Factors Influencing Seedling Establishment Rates in Zostera marina and Their Implications for Seagrass Restoration

Selection of strategies to efficiently utilize limited seed supplies in efforts to restore the seagrass Zostera marina (eelgrass) requires a better understanding of the processes that limit seedling establishment at potential restoration sites. We investigated the effect of seed distribution timing on seedling establishment and tested for interactive effects of seed burial and distribution timing. We also investigated the effect of habitat type on seedling establishment by distributing Z. marina seeds inside and outside of established Ruppia maritima (widgeongrass) patches and examined mechanisms causing habitat differences by manipulating seed position (buried or unburied) and vulnerability to seed predators (unprotected or protected in packets). Seeds distributed on the sediment surface in the summer (July or August) produced fewer seedlings than seeds distributed in fall (October) in five of six trials over 3 years. Seed burial increased success rates for seeds distributed in summer at one of two sites tested, eliminating the effect of season, but reduced success at the other site. Seeds placed in R. maritima generally produced fewer seedlings than seeds in bare sand, and although seed burial and protection in packets increased success in bare sand at three of four sites, the effect was less consistent in R. maritima. We conclude that seed predation and physical interactions were influential in reducing seedling establishment in R. maritima, contrary to hypotheses positing a nursery role for existing vegetation. Efficient restoration efforts with Z. marina seeds should target unvegetated areas after summertime sources of mortality have diminished. Direct seed burial may enhance seedling establishment rates.     

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.     

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.     

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.     

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.     

Factors influencing the natural regeneration of the pioneering shrub Calligonum mongolicum in sand dune stabilization plantations in arid deserts of northwest China

Calligonum mongolicum is a successful pioneer shrub to combat desertification, which is widely used for vegetation restoration in the desert regions of northwest China. In order to reveal the limitations to natural regeneration of C. mongolicum by asexual and sexual reproduction, following the process of sand dune stabilization, we assessed clonal shoots, seedling emergence, soil seed bank density, and soil physical characteristics in mobile and stabilized sand dunes. Controlled field and pot experiments were also conducted to assess germination and seedling emergence in different dune soil types and seed burial depths. The population density of mature C. mongolicum was significantly different after sand dune stabilization. Juvenile density of C. mongolicm was much lower in stabilized sand dunes than mobile sand dune. There was no significant difference in soil seed bank density at three soil depths between mobile and stabilized sand dunes, while the emergence of seedlings in stabilized dunes was much lower than emergence in mobile dunes. There was no clonal propagation found in stabilized dunes, and very few C. mongolicum seedlings were established on stabilized sand dunes. Soil clay and silt content, air‐filled porosity, and soil surface compaction were significantly changed from mobile sand dune to stabilized dunes. Seedling emergence of C. mongolicm was highly dependent on soil physical condition. These results indicated that changes in soil physical condition limited clonal propagation and seedling emergence of C. mongolicum in stabilized sand dunes. Seed bank density was not a limiting factor; however, poor seedling establishment limited C. mongolicum's further natural regeneration in stabilized sand dunes. Therefore, clonal propagation may be the most important mode for population expansion in mobile sand dunes. As a pioneer species C. mongolicum is well adapted to propagate in mobile sand dune conditions, it appears unlikely to survive naturally in stabilized sand dune plantations.     

The chemical nature of P accumulation in agricultural soils��implications for fertiliser management and design: an Australian perspective

Sand burial, persistent seed bank and soil water content (SWC) are three factors that potentially can affect regeneration in sand dune plant populations. To evaluate the effects of these three factors on population regeneration of Eremosparton songoricum, a rare and endangered legume, we investigated seed germination, seedling emergence and seedling survival in greenhouse and controlled field experiments in different sand dunes microsites. Freshly matured seeds are physically dormant, and the highest germination was only 9.3?±?5.8% at 25/10°C. Seed germination occurred at burial depths from 0 to 10 cm, but the maximum depth from which seedlings emerged was 6 cm; from 1 to 6 cm, the deeper the burial, the lower the percentage of seedling emergence. Only 36.2% of the total soil seed banks occurred at depths of 0?C6 cm. For artificially sown seeds at different dune microsites, mean seedling emergence percentage was 6.8%. Of 150 seedlings that emerged in the field investigation at the study site, only those germinating in flat sandy areas survived, and mean survival percentage was only 2.0%. Thus, the proportion of non-dormant seeds in soil seed banks that developed into seedlings and survived to the end of the growing season was only 0.2%. Minimum SWC for seed germination, seedling emergence and seedling survival was 2.0%. During monitoring of emergent seedlings in the field, low seedling recruitment was at least partly due to the rate of root extension (1.6?±?0.3 cm day^?1) into the sandy soil, which was slower than that of the downward movement of plant-available moisture (2.8?±?0.6 cm day^?1). Thus, population regeneration under natural conditions rarely occurred via sexual reproduction, owing to the limited water resources available for seedling establishment. Rational field seeding practices, including manually scarified or dry stored seeds before sowing, sowing the seeds at right time and suitable place, are suggested for ecological restoration of endangered E. songoricum populations.     

沙丘生态系统种子库研究现状、趋势与挑战

风蚀和沙埋赋予沙丘生态系统独特的自然物理和生物过程。沙丘生态系统种子库研究对于沙区植物多样性保护和生态恢复至关重要。但是人们对沙丘生态系统种子库的时空格局、形成机制和生态功能的认识还很肤浅。近些年, 沙丘生态系统种子库开始受到关注。该文对20年来沙丘生态系统种子库研究进行了回顾总结, 论述了沙丘生态系统种子库研究现状, 剖析了沙丘生态系统种子库现有结论, 阐述了沙丘生态系统种子库研究趋势与面临的挑战。沙丘生态系统种子库研究已获得以下认识: 1)沙丘生态系统类型、部位和深度决定种子库大小与组成; 2)沙丘生态系统种子库季节、年际变化受风沙活动和降水调控; 3)沙丘生态系统种子库受各种干扰的综合影响; 4)生境对沙丘生态系统种子库具有重要影响; 5)沙丘生态系统种子库结构和组成与植物繁殖对策密切关联; 6)种子库对沙丘生态系统植被恢复贡献率因沙丘类型而异。沙丘生态系统种子库研究应强化如下方面: 1)种子库区域分异规律研究; 2)种子库与植物生活史阶段的联系的研究; 3)多种干扰方式综合影响下的种子库研究; 4)种子库对沙丘生态系统植被恢复贡献的研究; 5)沙丘生态系统自然物理过程与种子库耦联关系的研究。     

Habitat degradation,topography and rainfall variability interact to determine seed distribution and recruitment in a sand dune grassland

Question: Understanding the mechanisms underlying how habitat degradation, topography and rainfall variability interactively affect seed distribution and seedling recruitment is crucial for explaining plant community patterns and dynamics. Interactions between these major factors were studied together in a semiarid sand dune grassland. Location: Eastern Inner Mongolia, China. Methods: The study system used four sites of fixed, semifixed, semishifting and shifting sand dune grasslands, representing a gradient of habitat degradation. We investigated the density of germinable seeds deposited in the top 5 cm of soil and in situ seedling emergence (number of seedlings emerging early in the growing season) and establishment (number of plants recruited at the end of the growing season) at three topographic positions (dune top, windward and leeward sides) within each site over 2 years that differed in rainfall. Habitat characteristics (i.e. vegetation cover, plant species composition and diversity, soil moisture and nutrient availability and soil erodibility) of the four sites were also measured. Results: Habitat degradation (i.e. decreased vegetation cover and enhanced wind erosion rate) significantly reduced the size of the germinable soil seed bank. On average, germinable seed number from the high‐vegetation cover fixed dune was 36‐fold larger than the low‐vegetation cover shifting dune, and eight‐ and two‐fold larger, respectively, than the semishifting and semifixed dunes with intermediate vegetation cover. We observed within‐habitat variability in seed distribution, but among‐topographic position variation differed among habitats. Seedling recruitment showed large between‐year, and among‐ and within‐habitat variability, but these variations varied significantly depending on the response variables evaluated (i.e. initial seedling density, final plant density, emergence rate and recruitment rate). Path analysis revealed complex density‐dependent positive and negative, direct and indirect effects of germinable seed density and initial seedling density on recruitment, but the relative importance of these density‐dependent effects varied depending on habitat type and rainfall availability. Conclusion: Our results suggest that habitat degradation, microtopography and rainfall availability interact in shaping sand dune seed bank and plant community recruitment patterns and dynamics. Their effects were mainly mediated through changes in both the biotic and abiotic environment during the process of habitat deterioration.     

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。     

Fine-scale spatial heterogeneity and seed size determine early seedling survival in a desert perennial shrub (<Emphasis Type="Italic">Ericameria nauseosa:</Emphasis> Asteraceae)

In heterogenous landscapes, seeds settle in some microsites more readily than others, independently of whether those microsites are suitable for germination and subsequent growth, and survival of seedlings. Wild seeds and seedlings of Ericameria nauseosa var. oreophila in the sand-dune ecosystem near Mono Lake, California show both concordance and conflict in where seeds are retained and where seedlings flourish. Using a field experiment, we followed performance of seeds and seedlings (i.e., the rate of germination and successful emergence, initial seedling size, seedling growth and survival over first growing season), by planting seeds of known sizes in pre-identified microsites (windward and lee aspects of interspaces between shrubs, under shrub canopies, and under snag canopies, respectively). Heavier seeds were more likely to germinate and emerge successfully and resulted in initially larger seedlings than lighter seeds. These initially larger seedlings subsequently remained larger over the growing season and lived longer than initially smaller seedlings. Independent of seed and initial seedling size, seedlings that germinated under the canopy of adult shrubs grew larger and survived longer than seedlings that germinated in interspaces (open space with little or no vegetation) or under snags (dead adult shrubs). Seedlings grown under the canopy of adult shrubs experienced significantly less solar radiation and wind-deposited sand than seedlings grown in interspace or snag microsites. Sand burial more than 1 cm was lethal for seedlings. The two variables of seed size and microsite type had by far greater impact on eventual growth and survival of seedlings than did aspect, and the effects of the former two variables were independent of each other. This study amplifies the body of work on E. nauseosa at this site demonstrating that the earliest events in the life history of this long-lived perennial shrub appear to persist through time despite the harsh and variable environment.     

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。     

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。     

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.     

The role of saltwort (Batis maritima L.) in regeneration of degraded mangrove forests

While saltwort (Batis maritima L.) is common in the fringe mangrove forests of southwest Florida, its role in regeneration of degraded mangrove communities is not known. Given the potential encroachment and subsequent degradation of mangrove communities by sea-level rise, it is important to quantify the effect of early-colonizing vegetation to early mangrove seedling survival. A greater number of mangrove seedlings were observed in existing B. maritima patches compared to surrounding mudflats. A planting experiment was designed to determine whether B. maritima was responsible for the observed pattern. Black mangrove (Avicennia germinans L.) seedlings, raised in a nursery, were planted in previously established B. maritima patches and on mudflats with and without nursery-raised B. maritima. There was significantly lower mortality of A. germinans seedlings when planted in existing B. maritima patches (69%), compared to seedlings planted on the mudflats (93%), demonstrating that existing B. maritima improved A. germinans seedling survival. Nursery-raised B. maritima had lower mortality on open mudflats (28%), suggesting that it can tolerate conditions, which make it an early colonizer of newly available habitats. The primary mechanism proposed for improving seedling success is a slight increase in elevation provided by the dense root network of established B. maritima. These findings have implications for scientists and managers anticipating the response of mangroves to sea-level rise.     

沙埋对两种沙生植物幼苗生长的影响及其生理响应差异

为了解沙埋对沙生植物生长的影响及其生理响应特征, 比较不同沙生植物耐沙埋能力及其机制, 2010年在内蒙古科尔沁沙地研究了不同深度沙埋下沙蓬(Agriophyllum squarrosum)和盐蒿(Artemisia halodendron)幼苗的存活率、株高等生长特性及其渗透调节物质含量, 保护酶活性和膜透性的变化, 得到以下结果: 沙蓬和盐蒿幼苗均具有较强的耐沙埋能力, 其中沙蓬幼苗最大耐沙埋深度超过幼苗10 cm, 盐蒿幼苗最大耐沙埋深度超过其株高8 cm; 随着沙埋深度增加, 沙蓬和盐蒿幼苗的存活率和株高均显著下降, 沙蓬的下降幅度显著小于盐蒿; 沙埋处理下两种植物均未表现出受水分胁迫, 沙埋导致其光合面积下降, 幼苗顶土困难, 是影响其存活和高生长的主要生态机制; 随着沙埋深度增加, 沙蓬幼苗丙二醛(MDA)含量显著增加, 盐蒿幼苗MDA含量下降, 虽然二者膜透性均呈增加趋势, 但沙蓬膜透性增加幅度显著低于盐蒿, 说明细胞膜受损是导致二者幼苗存活率下降和生长受到抑制的主要生理机制, 沙蓬膜透性受损程度较低是其耐沙埋能力较强的主要生理机制; 沙埋胁迫下, 虽然两种植物都通过提高过氧化物酶活性和脯氨酸含量减轻细胞膜受损程度, 但沙蓬体内超氧化物歧化酶也表现出重要协调作用, 使之酶促系统在保护细胞膜免受胁迫损伤过程中的作用更有效。     

Factors affecting the population dynamics of Suaeda maritima at initial stages of development

This study describes life history and population dynamics of the annual halophyte Suaeda maritima, at initial stages of seedling development, in salt-marshes of the Mont-Saint-Michel Bay. Effects of salinity on seed germination and seedling development were studied both in the laboratory and in the field. Salinity was measured as sediment electrical conductivity. Experiments in the laboratory showed that Suaeda maritima was relatively tolerant to salinity during the germination stage. In the field, germination occurred when soil salinity was minimal. Almost all seeds germinated in the field; this may explain the absence of a persistent seed bank. Salinity did not seem to affect the growth of seedlings either. Mortality in the field was attributed to physical factors such as anoxia during temporary immersion, burial by sediment and tidal uprooting. The intensity of these effects varied according to the location of Suaeda maritima seedlings: in hollows, along the edge of tidal creeks, in gaps among dense vegetation cover, or in pattern with Puccinellia maritima. Perennial vegetation did not restrict Suaeda maritima germination. In contrast, perennials such as Puccinellia maritima and Halimione portulacoides limited the development of Suaeda maritima seedlings since only a small number of this annual emerged beneath their canopies. The fate of seedlings depended to a great extent on the permeability of the perennial canopy to light. Thus, disturbance resulting in bare patches inside the perennial vegetation seemed essential for the development of this annual species. Suaeda maritima can grow in the presence of Puccinellia maritima so long as the latter present as an open matrix. Suaeda maritima may also benefit from protection against desiccation and tidal action where this occurs.     

Seed germination and seedling emergence of three annuals growing on desert sand dunes in China

BACKGROUND AND AIMS: Information on the initial growth characteristics of annuals found in Chinese deserts is very limited. The aim of this study was to investigate seed germination and interactive effects of irrigation and seed burial depth in sand on seedling emergence and seedling survival in three annuals (Agriophyllum squarrosum, Bassia dasyphylla and Aristida adscensionis) commonly growing on sand dunes in these regions. METHODS: Effects of temperature, light and polyethylene glycol-6000 on seed germination were examined by irrigating seeds sown on filter paper in Petri dishes. Seedling emergence was examined for seeds sown on the surface of, or at different depths (5, 10, 20, 30, 40 and 50 mm) in, sand-filled pots, which were irrigated under different regimes. For seeds buried at a depth of 50 mm, seed viability was examined after irrigation of the pots. KEY RESULTS: Seeds of three species germinated at most temperatures recorded between spring and autumn in their native habitats. No seed dormancy was found in any species. For all three species, seedling emergence was most favoured when seeds were buried at a depth of 10 mm. When seeds sown on the sand surface were irrigated, seed germination was considerably suppressed due to water deficiency, but many seeds remained viable. For A. squarrosum and B. dasyphylla, many seeds that were deeply buried and irrigated remained ungerminated but viable, while for A. adscensionis deeply buried seeds germinated, but the seedlings did not emerge due to unfavourable seedling growth in deep sand. CONCLUSIONS: Precipitation is the most crucial factor in determining the seasonal emergence of seedlings of the three tested species in the field. The vertical distribution of seeds in sand determines the proportion of seeds that germinate after precipitation and acts to maintain seed banks over multiple years.     

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。     

Seed mucilage improves seedling emergence of a sand desert shrub

The success of seedling establishment of desert plants is determined by seedling emergence response to an unpredictable precipitation regime. Sand burial is a crucial and frequent environmental stress that impacts seedling establishment on sand dunes. However, little is known about the ecological role of seed mucilage in seedling emergence in arid sandy environments. We hypothesized that seed mucilage enhances seedling emergence in a low precipitation regime and under conditions of sand burial. In a greenhouse experiment, two types of Artemisia sphaerocephala achenes (intact and demucilaged) were exposed to different combinations of burial depth (0, 5, 10, 20, 40 and 60 mm) and irrigation regimes (low, medium and high, which simulated the precipitation amount and frequency in May, June and July in the natural habitat, respectively). Seedling emergence increased with increasing irrigation. It was highest at 5 mm sand burial depth and ceased at burial depths greater than 20 mm in all irrigation regimes. Mucilage significantly enhanced seedling emergence at 0, 5 and 10 mm burial depths in low irrigation, at 0 and 5 mm burial depths in medium irrigation and at 0 and 10 mm burial depths in high irrigation. Seed mucilage also reduced seedling mortality at the shallow sand burial depths. Moreover, mucilage significantly affected seedling emergence time and quiescence and dormancy percentages. Our findings suggest that seed mucilage plays an ecologically important role in successful seedling establishment of A. sphaerocephala by improving seedling emergence and reducing seedling mortality in stressful habitats of the sandy desert environment.