埋深度显著影响沙鞭的种子萌发率、幼苗出土率和种子休眠率。沙子表面的种子不能萌发。2cm的浅层沙埋时的种子萌发率和幼苗出土率最高,1cm沙埋的种子萌发率和幼苗出土率次之。沙埋深度超过2cm之后,沙鞭的种子萌发率和幼苗出土率与沙埋深度呈负相关。2cm的种子休眠率最低。从2～12cm,种子休眠率随着沙埋深度的增加而增加。在幼苗能够出土的深度(1～6cm),幼苗首次出土所需的时间随着沙埋深度的增加而延长。种子大小对沙鞭的种子萌发率没有显著影响。但是在深层沙埋(6cm)时,与小种子相比,大种子产生的幼苗的出土率较高。从2～6cm,大种子形成的幼苗的茎长度都较长。

沙埋对沙米幼苗生长、存活及光合蒸腾特性的影响

沙米(Agriophyllum squarrosum)是藜科沙蓬属1年生沙生植物,广泛分布于我国各主要沙漠和沙地中。为了解沙埋对沙米生长、存活和光合蒸腾特性影响,2010—2011年在科尔沁沙地研究了不同沙埋深度下沙米幼苗高度、存活率、光合速率、蒸腾速率、气孔导度和水分利用效率的变化,结果表明:沙米具有极强的耐沙埋能力。埋深为株高25%时,沙米幼苗存活率和株高显著增加,埋深为株高50%—100%时,其株高和存活率虽有下降,但与非沙埋对照差异不显著。当沙埋深度超过株高后,其株高和存活率急剧下降,但沙埋达到株高266%时仍然有部分幼苗存活。沙埋第5天,随着沙埋深度的增加,沙米幼苗的光合速率缓慢下降,蒸腾速率、气孔导度和水分利用效率均呈波动式变化。随着沙埋时间的延长,和对照相比,沙米幼苗的光合速率、蒸腾速率和气孔导度均随沙埋深度增加而大幅度下降,但水分利用效率仍呈波动式变化。沙埋第15天时其光合速率、蒸腾速率、气孔导度和水分利用效率分别较非沙埋对照下降了86.7%、89.0%、90.0%和4.2%。相关分析表明,沙米幼苗的存活率和高生长与其光合速率、蒸腾速率、气孔导度变化呈显著正相关,而其光合速率、蒸腾速率、气孔导度之间也呈显著正相关,但与水分利用效率的相关性未达到显著水平。沙埋胁迫下沙米幼苗存活率下降和生长抑制不仅源于沙埋造成幼苗顶土困难和光合面积减少,光合速率、蒸腾速率和气孔导度下降也是重要原因。

海滨沙滩单叶蔓荆对沙埋的生理响应特征

海滨沙滩单叶蔓荆(Vitex trifolia L.var.simplicifolia)是优良的抗沙埋地被植物。以烟台海岸沙地单叶蔓荆为材料,通过不同厚度沙埋过程中沙上和沙下叶片抗逆生理指标的测定以揭示其抗沙埋生理调控机制。结果表明,轻度和中度沙埋5d,成株和幼株整株叶片细胞膜透性增大、POD和SOD活力增高、MDA和脯氨酸含量和叶片相对含水量(RWC)增加、可溶性糖含量下降。但同株沙上叶片细胞膜透性、MDA含量、SOD和POD活力和可溶性糖含量均高于沙下,而沙上叶片脯氨酸含量低于沙下叶片。在轻度和中度沙埋10d,沙上叶片细胞膜透性、MDA和可溶性糖含量、叶片POD活力降低,叶片SOD活力仍有小幅度增高,但脯氨酸含量增加,沙上叶片生长旺盛。研究表明,沙埋下叶片抗氧化酶活力和脯氨酸含量与细胞膜透性和膜脂过氧化成正相关。沙埋使植株上部叶片接近沙表面而经受干旱和地面热辐射胁迫引起细胞膜脂过氧化加剧和细胞膜透性加大。同时沙埋也使沙下叶片遭遇黑暗和缺氧胁迫诱导细胞内膜脂过氧化,但也激活了叶片抗氧化酶保护系统和叶片脯氨酸的积累抑制细胞膜脂过氧化维护细胞膜的稳定。因此在沙埋过程中,叶片快速响应沙埋胁迫激活叶片抗氧化酶系统抑制膜脂过氧化作用维持氧自由基和抗氧化酶系统的动态平衡在单叶蔓荆适应轻度和中度沙埋,维护沙上叶片旺盛生长中起重要作用,也是重度全埋下沙下植株茎顶端能快速延伸弯曲生长最后顶出沙面再生的主要生理保护原因。

毛乌素沙地游击型克隆半灌木羊柴对局部沙埋的反应

不同程度的沙埋是生长在于旱和半干旱区内陆沙丘的植物经常遭遇的事件，沙埋可以改变植物所处的生物和非生物环境条件。已有研究表明不同程度的沙埋对于植物的影响不同。轻微程度的沙埋可以增加植物高度、促进生物量的积累和新生分株的产生。如果沙埋强度不断增加，对植物的影响由正效应逐渐转变为负效应。即超过一定沙埋阈值后，沙埋会削弱植物的生长，甚至影响植物的存活。干旱和半干旱区内陆沙丘中常常生长着许多克隆植物，克隆整合常常可以缓解克隆植物分株所遭受的局部环境胁迫。根茎型克隆植物羊柴（Hedysarum laeve）是毛乌素沙地的优势半灌木之一，也是当地重要的固沙植物。为了探讨克隆整合的作用是否可以提高沙埋阈值，并有助于羊柴忍受高强度的沙埋，以其为研究对象开展了野外实验。结果表明：轻微程度的沙埋（例如沙埋深度是原始羊柴分株高的10％-20％）可以加速羊柴分株的高生长，提高叶片生物量、茎生物量以及整个地上部分的生物量。高强度的沙埋（例如沙埋深度是原始羊柴分株高的80％-100％）会削弱羊柴分株的存活和生长。在与不遭受沙埋分株相连的情况下，羊柴分株遭受沙埋的阈值高于没有分株相连的，而且在高强度的沙埋下，前者（有分株相连的遭受沙埋的分株）比后者（没有分株相连的遭受沙埋的分株）在株高增量、茎生物量、叶片生物量以及地上分株生物量上都要显著高。这暗示着克隆整合提高了羊柴遭受沙埋的阈值并有助于羊柴分株忍受高强度的沙埋。