地中海沿岸沙丘土壤种子库的异质性

应用自然萌发法和物理筛选法相结合的方法,调查了地中海沿岸沙丘三种微生境土壤总种子库和永久种子库特征,用功能群的方法分析了微生境之间的土壤总种子库和永久土壤种子库的组成和结构差异,也分析了主要物种土壤种子库的微生境分布格局,分析了种子大小和土壤种子库中的种子永久性关系.发现地中海沿岸沙丘土壤种子库具有明显的空间异质性,即使在相同的微生境内,土壤种子库的分布也不是均匀的,甚至具有很大的差异;微生境对总的土壤种子库和各功能群的土壤种子库的分布格局具有显著的影响,开阔地区域具有最大的种子密度,而灌丛下和路径具有较小的种子密度;在永久土壤种子库中,豆科类植物种子在休眠种子中占有最大的比例,约为76%,而一年生和多年生禾草未发现休眠种子,豆类、伞形科类、多年生阔草、菊科类、一年生阔叶草和十字花科类等种子在土壤中的永久性比例依次下降,分别为50%、46%、41%、6%、6%和1%,微生境影响各功能群种子在土壤中的永久性.微生境显著影响大部分主要物种的土壤种子库分布;种子大小与其在土壤中的永久性为正相关,即大种子具有较高的永久性,小种子具有较低的永久性.     

地中海沿岸沙丘土壤种子库的异质性

应用自然萌发法和物理筛选法相结合的方法,调查了地中海沿岸沙丘三种微生境土壤总种子库和永久种子库特征,用功能群的方法分析了微生境之间的土壤总种子库和永久土壤种子库的组成和结构差异,也分析了主要物种土壤种子库的微生境分布格局,分析了种子大小和土壤种子库中的种子永久性关系。发现地中海沿岸沙丘土壤种子库具有明显的空间异质性,即使在相同的微生境内,土壤种子库的分布也不是均匀的,甚至具有很大的差异;微生境对总的土壤种子库和各功能群的土壤种子库的分布格局具有显著的影响,开阔地区域具有最大的种子密度,而灌丛下和路径具有较小的种子密度;在永久土壤种子库中,豆科类植物种子在休眠种子中占有最大的比例,约为76％,而一年生和多年生禾草未发现休眠种子,豆类、伞形科类、多年生阔草、菊科类、一年生阔叶草和十字花科类等种子在土壤中的永久性比例依次下降,分别为50％、46％、41％、6％、6％和1％,微生境影响各功能群种子在土壤中的永久性。微生境显著影响大部分主要物种的土壤种子库分布;种子大小与其在土壤中的永久性为正相关,即大种子具有较高的永久性,小种子具有较低的永久性。     

持久土壤种子库研究综述

土壤种子库是指存在于土壤上层凋落物和土壤中全部存活种子,简单地可分为短暂土壤种子库和持久土壤种子库。即使给予理想的萌发条件如季节、温度、湿度等,土壤种子库中也仍有部分种子保持休眠状态,休眠的种子组成了持久土壤种子库。持久土壤种子库具有在承受了空间或时间上不可预测的干扰的植被中发挥繁殖能力的潜势,因此,其在植被承受干扰后的恢复、管理和重建中常常起关键作用。研究持久土壤种子库能丰富生物多样性的内容,同时,可以为深入了解植被更新提供更多的信息。本文从持久土壤种子库的研究方法、分类方法、指示因子、生态意义,以及持久土壤种子库研究中存在的问题和今后的工作进行综述,试图为将来的工作奠定基础。     

土壤种子库的研究进展及若干研究热点

 土壤种子库是指存在于表层土壤（包括凋落物）中的有生命的种子。土壤种子库的研究已是植物生态学研究不可缺少的一部分,现已成为植物种群生态学中比较活跃的领域。土壤种子库时期是植物种群生活史的一个重要阶段,有人称之为潜种群阶段。土壤种子库对一年生植物来说尤其重要。土壤种子库简单地可分为瞬时土壤种子库和长久土壤种子库,即使给予理想的萌发条件如季节、温度、湿度等,土壤种子库中也仍有部分种子保持休眠状态,休眠的种子组成了土壤长久种子库的成分。时空异质性是土壤种子库的基本特性之一,不仅不同植被类型的土壤种子库具有不同的组成、大小和多样性,而且微环境也影响土壤种子库的分布格局。由于萌发、捕食和衰老等原因,土壤种子库具有季节动态,一般在旧种子萌发之后,新种子散布之前达到最低点。在高等植物占据的大多数生境中,以休眠繁殖体形式存在的个体远远超过地上植株的数量;土壤种子库、幼苗库和成年植被相互联系相互影响。由于各种原因如群落类型的差异、群落的演替阶段、取样的时间等,地上植被和土壤种子库之间关系大体上可分为两种情况,即相似性和差异性;研究土壤种子库的方法通常有萌发法和物理分离法。土壤种子库能部分反应群落的历史,对退化生态系统的恢复起着重要的作用。目前土壤种子库的主要研究热点问题可分为以下几个方面：1）土壤种子库的研究方法,2）土壤种子库的分类问题,3）土壤种子库分布的时空格局,4）地上植被和土壤种子库的关系,5）土壤种子库的动态等。     

荒漠草原区4种植物群落土壤种子库特征及其土壤理化性质

土壤种子库作为地上植被更新的潜在种源,在植被自然恢复和演替过程以及生态系统建设中起着重要作用。以宁夏盐池县荒漠草原区4种植物群落为研究对象,通过对封育16年后植物群落土壤理化性质的变化与其土壤种子库的特征之间的关系的探讨,揭示荒漠草原植物群落的土壤种子库分布特征和演替趋势,以及土壤质量的改变如何影响土壤种子库特征。研究结果表明：（1）不同植物群落土壤种子库物种组成及种子密度不同,同一物种在不同植物群落土壤种子库中出现时种子密度也存在差异;禾本科、菊科、藜科植物种在4种植物群落土壤种子库中出现比例均较高,分别占到26.19%、21.43%和19.05%,占总物种数的66.7%;灰绿藜、冰草、碱蓬在4种植物群落土壤种子库中均有出现。（2）4种植物群落土壤种子库中多年生植物和一年生植物种子居多,灌木、半灌木植物种较少。（3）4种植物群落土壤种子库中物种数和种子密度均表现为：芨芨草群落 > 苦豆子群落 > 油蒿群落 > 盐爪爪群落;且随着土层的加深,4种植物群落土壤种子库中物种数和种子密度均呈递减趋势。（4）4种植物群落土壤种子库中Shannon-Wiener指数、Simpsin指数、Pielou指数和Patrick指数均表现为芨芨草群落 > 苦豆子群落 > 油蒿群落 > 盐爪爪群落。此外,芨芨草群落土壤种子库与油蒿群落土壤种子库相似性最高,油蒿群落与盐爪爪群落的最低。（5）土壤种子库物种多样性特征指数与土壤pH、含水量、碱解氮呈正相关,与土壤电导率呈负相关,其中土壤pH和电导率对种子库物种多样性的影响较大。荒漠草原封育以后,土壤种子库中植物种数、种子密度和物种多样性均呈增加趋势,且以多年生植物和一年生植物种子居多;禾本科和豆科植物的物种数和种子密度大于菊科和藜科,这表明围封后牧草品质改善,植物群落正向演替。     

锡林郭勒退化草原不同禁牧恢复演替阶段土壤种子库比较

以冷蒿(Artemisia frigida) 糙隐子草(Cleistogenes squarrosa)退化草原经过长期围栏禁牧恢复演替(23 a和10 a)群落为研究对象,并对比围栏外退化的冷蒿 糙隐子草草原,揭示锡林郭勒退化草原长期围栏禁牧后的土壤种子库特征.随着群落的恢复演替,土壤总种子库物种组成和密度均增加,但持久性土壤种子库占土壤总种子库的比例下降.代表土壤总种子库的4月份取样,围栏禁牧23 a、10 a和围栏外退化样地土壤种子库密度平均值分别为4433、4756粒· m-2和856粒· m-2, 在0.001水平上差异显著,基本代表持久土壤种子库(persistent soil seed bank)的6月底取样,3个样地土壤种子库密度在0.05水平上差异显著;多数植物土壤种子库主要分布在0～3 cm层,各样地种子库密度随土壤深度的增加而减少;围栏禁牧23 a、10 a和围栏外重度退化样地的持久土壤种子库密度分别为土壤总种子库的52.6%、47.4%和90.9%,;各样地一年生植物,构成了草原土壤种子库密度的主要组成部分.结果表明,对于严重退化的冷蒿 糙隐子草群落,只要在恢复起点仍有零星分布的羊草和大针茅植株,经过长期围栏禁牧可恢复其土壤种子库物种组成和种子库规模.     

植冠与土壤种子库储存种子的萌发特性及策略

为明确黄土丘陵沟壑区植物种子库如何调控种子萌发来提高个体适合度,选择研究区7种具有种子库的主要物种为研究对象,以刚成熟和室内储存种子为对照,比较植冠宿存(5个宿存期)和土壤埋藏(5a埋藏期)对植物种子萌发特性的影响,探讨植冠种子库与土壤种子库储存下的种子萌发策略。结果表明:7种植物种子经过不同种子库储存后萌发特性表现出明显的种间差异,黄刺玫(Rosa xanthina)和水栒子(Cotoneaster multiflorus)种子萌发力表现为植冠宿存不变型、土壤储存增强型,土壤储存明显提高水栒子种子萌发速率;达乌里胡枝子(Lespedeza davurica)和狼牙刺(Sophora davidii)种子萌发力表现为植冠宿存增强型、土壤储存减弱型,种子萌发历时表现为植冠宿存延长型,土壤种子库储存还可加快达乌里胡枝子萌发速率、缩短萌发历时;茭蒿(Artemisia giralaii)和铁杆蒿(Artemisia gmelinii),种子萌发率随植冠宿存时间先升高后降低,随土壤储存时间先降低后升高,土壤储存可推迟其萌发,铁杆蒿种子萌发速率在植冠与土壤储存后均加快;紫丁香(Syringa oblata)种子萌发率随植冠宿存先升高后降低,土壤储存明显加快其种子启动萌发与速率。在黄土丘陵沟壑区,植物种子经过植冠或土壤种子库储存,或增加、加快、提早萌发充分利用有利条件提高占据性,或减少、减缓、推迟萌发分摊不利条件的风险;而且该区植物植冠与土壤储存后种子萌发特性间的关系,体现各自适应环境与应对干扰的分工与协作策略,主要表现为:单一主导型和相辅相成型。     

安太堡露天煤矿排土场土壤种子库

采用种子萌发法研究了安太堡露天矿排土场的土壤种子库,共统计到17种植物种子,分属5科,以1年生草本植物种子为主.排土场土壤种子库具有极大的时空异质性,地上植被与土壤种子库群落组成上的相似性程度较低,复垦后期土壤种子库间相似性系数较低.随着复垦时间、复垦模式或立地条件的不同,土壤种子库的种类、密度各不相同.选择正确的复垦模式,有利于土壤种子库自我更新能力的增加和排土场生态系统稳定,对排土场的土地复垦和生态重建具有重要意义.     

黔西北喀斯特土壤种子库季节动态及种子库对策

采用野外样线取样与温室萌发相结合的方法,研究了贵州省西北部喀斯特区域处于不同演替阶段群落的土壤种子库季节动态与种子库对策.实验共检测出130个物种,种子库密度变动于541～11180粒·m-2,草本种子的密度和物种数远大于木本种子.采样时间和地点对土壤种子库平均物种数和密度有显著影响.各群落类型的土壤种子库平均物种数都在4月达到最高,除草地外在12月下降到最低,草地在8月下降到最低.各群落类型土壤种子库密度都在4月达到最高,8月、12月以及次年5月的密度差异不显著.至少两种种子仅存在于1年中某些时段的短暂性土壤种子库类型和两种种子在全年的土壤种子库中都存在的持久性土壤种子库类型.植物的种子库对策是物种本身固有的生活史对策,但某些物种的种子库对策可能受干扰历史及微环境因素的影响,使其在种子数量上表现出不同的季节动态.     

准噶尔盆地梭梭土壤种子库基本特征

土壤种子库是种子的安全岛,是干旱荒漠区植被更新与恢复的基础。以梭梭为主的荒漠植被是准噶尔盆地绿洲区重要的生态屏障,但是,对于与梭梭种群自然更新紧密联系的土壤种子库特征的研究却相对缺乏。本研究通过野外采样、物理筛分并结合室内实验分析,对准噶尔盆地梭梭土壤种子库基本特征进行了研究。结果表明：（1）梭梭种群平均土壤种子库密度从71~696 粒·m^-2,局部小环境甚至达到7 534 粒·m^-2;（2）变异系数分析表明,各种群土壤种子库密度变异系数均大于1,说明梭梭土壤种子库的数量分布是非常不均匀的;（3）梭梭种群土壤种子库中种子萌发率从2.08%到47.62%不等,平均约18%;（4）变异函数分析表明,梭梭土壤种子库中种子分布的空间变异较大。综上,准噶尔盆地梭梭土壤种子库密度大且种子萌发能力强,同时其时空分布差异明显,这些特征均将影响梭梭种群的分布格局和种群更新。     

埋藏条件下3种干旱荒漠植物的种子休眠释放和土壤种子库

对种子休眠的自然释放及其作用因素的研究, 是了解种子休眠生态学、种群适应机制的重要途径。以内蒙古阿拉善干旱荒漠区的3种主要植物牛枝子(Lespedeza potaninii)、唐古特白刺(Nitraria tangutorum)和骆驼蒿(Peganum nigellastrum)为材料, 研究了种子在野外埋藏18个月期间和4个埋深条件下的休眠释放特性和土壤种子库。3种植物种子在野外埋藏时(采收后5 ℃冷藏6个月)的休眠率分别为98%、95%和3%。结果显示, 埋藏过程中, 3种植物种子的休眠释放表现出不同的变化特性。对牛枝子而言, 置于地表(0 cm)的种子比埋藏于土中的种子的休眠释放快, 埋藏期末, 埋深0、2、5和10 cm的种子的休眠率分别为64%、87%、86%和82%。唐古特白刺种子埋藏6个月后, 各埋深的休眠已完全释放, 释放速率随埋深增加而加快。骆驼蒿种子具有典型的季节性休眠循环特性, 休眠率各年度最高点出现在10月份, 释放速率随埋深增加呈减慢趋势。埋藏期末不同埋深条件下, 牛枝子、唐古特白刺和骆驼蒿种子的平均田间萌发率分别为11%、12%和8%; 平均室内萌发率分别为3%、74%和42%; 而平均死种子率分别为3%、15%和10%。根据Thompson和Grime (1979)的土壤种子库分类体系, 供试的3种植物都属于持久土壤种子库类型。     

Ecological correlations between the persistence of the soil seed bank and several plant traits,including seed dormancy

It was empirically showed that seed size and life history correlate with the formation of a soil seed bank. Although no empirical data are available that indicate a close relationship between seed dormancy and the soil seed bank, dormancy has been considered essential to the formation of a soil seed bank. I have considered the formation of the soil seed bank and survival of seeds for more than a year in the soil, and the persistence and survival of the seed bank for more than 5 years. These periods were derived from the definition of a persistent seed bank and the criterion for seed banks of long-term persistence. Plant traits that are closely related to the formation or persistence of a seed bank and their relationships to dormancy were analysed using two pre-existing databases of seed longevity in soil and comparative ecology. The integrated database comprised 18 plant traits and seed bank formation or persistence data. This approach was used to identify more reliable general empirical rules. The results of a regression tree analysis and common statistical tests of plant traits indicated that only life history and seed size were closely related to seed bank formation, and dormancy was not essential for the formation and persistence of a seed bank. However, the contribution of dormancy differed slightly between dormancy types. Scarification or dry storage requirements to break dormancy slightly enhanced the formation and persistence of a seed bank, whereas a chilling requirement decreased the formation and persistence of a seed bank. In contrast, fluctuating temperature requirements clearly contributed to the formation and persistence of a seed bank.     

土壤种子库研究综述

土壤种子库是指存在于土壤表面和土壤中全部存活种子的总和。土壤种子库研究是植物生态学研究的一个重要领域。研究土壤种子库是对生物多样性研究的一个重要补充 ,有助于对植被更新和植被演替动态的了解 ,对于植被重建与恢复具有指导意义 ,有利于指导我们的生产实践。本文从土壤种子库研究的取样方法、研究内容及进展等方面进行一个较系统的总结 ,为将来的研究提供一些有益的途径。     

Assessing soil seed bank persistence in flood‐meadows: The search for reliable traits

Abstract. To assess seed bank persistence of target species in endangered flood‐meadows (alliances Cnidion and Molinion), we investigated established vegetation and soil seed bank of 46 plots for 3 yr and 2 yr, respectively. As traits of seed persistence we calculated various continuous indices that refer to the frequency and abundance of species in above‐ground vegetation and at different soil depths. Furthermore, we tested the significance and soundness of easily observed traits such as maximum seed density per plot and seed attributes (mass, size and shape) as predictors of soil seed bank features. In linear regression, SAI, the seed accumulation index, showed the best agreement (R²= 0.64) with the seed longevity index that was derived from the database by Thompson et al. (1997) for a set of 115 species. The second best predictor (R²= 0.39) of the seed longevity index was maximum seed density per plot in the lower soil layer (5–10 cm). Depth distribution indices and seed attributes showed weaker but still significant relations. The dynamic character of flood‐meadows was reflected by a large proportion of species with a strong tendency to accumulate seeds in the soil relative to their importance in above‐ground vegetation. Most of these species have a ruderal strategy, exploiting gaps after flood disturbances, while the dominants of flood‐meadows tended to have short‐lived seed banks. Compared to other grassland types, a relatively large proportion of rare and endangered target species can be expected to form long‐term persistent seed banks. However, only under marginal conditions that facilitate seed survival in the soil (e.g. fallow) are these persistent seed banks likely to contribute to restoration. We conclude that easily observed traits of persistence such as seed weight, size and shape do not meet the accuracy needed in scientific and practical applications. Thus, there is a crucial demand for further seed bank studies in poorly investigated habitats and of rare species.     

Dormancy cycling and persistence of seeds in soil of a cold desert halophyte shrub

Background and Aims

Formation of seed banks and dormancy cycling are well known in annual species, but not in woody species. In this study it was hypothesized that the long-lived halophytic cold desert shrub Kalidium gracile has a seed bank and dormancy cycling, which help restrict germination to a favourable time for seedling survival.

Methods

Fresh seeds were buried in November 2009 and exhumed and tested for germination monthly from May 2010 to December 2011 over a range of temperatures and salinities. Germination recovery and viability were determined after exposure to salinity and water stress. Seedling emergence and dynamics of the soil seed bank were investigated in the field.

Key Results

Seeds of K. gracile had a soil seed bank of 7030 seeds m⁻² at the beginning of the growing season. About 72 % of the seeds were depleted from the soil seed bank during a growing season, and only 1·4 % of them gave rise to seedlings that germinated early enough to reach a stage of growth at which they could survive to overwinter. About 28 % of the seeds became part of a persistent soil seed bank. Buried seeds exhibited an annual non-dormancy/conditional dormancy (ND/CD) cycle, and germination varied in sensitivity to salinity during the cycle. Dormancy cycling is coordinated with seasonal environmental conditions in such a way that the seeds germinate in summer, when there is sufficient precipitation for seedling establishment.

Conclusions

Kalidium gracile has three life history traits that help ensure persistence at a site: a polycarpic perennial life cycle, a persistent seed bank and dormancy cycling. The annual ND/CD cycle in seeds of K. gracile contributes to seedling establishment of this species in the unpredictable desert environment and to maintenance of a persistent soil seed bank. This is the first report of a seed dormancy cycle in a cold desert shrub.     

Patterns of seed persistence in South African fynbos

In fire-prone communities such as fynbos, many species rely on regeneration from seed banks in the soil. Persistent seed banks are particularly important for species with life spans shorter than the average fire cycle, in order to counter local extinction. Persistent seed banks also give potential for restoring ecosystems following disturbances such as alien plant invasion. This study investigated the seed persistence patterns of 25 perennial species, representing several growth forms and life histories, during a three-year burial. Long-term persistence (i.e., seed bank half-life exceeding two years) was found in the hard-seeded Fabaceae and Pelargonium, and the nut-fruited Proteaceae. In this group, germinability was low and dormancy increased further following burial, resulting in a highly viable, dormant seed bank after three-year's burial. A second group with potentially long-term persistent seeds includes four taxa (Pseudopentameris, Passerina, Elegia and Restio) that either have low germinability or develop secondary dormancy following burial. Dormancy in the latter group was partially countered by exposure to smoke-seed primer. Of the small-seeded species, only two Erica species with high initial dormancy had long-term persistent seed banks. The other species mostly displayed high initial germinability and short-term persistent seed banks (i.e., seed bank half-life less than two years). This group included taxa with short to medium life-spans (Syncarpha, Roella) that were expected to have long-term persistent seeds in order to buffer against local extinction following average to long fire-return intervals. We hypothesize that light may play a role in overcoming secondary dormancy in those species, and could have resulted in an underestimate for seed persistence in this study. Alternatively, those short to medium life-span species persist via inter-fire recruitment in gaps or long-distance dispersal (of the smallest seed). No correlations were found between seed persistence and seed mass or variance in seed dimensions. Nor was a correlation found between seed persistence and phenol concentration. In fynbos, seed burial of larger seeds by ants and rodents are major processes that operate in conjunction with passive burial of small seeds. Selection for persistence can be expected to operate across all seed sizes and shapes in fire-prone communities.     

Climate change and bet-hedging: interactions between increased soil temperatures and seed bank persistence

In order to predict the long-term consequences of climate change, it is necessary to link future environmental changes to mechanisms that control plant population processes. This information can then be incorporated into strategies to more accurately model climate change impacts on species or to estimate future extinction risks. We examined the impact of increased temperatures on the longevity and dynamics of the persistent soil seed banks of eight ephemeral species from arid Australia. We found that the predicted global temperature increases under climate change will be reflected in increased soil temperatures, and that seeds in the soil seed bank will be exposed to long durations of high temperatures over the summer months. Three of the eight species studied had significantly greater levels of germination after exposure to predicted increased soil temperatures. Another species displayed a dramatic decrease in seed viability after such exposure. The capacity of such species to use the seed bank to bet hedge against rainfall events that cause germination but are insufficient to allow plant maturation, is compromised by increased germinability and subsequent loss or reduction of seed bank persistence. These predicted changes in the dynamics of soil seed banks increase the risk of local extinctions of these species, while the composition of the community may be altered by changes in species abundance. Our results show that the risk spreading mechanism provided by persistent seed banks could be compromised by the mechanistic impact of forecast temperature increases in arid habitats, and highlight the need to understand mechanisms that control population dynamics when attempting to address likely future impacts of climate change on biodiversity.     

Ecological longevity of Polaskia chende (Cactaceae) seeds in the soil seed bank,seedling emergence and survival

• Soil seed banks are essential elements of plant population dynamics, enabling species to maintain genetic variability, withstand periods of adversity and persist over time, including for cactus species. However knowledge of the soil seed bank in cacti is scanty. In this study, over a 5‐year period we studied the seed bank dynamics, seedling emergence and nurse plant facilitation of Polaskia chende, an endemic columnar cactus of central Mexico.
• P. chende seeds were collected for a wild population in Puebla, Mexico. Freshly collected seeds were sown at 25 °C and 12‐h photoperiod under white light, far‐red light and darkness. The collected seeds were divided in two lots, the first was stored in the laboratory and the second was use to bury seeds in open areas and beneath a shrub canopy. Seeds were exhumed periodically over 5 years. At the same time seeds were sown in open areas and beneath shrub canopies; seedling emergence and survival were recorded over different periods of time for 5 years.
• The species forms long‐term persistent soil seed banks. The timing of seedling emergence via germination in the field was regulated by interaction between light, temperature and soil moisture. Seeds entered secondary dormancy at specific times according to the expression of environmental factors, demonstrating irregular dormancy cycling.
• Seedling survival of P. chende was improved under Acacia constricta nurse plants. Finally, plant facilitation affected the soil seed bank dynamics as it promoted the formation of a soil seed bank, but not its persistence.