沙埋对半灌木羊柴幼苗的存活、生长和生物量分配的影响

羊柴(Hedysarum laeve Maxim.)是中国北方沙地飞播进行植被恢复与重建的主要植物种.在晚春与夏初的出苗期 ,羊柴幼苗经常遭受不同深度的沙埋.研究旨在探明沙埋对羊柴幼苗存活、生长和生物量分配格局的影响.6周的沙埋试验结果显示:当沙埋深度达到其株高时,约有70%的幼苗死亡 ;沙埋深度达到其株高的133%时,可使羊柴幼苗全部死亡;沙埋深度分别为株高的33%和67% 时,羊柴幼苗的整株生物量、叶片生物量、根系生物量以及相对生长率都相应地高于非沙埋的对照(即:0%沙埋).与非沙埋的对照相比,羊柴幼苗在33%和67%沙埋条件下并不显著地改变其生物量分配格局;并且羊柴幼苗的叶数和株高与对照无明显差异,但试验期内的新生叶数是沙埋处理(即:33%、67%和100%沙埋)的幼苗明显地高于非沙埋的对照.     

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干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。     

沙埋对花棒种子萌发和幼苗生长的影响

为探讨花棒(Hedysarum scoparium)种子萌发和幼苗生长对沙埋的响应,并为固沙造林、水土保持提供理论基础,研究了6种沙埋深度(0、1、2、3、4和5 cm)对花棒种子萌发和幼苗生长的影响.结果表明,埋深对花棒幼苗出土率、首次出苗时间、单株叶片数、幼苗生长高度以及生物量的分配均有极显著影响(P<0.001).种子出苗率在2cm沙埋下达到最高(95.6%),在5 cm沙埋下最低(43.4%);幼苗最大高度(11.6cm)、绝对株高(13.9cm)和最大地上生物量(26.7mg)均出现在2cm的埋深,幼苗最小高度(3.3cm)、最小根长(4.3cm)和最小地上生物量(5.3mg)出现在5cm的埋深;生物量分配随沙埋深度增大而更多地分配给地下部分.2cm的沙埋是花棒种子萌发和幼苗生长的最佳深度.     

沙埋对无芒雀麦种子萌发和幼苗生长的影响

研究了沙埋对浑善达克沙地植物群落中多年生禾草无芒雀麦种子萌发、出苗和幼苗生长的影响.结果表明:无芒雀麦种子能在深度≤12cm的沙埋中萌发,≤8cm的沙埋中出苗.在此范围内,随着沙埋深度的增加,无芒雀麦种子的萌发率和出苗率逐渐降低.沙埋深度在植株高度的33%时,无芒雀麦1周龄和2周龄幼苗均全部存活,且总生物量≥对照(未沙埋幼苗);当沙埋深度增至植株高度的66%时,1周龄和2周龄幼苗的存活率分别降至70%和25%,生长也受到抑制;而遭受全部沙埋时,1周龄和2周龄幼苗均不能存活.遭受沙埋后,无芒雀麦幼苗分配较多的生物量用于地上部分的生长,其生物量分配模式改变可能是无芒雀麦幼苗对沙埋环境的重要适应对策.     

小叶锦鸡儿幼苗对沙埋的生态适应和生理响应

以中国半干旱地区固定、半固定沙地分布最广泛的灌木种之一小叶锦鸡儿(Caragana microphylla)为对象,于2010～2011年在内蒙古科尔沁沙地测定了不同深度沙埋下其幼苗的存活率、株高、渗透调节物质含量、保护酶活性和膜透性变化,探讨小叶锦鸡儿对沙埋的生理响应特征。结果显示:(1)与对照相比,当沙埋深度为小叶锦鸡儿幼苗株高的25%～75%时,存活率差异不显著;当沙埋深度为50%～75%时其存活率增加,但株高明显下降,沙埋对其生长具有抑制作用;当沙埋达到其株高100%时幼苗全部死亡。(2)不同沙埋深度对小叶锦鸡儿幼苗叶片细胞膜透性无显著影响,对沙埋深度为其株高25%时,叶片含水量增加,MDA含量下降,POD活性增强,细胞膜未受到伤害;沙埋深度为其株高50%～75%时,叶片含水量下降,MDA含量增加,但SOD和POD对细胞膜起到了协同保护作用,细胞膜也未受到损伤;沙埋深度为株高25%～75%情况下,细胞膜未受到损伤,可溶性糖和脯氨酸含量变化不明显。研究认为:小叶锦鸡儿幼苗只能耐受部分沙埋,完全沙埋则会导致其幼苗全部死亡,沙埋胁迫下可溶性糖和脯氨酸等渗透调节物质含量反应迟缓可能是其耐沙埋能力较弱的主要生理原因之一。     

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。     

沙埋与供水对毛乌素沙地两种重要沙生植物幼苗生长的影响

沙埋与水分是影响毛乌素沙地植物存活、生长的关键因子。通过沙埋与供水的野外控制实验,研究了毛乌素沙地重要物种柠条(Caragana korshinskii)与羊柴(Hedysarum laeve)幼苗存活及生长对沙埋和供水的反应。结果表明:适量的沙埋(0.25H-0.50H沙埋,H为苗高)与供水(柠条50mm/月,羊柴75mm/月)可以促进幼苗生长,过量的沙埋与供水则会抑制生长甚至导致幼苗死亡。柠条幼苗在50mm/月供水,0.25H、0.50H沙埋,羊柴幼苗在75mm/月供水,0.25H、0.50H沙埋下具有较高的相对生长速率和净同化速率;幼苗在1.0H和1.25H沙埋下全部死亡。供水量较小、沙埋较浅时,两种植物根冠比均较高;当供水量最大时,柠条幼苗根冠比在沙埋最深和没有沙埋时较大,而羊柴幼苗根冠比仍在没有沙埋时最高。     

毛乌素沙地4种沙生植物种子萌发及出苗对沙埋及单次供水的响应

飞播固沙作为一项快速而有效的现代化治沙措施,已成为毛乌素沙地植被恢复与重建的重要手段。但此项技术仍存在一些问题,如飞播后成苗率较低等。以毛乌素沙地优势植物柠条（Caragana korshinskii）、羊柴（Hedysarum laeve）、油蒿（Artemisia ordosica）与籽蒿（Artemisia sphaerocephala）为研究对象,人工控制7个沙埋深度和9个水分梯度,研究4种植物种子萌发和幼苗出土对沙埋和单次供水的响应,阐明多大的单次降雨才能保证种子萌发与幼苗出土及出土幼苗的维持时间。结果表明,对于柠条和羊柴质量相对较大的种子而言．种子主要在供水量10．20mm,埋藏深度为0．5．2cm的条件下出苗,但在埋藏深度为5cm时这两个物种仍有少量种子能够出苗;而对于油蒿和籽蒿质量相对较小的种子而言,种子主要在供水量10．20mm,埋藏深度为0．5cm时出苗,当埋藏深度为1．5cm和2cm时不能顺利出苗。沙埋深度0．5cm条件下,4个物种具有最高的出苗率和出苗速率。在试验期内（30d）,柠条、羊柴和油蒿出土幼苗分别在单次供水量为5、7．5mm和7．5mm时全部死亡,在单次供水量高于15mm时4个种的出土幼苗死亡率大多低于50％,且随供水量的增加出土幼苗死亡率降低。结合毛乌素沙地降雨特点与本次实验结果,0．5cm的沙埋及7．5mm以上的单次降雨是上述4种植物自然条件下出苗较好的重要条件。     

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0.5-2cm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.     

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。     

Effects of sand burial on survival, growth, gas exchange and biomass allocation of Ulmus pumila seedlings in the Hunshandak Sandland, China

BACKGROUND AND AIMS: In the last decade, the number of young plants of Ulmus pumila in the Hunshandak Sandland has decreased sharply because of severe sand burial, and their ecological protective function has been weakened. In order to develop an understanding of the tolerance of U. pumila to sand burial and to suggest reasonable measures to protect the sparse-elm-grassland ecosystem, the effects of burial on the survival, growth, photosynthesis and biomass allocation in U. pumila were studied. METHODS: Seedlings were buried at five different depths in pot experiments: no burial (control), partial burial (33 % and 67 % stem height), and complete burial (100 % and 133 % stem height). Growth analyses and measurements of photosynthesis were carried after the plants had been uncovered. KEY RESULTS: All the plants survived partial burial, but about 30 % and 80 % of the seedlings died as a result of the 100 % and 133 % sand burial treatments, respectively. The numbers of newly produced leaves and branches, and the height of the stems of the seedlings in the 33 % and 67 % burial treatments during the period of the experiment were significantly greater than those in the control. Furthermore, net photosynthetic rate, transpiration rate and water use efficiency were also elevated by the partial burial, but not affected by burial time. This might be attributed to the increased root length, which improved water acquisition. The biomass and biomass allocation of the seedlings were significantly changed by the burial treatments and burial times. The biomass was enhanced by partial burial but was reduced by complete burial at each burial time. However, the biomass allocation was not significantly changed by the 33 % and 67 % sand burial treatments 2 or 4 weeks following the burial. CONCLUSIONS: Ulmus pumila was shown to be tolerant to partial sand burial, but must be protected from complete burial.     

Effects of sand burial on purple sandgrass (Triplasis purpurea): the significance of seed heteromorphism

The effects of sand burial on seedling emergence, growth, and reproduction of the annual dune grass Triplasis purpurea (Walt.) Chapm. were examined. This species matures heteromorphic seeds on panicles enclosed by leaf sheaths along tiller nodes in a characteristic pattern: seeds at the lower nodes are heaviest, whilst those at the upper nodes are lightest. Field excavation of seedlings revealed that seedlings can emerge from seeds buried to 4 cm.In one experiment in an incubator, seeds collected from the upper and lower tiller nodes of glasshouse-grown parental plants were buried in sand at depths of 0, 3, and 6 cm. As expected, the effects of seed depth on seedling growth were due to delayed emergence for seedlings from buried seeds. However, seedlings from the larger seeds from the lower nodes of parental plants had significantly greater dry mass than those from the lighter seeds from upper nodes. Because lower node seeds are the most likely to become buried by sand in nature, the seed heteromorphism may be adaptive in the coastal environment.A second experiment involved the burial of seedlings to 0, 50, or 100% of their height. Although most seedlings did not survive complete burial (100% height), for partially buried seedlings (50% height) there was high survival and a marked stimulation of growth and eventual reproduction compared to unburied seedlings. This stimulation could be linked to increased root growth for seedlings that had been buried. The ability to respond positively to sand accumulation may represent an adaptation to the dynamic dune environment.     

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 microenvironments 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 environmental stress encountered by ramets.A rhizomatous clonal semishrub,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 biomass 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.     

沙埋对中间锦鸡儿幼苗生长发育的影响

在沙地环境中,沙埋是影响植物幼苗存活的一个重要因素。中间锦鸡儿(C arag ana interm ed ia)是浑善达克沙地和毛乌素沙地固定和半固定沙丘上常见的一种沙生灌木。为了研究沙埋对其幼苗存活以及生长的影响,对出土后生长了1个月的中间锦鸡儿实生苗进行沙埋实验,实验处理:TC为对照,T1为沙埋幼苗地上部的1/3,T2为沙埋幼苗地上部的1/2,T3为沙埋到幼苗的近顶端。实验持续4周,每周收获1次。结果表明:TC、T1和T2没有幼苗死亡,T3有20%的幼苗死亡;沙埋对中间锦鸡儿幼苗的生物量、根冠比、根生物量比和茎生物量比有显著影响,叶片生物量比各处理没有显著差异,沙埋对幼苗的叶面积没有显著影响,在第1周T2和T3的小叶片面积显著大于TC,在其余3周,TC与T1、T2的小叶片面积没有显著差异,与全埋则有显著差异;沙埋对幼苗的相对生长速率(RGR)和净同化速率(N AR)产生显著影响,实验结束时,T3处理幼苗的RGR和N AR显著地小于其它3个处理。上述结果表明,部分沙埋(T1、T2)对沙生灌木中间锦鸡儿幼苗的生长发育没有显著的影响,而全部沙埋(T3)对其幼苗的生长发育产生显著的抑制,中间锦鸡儿是一个比较耐沙埋的物种。     

Seedling growth variation in response to sand burial in four <Emphasis Type="Italic">Artemisia</Emphasis> species from different habitats in the semi-arid dune field

We compared seedling growth of four Artemisia species dominated at different habitats to determine whether interspecific seedling growth variation of a same genus in tolerance to burial can be used to explain plant distribution in the sand dune field. Interdune lowland species, Artemisia gmelinii, stabilized dune species, A. frigida, semi-stabilized dune species, A. halodendron, and active dune species, A. wudanica were selected. Seedlings grown for 3 weeks were treated at five burial depths for three burial times in pot experiments. Species from the habitats with little burial had smaller survival rate, dry weight and stem elongation speed than those from the habitats with intensive burial when buried. Furthermore, when buried, the former tended to adjust biomass allocation between shoot and root and produce adventitious buds, while the latter tended to maintain a constant root:shoot ratio and produce adventitious roots. We conclude that (1) seedlings of species with a long evolutionary history of exposure to sand burial (from the active sand dune), show quicker stem growth when buried than do seedlings of species from the habitats with little or no sand burial; (2) seedlings of species which do not change root:shoot ratio might be more tolerant of sand burial than those do; (3) seedlings of species from the habitats with intensive sand burial is prone to produce adventitious roots and seedlings of species from the habitats with little or no sand burial tend to produce adventitious buds when buried.