种子萌发和幼苗生长对沙丘环境的适应机制

综述了植物沙生适应机制的研究及其进展.一些植物的种子在刚成熟时具有休眠特性.种子需要适度沙埋促进萌发并实现幼苗定居,但过度沙埋则会抑制种子萌发和出苗.在沙层深处,没有萌发的种子会进入休眠状态,形成土壤种子库.幼苗通过增加节数和延长节间来适应沙埋.沙埋深度超过植物的忍耐限度会抑制幼苗生长,甚至导致幼苗死亡.沙生植物必须适应其他环境因子,如盐风与土壤盐分、昆虫采食、土壤养分亏缺等,才能在沙丘上成功生长.沙蚀可导致幼苗根系暴露并干燥脱水.一些沙漠植物的幼苗在萌发后可忍耐一段时期的干燥,水分条件得到满足之后,幼苗能够恢复生长.     

Is zonation on coastal sand dunes determined primarily by sand burial or by salt spray? A test in New Zealand dunes

There has long been controversy on which environmental factor is the predominant determinant of community zonation on sand dunes. It is demonstrated here that, on a dune system in southern New Zealand, several environmental factors that could limit growth all vary along the sea-to-inland sand dune zonation: soil moisture, soil nutrients, wind exposure, sand burial, salt spray and soil salinity. Correlation of the responses of 30 species to experimental stress (burial, darkness, rooting-medium salinity and salt spray) with the zonation of the species in the field indicates that, in the four dune systems studied, sand burial and salt are both important, with salt generally being the more important. However, the relative importance of the factors differs between sites.     

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.     

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.     

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.     

Burial of canopy-stored seeds in the annual psammophyte Agriophyllum squarrosum Moq. (Chenopodiaceae) and its ecological significance

Agriophyllum squarrosum Moq. is a dominant annual on sand dunes in the arid regions of central Asia. A high percentage of seeds is retained on dead plants which become covered by moving sand, but little is known about the ecological significance of burial of canopy-stored seeds. We investigated the size and dynamics of the buried canopy-stored seed bank and effects of burial on seed germination. In March (during the windy season), May (beginning of the germination season), and July (middle of the growing season), the number of seeds per square meter in sample plots in the dunes was 623, 223 and 22, respectively, with 54.6, 30.6 and 12.9% of the total seeds retained on buried plant canopies. In a controlled experiment, more seedlings emerged from released (dispersed) than from canopy-stored seeds when burial depth was the same. No viable ungerminated released seeds were found, but 45–80% of the ungerminated canopy-stored seeds were viable. In general, with an increase in applied water germination of released seeds buried at a depth of 1 or 2 cm and of canopy-stored seeds buried at 1 cm increased, but regardless of watering regime few or no released seeds at 4 cm or canopy-stored seeds at 2 or 4 cm germinated. Significantly more seedlings emerged from plants buried in a horizontal than in a vertical position. Seedlings originating from buried canopy-stored seeds on an active dune accounted for only 5.4% of the total seedlings emerging, and most of them emerged later than those from released seeds. Thus, seed release is more effectively postponed in buried than in exposed canopies, and burial of canopy-stored seeds is a mechanism that helps regulate seed germination and seedling emergence of A. squarrosum on active dunes.     

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(Psammochloa villosa)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。2 cm的浅层沙埋时的种子萌发率和幼苗出土率最高,1 cm 沙埋的种子萌发率和幼苗出土率次之。沙埋深度超过2 cm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。2 cm的种子休眠率最低。从2 ~12 cm,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(1~6 cm),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(6 cm)时,与小种子相比,大种子产生的幼苗的出土率较高。从2~6 cm,大种子形成的幼苗的茎长度都较长。     

Emergent interactions influence functional traits and success of dune building ecosystem engineers

Aims Dune building processes are affected by interactions between the growth of ecosystem engineering dune grasses and environmental factors associated with disturbance such as sand burial and sea spray. Research investigating how species interactions influence dune community structure and functional trait responses in high abiotic stress environments is minimal. We investigated how species interactions influence the functional trait responses of three dominant dune grasses to common abiotic stressors.Methods We performed a multi-factorial greenhouse experiment by planting three common dune grasses (Ammophila breviligulata Fern., Uniola paniculata L. and Spartina patens Muhl.) in different interspecific combinations, using sand burial and sea spray as abiotic stressors. Sand burial was applied once at the beginning of the study. Sea spray was applied three times per week using a calibrated spray bottle. Morphological functional trait measurements (leaf elongation, maximum root length, aboveground biomass and belowground biomass) were collected at the end of the study. The experiment continued from May 2015 to August 2015.Important findings Species interactions between A. breviligulata and U. paniculata negatively affected dune building function traits of A. breviligulata, indicating that interactions with U. paniculata could alter dune community structure. Furthermore, A. breviligulata had a negative interaction with S. patens, which decreased S. patens functional trait responses to abiotic stress. When all species occurred together, the interactions among species brought about coexistence of all three species. Our data suggest that species interactions can change traditional functional trait responses of dominant species to abiotic stress.     

Seedling growth strategies and seed size effects in fourteen oak species native to different soil moisture habitats

 Seedling growth and morphology are thought to reflect evolutionary responses to habitat or influences of seed size. To test these hypotheses, we selected fourteen species of North American oaks differing in soil moisture habitat preference and seed size. Seedlings were grown for 1 – 2 years with abundant soil water and moderate soil nutrition in pots placed outdoors and in a common garden. Oak species native to xeric environments produced the smallest seedlings. Oaks from hydric soils had more shoot weight per unit of root weight and more height per unit of total plant weight than did mesic or xeric oaks. Essentially no differences in leaf area per unit of total plant weight were detected. Species with thinner and larger individual leaves tended to produce larger seedlings. Within species, seed size was generally unrelated to seedling growth, although results may have been complicated by uncontrolled genotypic variability. However, when species were compared, those with larger mean seed size produced larger seedlings. Root/shoot allometry, height growth and leaf thickness in the tested species may reflect evolutionary responses to soil moisture and flooding. Although seed size influenced seedling growth, no clear relationship between seed size and soil moisture habitat was found. Received: 26 March 1995 / Accepted: 30 November 1995     

广东亚热带森林木本植物幼苗生长特性研究

研究了广东亚热带42种木本植物幼苗的生长及其与物种的生态特性、生活型、种子大小的相互关系.较强光下(H,66.8 μmol m-2s-1)乔木幼苗的茎高和茎生物量显著高于灌木幼苗的相应值,但在较低光下(L,33.7 μmol m-2s-1)两者无显著差异.而阳性植物、耐阴植物和中间型植物之间,茎高和茎生物量无显著差别.乔木幼苗的叶面积和叶生物量比灌木幼苗大,但灌木幼苗的叶片数较乔木幼苗多.大种子种和小种子种幼苗之间,阳性植物、耐阴植物和中间型植物幼苗之间的叶片数和叶面积一般无显著差异.阳性植物幼苗比耐阴植物幼苗侧根数多.乔木幼苗的根生物量和根/茎比显著高于灌木幼苗在较高光下,阳性植物幼苗的根/茎比较耐阴植物幼苗高,但在较低光下无明显差异.45 d幼苗的根生物量与种子重量呈显著的正相关,而90 d幼苗无明显的相关.乔木幼苗个体生物量显著高于灌木幼苗.幼苗相对生长率和叶面积比的大小呈现如下顺序阳性植物＞中间型植物＞耐阴植物,但只有阳性植物和耐阴植物之间有显著差异.阳性植物、中间型植物和耐阴植物幼苗之间的单位叶率无显著差异.     

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。     

The role of reproductive phenology, seedling emergence and establishment of perennial Salix gordejevii in active sand dune fields

BACKGROUND AND AIMS: The function of sexual reproduction of perennials in restoration of vegetation of active dune fields frequently has been underestimated. The objective of this study was to evaluate the role of sexual reproduction of the perennial Salix gordejevii in the revegetation of active dunes. METHODS: Seedling emergence and establishment of S. gordejevii were examined both in controlled experiments (germination at different burial depths with different watering regimes) and in field observations in three dune slacks. The reproductive phenology and soil seed bank of S. gordejevii, the dynamics of soil moisture, the groundwater table and the landform level of three dune slacks were monitored. KEY RESULTS: Seeds of S. gordejevii began maturation on 1 May, and seed dispersal lasted from 8 May to 20 May. Seeds on the soil surface germinated significantly faster than those buried in soil (P<0.05). Seedling emergence was negatively correlated with landform level. When most seedlings emerged, there was a significantly positive correlation between soil moisture and seedling emergence (P<0.01). Rainfall was negatively correlated with seedling emergence. Seedling establishment was significantly and positively correlated with seedling emergence (P<0.05), and 72.3 % of the emergent seedlings were established at the end of the growing season. These results indicated that (a) seeds matured and dispersed before the rainy season; (b) seeds germinated as soon as they contacted a moist surface and relied more on soil moisture than on rainfall; and (c) more seedlings emerged at lower sampling points in dune slacks. CONCLUSIONS: In natural conditions, restoration of active sand dune fields generally commences with revegetation of dune slacks where sexual reproduction of perennials contributes greatly to species encroachment and colonization and hence plays an important role in restoration of active dune fields. Furthermore, aeolian erosion in dune slacks, leading to good soil moisture, facilitates seed germination, seedling emergence and establishment of S. gordejevii.     

The sediment burial depth and salinity control the early developments of Suaeda salsa in the Yellow River Delta

Sediment deposition is a common phenomenon in the estuary area. Pot control experiments were conducted to evaluate the interaction effects of sediment burial depth and salt stress on the seed germination and early seedling growth of Suaeda salsa (L.) Pall., an pioneer species of tidal wetland near the Yellow River Delta. The results showed that the percentage of seedling emergence, seedling emergence rate, seedling height, branch number, shoot biomass and root biomass were all significantly affected by salt stress and sediment burial depth. While the interaction of salt and burial depth significantly influenced the branch number, leaf biomass, shoot biomass and total plant biomass. Only 5 cm burial depth without salt stress should 6.25 ± 3.61% seedlings emergence. With the increasing of sediment burial depth and salt stress, percentage of seedling emergence, seedling emergence rate and plant height decreased significantly. However, under the salt treatment of 0 and 1%, the branch number increased dramatically with the increasing of sediment burial depth from 0 to 3 cm. The ratio of leaf to total biomass increased with increasing of burial depth, on the contrary, the ratio of root to total biomass decreased. 0–1 cm sediment burial depth was proved the suitable depths for seed germination of S. salsa in the coastal wetland of the Yellow River Delta. Our findings contribute to a better understanding of how to improve the seedling establishment of S. salsa under the dynamic changes of sediment deposition and salinity in the coastal wetland of the Yellow River Delta.     

西鄂尔多斯珍稀濒危植物长叶红砂种子萌发特征

珍稀濒危灌木长叶红砂是我国西北干旱地区荒漠植被中的古老残遗物种,对维持荒漠植被稳定具有重要作用.本文研究了长叶红砂种子特性,及其在不同光照、温度、土壤水分和沙埋等环境因子中的萌发策略.结果表明:长叶红砂种子具有高活力、高萌发率、耐贮存的特点,在光照和黑暗条件下均能很好地萌发.种子萌发的适宜温度为恒温20 ℃～25 ℃或变温15 ℃/25 ℃,萌发率高达93%.土壤含水量为2%时,种子开始萌发;土壤含水量为12%时,其萌发率最高,达89%.长叶红砂种子适宜的沙埋深度为1 cm,＞5 cm沙埋不出苗.沙埋深度对出苗率和幼苗生长高度的影响显著,而对幼苗质量的影响不显著.水分条件和沙埋深度是制约长叶红砂种子萌发和出苗的主要因素,而种子的高萌发率增加了幼苗生存的风险,不利于其应对荒漠极端环境变化.这种特性是长叶红砂濒危的重要原因之一.     

Zonation of vegetation on lacustrine coastal dunes: effects of burial by sand

More than 50 years ago it was proposed that zonation of major plant species on coastal dunes was determined by salt spray. Here, we argue against this hypothesis because (i) salt concentrations rarely exceed toxic levels; (ii) high precipitation in temperate latitudes washes the salt off the plants; (iii) major salt spray events occur in late autumn and winter when plants are dormant; and (iv) zonation also occurs on lacustrine dunes. Instead, we show evidence that zonation may be caused by burial because plant distribution was correlated with sand deposition and species were eliminated when burial exceeded their limits of tolerance, thus creating zones of different plant species. We conclude that in temperate regions (i) burial by sand may be among the most important factors in zonation, while salt spray may play a secondary role; and (ii) single environmental factors cannot be the determinants of a phenomenon as complex as species zonation.     

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。     

Soil Moisture Rather than Soil Nutrient Regulates the Belowground Bud Bank of Rhizomatous Species <i>Psammochloa villosa</i> in Arid Sand Dunes

In arid and semi-arid sand dune ecosystems, belowground bud bank plays an important role in population regeneration and vegetation restoration. However, the responses of belowground bud bank size and composition to sand burial and its induced changes in soil environmental factors have been rarely studied. In arid sand dunes of Northwestern China, we investigated belowground bud bank size and composition of the typical rhizomatous psammophyte Psammochloa villosa as well as three key soil environmental factors (soil moisture, total carbon and total nitrogen) under different depths of sand burial. Total buds and rhizome buds increased significantly with increasing burial depth, whereas tiller buds first increased and then decreased, with a peak value at the depth of 20–30 cm. Soil moisture increased significantly with sand burial depth, and was positively correlated with the number of all buds and rhizome buds. Soil total carbon concentration first increased and then decreased with sand burial depth, and total nitrogen concentration was significantly lower under deep sand burial than those at shallow depths, and only the number of tiller buds was positively correlated with soil total nitrogen concentration. These results indicate that soil moisture rather than soil nutrient might regulate the belowground bud bank of P. villosa, and that clonal psammophytes could regulate their belowground bud bank in response to sand burial and the most important environmental stress (i.e., soil moisture). These responses, as the key adaptive strategy, may ensure clonal plant population regeneration and vegetation restoration in arid sand dunes.     

Potential for seed bank formation in seven Great Lakes sand dune species

Seeds of seven dune species were collected from sand dunes of Lakes Erie and Huron and buried to various depths in a natural sand dune habitat along Lake Huron. The seed samples were then retrieved after varying lengths of time and examined for their germinability and dormancy. Results showed that buried seeds remained viable for at least 2.5 years and had the potential to form a persistent seed bank. Seed banks were larger and longer lasting at greater depths of burial than those at shallow burial depths. The results suggested that failure to verify the existence of effective seed banks in previous studies may be due to insufficient number of samples, shallow sample depth, local population variations, and fruiting events. Several species also possessed a temporary, aboveground seed reserve formed by retention of a small proportion of viable seeds on the previous year's inflorescences. In some species, seeds retained aboveground were dormant and thus capable of forming a persistent seed bank when they entered the soil.     

Effects of sand burial depth on seed germination and seedling emergence of Cirsium pitcheri

A greenhouse study was conducted to determine the effects of sand burial on seed germination and seedling emergence of Cirsium pitcheri, a threatened species along Lake Huron sand dunes. In October 1996, seeds of C. pitcheri were sorted into three groups (small, medium and large) and artificially buried at 2, 4, 6, 8, 10 and 12 cm depths in plastic pots filled with unsterilized sand from Lake Huron sand dunes. These pots were placed outside in sand boxes for natural stratification. In early February, the pots were brought into the greenhouse for germination of seeds. Data showed that percent seed germination and emergence of seedlings were not related to seed size. However, both variables were negatively correlated with depth. Seedling emergence occurred from a maximum depth of 6 cm with most seedlings emerging from 2 cm depth. Only one seedling emerged from 8 cm depth. Seedlings from large seeds produced longer roots than those of small seeds. Larger root system would probably enhance seedling establishment.     

Effects of sand burial depth on the root system of Salix cheilophila seedlings in Mu Us Sandy Land, Inner Mongolia, China

Salix cheilophila Schneid. is a naturally occurring Salix species in Mu Us Sandy Land, Inner Mongolia, China. We focused on the morphological adaptability of S. cheilophila to sand dune burial. For morphological measurements, 32 S. cheilophila seedlings were removed from a community which was in the process of being buried by a shifting sand dune. Each seedling collected included the entire root system. We measured the number, length, and biomass of the adventitious roots, primary lateral roots, and taproot, and compared the morphological characteristics of the root system, including adventitious roots, for seedlings buried to various levels in the sand. The growth range of adventitious roots increased as the length of the buried portion of the main shoot increased. In addition, the total dry weight of all current-year shoots tended to increase gradually with increasing total dry weight of the adventitious roots. These results suggest that S. cheilophila tends to make use of the sedimentary sand layer that accompanies shifting sand dunes. However, there was no correlation between biomass or number of adventitious roots and the length of the buried part of the main shoot. Thus, S. cheilophila does not grow adventitious roots proportional to the buried part. These morphological characteristics of the root system, including the adventitious roots, may indicate that S. cheilophila has poor morphological adaptability to sand dune burial.