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。     

荒漠生境下柠条幼苗存活与生长对水分添加和沙埋的响应

为了解柠条幼苗生长与存活对荒漠生境下沙埋和水分添加的响应,在宁夏灵武白芨滩国家级自然保护区南部荒漠灌丛群落,通过水分添加和持续沙埋处理,研究了柠条幼苗的存活与生长对水分添加和沙埋的响应。结果表明:水分添加有利于柠条幼苗的存活与生长,幼苗最终留存率显著提高(P0.05),且幼苗株高、单株叶面积和总干质量显著增大(P0.05),侧根数、小叶数、相对生长率和净同化率也均有不同程度增大。沙埋抑制柠条幼苗的存活与生长,试验结束时,沙埋幼苗在水分添加条件下的最终留存率显著低于无沙埋幼苗(P0.05);除株高外,沙埋幼苗的基径、小叶数、单株叶面积、总干质量、相对生长率等均显著减小(P0.05),主根长、侧根数和净同化率也不同程度减小。沙埋幼苗的比叶面积、比根长和比枝长均显著大于无沙埋幼苗(P0.05),但前者的根冠比显著小于后者(P0.05);沙埋幼苗的比根长在水分添加条件下显著减小(P0.05),但不论是否沙埋,幼苗根冠比、比叶面积和比枝长在有无水分添加间均差异不显著。     

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。     

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 (HedysarumlaeveMaxim .)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 6 7%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 6 7%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%、6 7%和 10 0 %沙埋 )的幼苗明显地高于非沙埋的对照。     

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.     

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.     

鄂尔多斯高原4种主要沙生植物种子萌发与出苗对水分和沙埋的响应

 羊柴 (Hedysarum leave)、柠条 (Caragana korshinskii)、籽蒿 (Artemisia sphaerocephala)与油蒿 (Artemisia ordosica)为鄂尔多斯高原广泛分布的植物种,也是该地区飞播选用的主要植物种。由于飞播为鄂尔多斯高原植被恢复的重要手段,而飞播后植物种子的出苗率与土壤水分及沙埋密切相关。因此,上述植物种子萌发与出苗对水分和沙埋反应的研究对当地退化生态系统的恢复具有重要意义。通过温室控制实验,对4种植物萌发特性及出苗与水分和沙埋深度的关系进行研究,阐明了种子萌发最适合的水分条件和沙埋深度。结果表明,4种植物种子萌发与出苗最适宜的供水量都接近于当地种子萌发时期的平均降水量, 最适宜的沙埋深度为0.5～1 cm, 过多的水分和过深的沙埋会降低种子萌发与出苗,反映了4种植物对沙区环境的生态适应。4种植物最适宜的供水量均为123 ml,羊柴、柠条、籽蒿和油蒿的最适沙埋深度分别为：0.5、1.0、0.5和0.5 cm。根据实验结果及当地气候特点,鄂尔多斯地区的飞播时间应由6月初提前至5月中下旬,以提高飞播植物的出苗率。     

沙埋与水分对科尔沁沙地主要固沙植物出苗的影响

蒿属半灌木乌丹蒿(Artemisia wudanica)、白沙蒿(A. sphaerocephala)、差不嘎蒿(A. halodendron)是科尔沁沙地的主要固沙植物。其中乌丹蒿和差不嘎蒿是科尔沁沙地的本土植物,白沙蒿为来自于库布齐沙漠、毛乌素沙地的飞播植物。设置了 5个沙埋深度(0.5、1.0、1.5、2.0和3.0 cm)和 4个水分梯度(86、171、257和 342 mL,分别模拟每月25、50、75和100 mm的降雨量),以探讨3种植物幼苗出土对沙埋和水分的响应。结果表明,沙埋与水分均显著影响着3种蒿属植物的幼苗出土(P < 0.001)。3种植物最适沙埋深度在0.5-1.5 cm范围内,萌发出土时适宜水量要高于当地种子萌发期的平均降水量(50 mm/月)。两种固沙先锋植物乌丹蒿和白沙蒿的种子出苗率均显著高于差不嘎蒿,乌丹蒿较白沙蒿也明显为高,尤其在水分缺乏时,表现出两种先锋植物种子出苗对干旱有更好的适应性。协方差分析表明,乌丹蒿幼苗死亡率显著高于白沙蒿和差不嘎蒿(P < 0.05),在达到75 mm/月降水量时,3种植物的出苗较好,但不能满足乌丹蒿幼苗生长对水分的需求,而实际种子萌发期的降水量平均只有50 mm/月。因而降水的缺乏导致乌丹蒿种群更新出现问题,加之飞播植物的竞争,使得近几年科尔沁沙地较多乌丹蒿种群出现衰退。     

差巴嘎蒿幼苗对沙埋的生态适应和生理响应

沙埋对于沙漠植物存活和生长影响的研究文献已有很多,但迄今有关沙漠植物对于沙埋生理生态适应的研究还很少见有报道。差巴嘎蒿(Artemisia halodendron)广泛分布于我国半干旱沙地中,是流动半流动沙地的优势种。为了解沙生植物对沙埋的生态适应及其生理响应,2010—2011年在内蒙古科尔沁沙地研究了不同沙埋深度下差巴嘎蒿幼苗的存活率、株高等生长特性和渗透调节物质含量,保护酶活性和膜透性的变化,得到以下结果:沙埋对差巴嘎蒿幼苗的存活与生长有着显著影响,特别是沙埋深度超过其株高后,其存活率和高生长受到严重抑制,但和非沙生植物相比,差巴嘎蒿幼苗具有很强的耐沙埋能力,即使沙埋深度达到其株高的200%时仍有部分幼苗存活。随着沙埋深度的增加,差巴嘎蒿幼苗叶片含水量没有显著变化,但过氧化物歧化酶活力降低,过氧化物酶活性增强,过氧化氢酶活性变化不明显,脯氨酸含量增加,可溶性糖含量下降,丙二醛含量下降,膜透性增强。和水分、盐分胁迫不同,沙埋并未造成差巴嘎蒿幼苗的水分亏缺,沙埋胁迫下差巴嘎蒿幼苗死亡率增加、生长受到抑制的主要原因是沙埋导致植物光合面积下降,沙埋叶片无法进行正常呼吸和部分幼苗无法破土生长,但沙埋胁迫下脯氨酸含量的和过氧化物酶活性增强分别在渗透调节和保护细胞膜免受损伤中起到了关键作用。     

Effects of watering regime and depth of burial on seedling emergence of four dominant psammophytes in the Mu Us sandy land, Inner Mongolia, China, and relevance to revegetation of a desertified region

Four dominant psammophyte species are air-seeded in attempts to revegetate the Mu Us sandy land in Inner Mongolia, but seedling emergence is low. This study sought to clarify the behaviour of seedling emergence under different water supply and sand burial regimes to improve the technology of air seeding. Seeds were buried in sand at depths of 0, 0.5, 1, 1.5, 2, 3 and 5 cm and supplied one time with 2.5, 5, 7.5, 10, 15, 20, 30 or 40 mm of water or with 2.5 mm once every 3 days for 30 days. Our study showed that a 0.5-cm burial depth and under 10- to 20-mm single-watering regimes resulted in highest seedling emergence percentage and rate of these four species. All emerged seedlings of Caragana korshinskii , Hedysarum laeve and Artemisia ordosica died under 2.5, 5 and 7.5 mm of watering, respectively, but seedling mortality was <50% when the single-watering regime was >10 mm. In the field, germination of the four species increased from the top of the dune to the middle of the leeward side and then to the bottom. Based on precipitation pattern in the Mu Us sandy land, our study indicates that 0.5-cm burial depth and a single-watering regime of >7.5 mm is the threshold condition for seedling emergence.     

Effects of burial in sand and water supply regime on seedling emergence of six species

BACKGROUND AND AIMS: Air seeding has long been regarded as a quick and successful measure for vegetation rehabilitation in China. However, seedling emergence of often-used species including Agriophyllum squarrosum, Artemisia sphaerocephala, Artemisia ordosica, Hedysarum fruticosum, Caragana korshinskii and Medicago sativa is low. Experiments were conducted under controlled conditions to study the effects of sowing depth and water supply on seedling emergence, in order to understand the requirements for increasing seedling emergence. METHODS: Seeds were exposed to different environments of burial and water supply regimes in PVC pots (7 cm in diameter and 11 cm in height) under the same light intensity and alternating temperature regimes in a growth chamber. KEY RESULTS: Seedlings of three species (Agriophyllum squarrosum, Artemisia sphaerocephala, Artemisia ordosica) with relatively light seeds emerged well at a 0.5 cm sowing depth under a 7.5 and 10 mm water supply regime. However, few seedlings of these species emerged when the sowing depth was over 1 cm or when water supply was 5 mm. Seedlings of Caragana korshinskii, Hedysarum fruticosum and Medicago sativa emerged from sowing depths of 0.5-4 cm, 0.5-3 cm, and 0.5-4 cm, respectively, under both 7.5 and 10 mm water supply regimes. Under a 5 mm water supply regime, seedlings of these species also emerged at over 1 cm sowing depth. Seeds of all six species sown on the surface of sand did not germinate, and seedlings did not emerge when they were sown at depths greater than 6 cm. CONCLUSIONS: Based on these experiments, a 0.5 cm sowing depth resulted in the highest seedling emergence and it is concluded that this is the optimal sowing depth for seedling emergence of all six species.     

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.     

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.