种子生态学: 种子在群落中的作用

种子是指由胚珠发育而成的繁殖器官。在生态学研究中,其涵义要比植物学上种子的概念宽泛有时也指果实。作为种子植物生活史的一个重要阶段,种子从其发生、发育到成熟、传播、萌发以至成苗都与周围的环境紧密联系着。因而种子生态学是研究种子与其环境相互关系的科学。种子生态学的研究内容主要包括:种子发育与成熟的环境调节(生殖分配和结实格局)、种子性状、种子捕食、种子传播(包括种子雨)、种子库、种子休眠、种子萌发、幼苗建成和种群更新等,其中种子传播、土壤种子库、种子休眠与萌发是研究的核心内容。     

植物种群更新的补充限制

补充限制基于生态位理论, 从种子萌发、幼苗存活和生长、繁殖体扩散等生活史阶段的种群统计特征及环境因素(土壤水分、养分、凋落物等)着手, 探讨种群的更新问题。种源限制和微生境限制是补充限制理论研究的核心内容, 但是哪个更为重要并没有统一的结论。种源限制与种子生产、土壤种子库和地下芽库中的繁殖体数量不足有关。其中, 气候的年际波动、土壤种子库寿命和动物的捕食都会影响种子生产在种群更新中的作用; 土壤种子库常被视为种群更新的保险库, 与地上种子雨共同促进种群更新, 但是, 如果土壤里种子具有较高的死亡率和休眠率, 将会降低种子库的作用; 地下芽库及其产生的无性分株对于种群更新的意义更多地体现在干扰后种群更强的恢复能力上。扩散限制是种群更新中的普遍现象, 与种子产量、散布能力、传播媒介、幼苗密度等因素有关。微生境限制主要表现为水分、养分、凋落物等非生物因素以及竞争、捕食等生物因素对种子的活力、萌发性、幼苗的存活力、物质分配等过程的影响, 其重要性随着植物生活史阶段而发生变化。未来需要进行综合的、长期的实验, 并应着重加强种源限制及相关生态过程的进化与生态相结合的机理性研究, 从而更深刻地认识和理解种群更新问题, 建立更为综合、系统的种群更新理论体系。     

濒危乔木红豆杉不同更新阶段的生境特征比较

以福建梅花山自然保护区濒危乔木红豆杉(Taxus chinensis(Pilger)Rehd.)为对象,对红豆杉的种子排放、幼苗更新、幼树更新3个更新阶段的生境特征进行研究,并比较生态位的变化,以此评价红豆杉不同更新阶段的生境需求。结果显示:随着个体发育生长,红豆杉更新生境发生了明显转变,种子排放地与幼苗生境空间一致性较高,而它们与幼树生境存在明显的空间不一致性。从生态位角度来看,种子排放地和幼苗生境重叠指数≤0.5的仅有海拔和坡度2个因子,这说明幼苗的空间格局受到鸟类传播的强烈影响,鸟类传播所排放的种子能萌发并生长成幼苗。幼苗和幼树生境重叠指数≤0.5的因子有海拔、坡向、植被类型,说明在大尺度景观因子和庇护植被层面上,植物的更新需求出现了明显的不一致。研究结果表明红豆杉的更新生境存在明显的阶段变化,可能造成植物更新出现建成限制,使幼树阶段成为更新的瓶颈阶段。     

海南霸王岭陆均松种子雨和土壤种子库特征

为了解陆均松(Dacrydium pierrei)的自然更新能力,对海南霸王岭陆均松进行为期3年的种子雨、种子库及幼苗幼树观测,并分析其与环境因子的关系。结果表明,陆均松种子雨和种子库有效种子数量太少并存在季节性差异;其结实存在大小年现象;种子雨的扩散密度随着距母树距离的增加而逐渐减少;土壤种子库种子数量在枯枝落叶层所占比例最大,但有活力种子数量极少。幼苗幼树适宜生存的海拔范围为800~1 400 m,经度、纬度、坡度与幼苗幼树分布有明显相关性。因此,选择适宜季节采摘种子,提高种源数量和质量,清除林下枯枝落叶,增加种子萌发能力等以减少陆均松更新的限制条件的人工抚育措施应适当考虑。     

秦岭壳斗科四种植物种子雨组成及其动态变化

种子雨组成及其动态变化是植物种群自然更新的一个关键阶段。2011年和2012年8-12月,在秦岭南坡佛坪国家级自然保护区内对壳斗科的四种优势植物(板栗、锐齿栎、栓皮栎和小橡子)的种子雨组成及其动态变化进行了调查研究。结果表明:(1)壳斗科四种植物的种子雨均起始于8月中上旬,至11月中下旬结束,种子掉落高峰集中在9月下旬至10月中旬之间,其中板栗的各类型种子雨掉落高峰均要早于其它三种植物;(2)无论2011年还是2012年,锐齿栎、栓皮栎和小橡子的各种子雨类型在时间动态上均存在极显著差异,而板栗的各种子雨类型在时间动态上则差异并不显著;(3)四种植物的种子雨动态在年际间存在显著差异,2011年各类型种子雨掉落密度远远大于2012年(栓皮栎除外),推测2011年可能是种子产量的相对丰年。     

植物天然更新过程中种子和幼苗死亡的影响因素

植物天然更新包括有种子搬运、种子库动态、种子萌发和幼苗定居等过程。从种子生产到幼苗定居的更新是植物生活史中最为敏感的阶段之一 ,多种因素的影响种子和幼苗的命运。其中包括 :( 1 )动物取食或病原体侵袭。种子在扩散和搬运过程中 ,易被小哺乳动物或无脊椎动物取食。蛀虫也可以使种子失去萌芽能力。病原体感染种子和幼苗 ,容易引起种子和幼苗的死亡。 ( 2 )异质生境的影响。在不同生境中 ,光照条件、土壤水分和化学成分等因子的组合严重影响种子和幼苗的命运。 ( 3 )干扰的影响。小尺度和大尺度的干扰都可以影响到植物更新时种子和幼苗的命运。林窗作为特殊的干扰体系 ,为不同种类植物提供了更新的机会。 ( 4 )繁殖体特征。种子大小、质量和保护色等特征影响种子和幼苗在更新过程中的生存。种子休眠期间 ,由于生理衰老和腐烂的原因使种子失去活力而不能萌发。 ( 5 )密度和距离制约。母株附近由于密度竞争的影响 ,种子和幼苗死亡率都较高。     

植物种子大小与幼苗生长策略研究进展

种子大小和幼苗更新对策是植物生活史策略的重要组成部分.本文从不同侧面综述了当前国内外关于种子大小和幼苗之间关系的生态学研究,包括:种子大小对萌发出土的影响,种子大小与幼苗形态生长特性之间的关系,种子大小与幼苗存活、竞争能力之间关系等,并对今后的相关研究进行了展望.在不同的微环境和植被类型下,种子大小与幼苗生长之间的关系可能有所差异;种子大小对植物幼苗生长的影响导致种子大小不同的植物对植被幼苗更新补充的贡献不同;种子大小与幼苗生活史策略关系在大尺度的群落空间水平上和小尺度的物种间与物种内的研究对于天然植被的更新恢复研究具有重要意义.     

以种子为繁殖体的植物更新模型研揪

种子植物的自然更新过程是20世纪70年代以来生态学领域研究的热点问题之一。对影响种子植物更新过程因子的研究主要涉及3个阶段,即种子生产、种子运动(主要为种子扩散的过程)和种子萌发、幼苗建成以及幼树生长到成树。一些完整的理论和实验模型被提出。反过来模型的研究促进了野外研究的进一步发展。关于种子植物更新的定性模型和定量模型陆续被提出,定量研究种子植物的更新过程将成为主要的发展方向。本文综述了近年来关于种子植物更新过程的3个较为完整的定性模型,即Chambers等的种子命运模型、Clark等的补充限制模型和Nathan等的时空模型,以及2个定量模型,即Vander和Boehning等的种子扩散模型。     

千岛湖片段化生境中木本植物种子雨基本特征及其影响因子

为探究片段化生境中木本植物种子雨的基本特征,该研究根据2015—2020年(研究期间)在千岛湖样岛上的植物群落长期监测样地内每月收集的种子雨数据,采用Kruskal-Wallis检验对木本植物的种子雨密度进行年际差异分析,对不同传播方式物种的种子雨密度进行月份间差异性分析,并利用线性混合效应模型,探究岛屿空间特征(岛屿面积、距最近岛屿的距离、距大陆的距离)以及气候因子(0 ℃以上积温、降水量)对木本植物以及不同传播方式物种的种子雨密度的影响。结果表明:(1)2015—2020年6年间,在29个样岛用240个收集器共收集到877 178粒木本植物的成熟种子,属于26科40属52种。(2)动物传播是木本植物主要的种子传播方式,不同传播方式物种的种子雨时间动态存在较大差异。(3)木本植物的种子雨年密度与岛屿面积和年积温呈显著正相关,与年降水量呈显著负相关。(4)自主传播物种的种子雨月密度与距最近岛屿的距离呈显著正相关,而动物传播物种的种子雨月密度则与距大陆的距离呈显著正相关,风力传播物种的种子雨月密度与月积温呈极显著正相关。综上表明,生境片段化通过岛屿空间特征影响了木本植物种子雨的时间动态。     

准噶尔盆地东南缘梭梭种子雨特征

为了研究梭梭种子散布规律,通过布设种子雨收集器结合室内实验分析,对准噶尔盆地东南缘梭梭种子雨特征进行了研究。研究显示:(1)梭梭种子雨的累积密度达到平均189粒/m2,其中有活力种子占约80%;(2)种子散布的高峰集中在11月初到11月15日时间段,其落种量占整个种子雨的65%。其后种子雨密度随时间逐渐减小;(3)整个种子雨过程,不同时期散落的种子雨质量存在差异,表现为不同时期散落种子的萌发率呈现出先增大后减小的趋势;(4)变异函数分析表明,梭梭种子雨在8.12 m的有效变程内,种子雨具有明显的空间格局,其由空间自相关和随机因素引起的空间异质性各占50.0%。准噶尔盆地东南缘梭梭种子雨密度大且质量较高,同时其时空分布异质性较高,这些特征均将影响梭梭种群的分布格局和种群更新。     

Seed dynamics of resprouting shrubs in grassy woodlands: Seed rain,predators and seed loss constrain recruitment potential

Abstract Measuring the fate of seeds between seed production and seedling establishment is critical in understanding mechanisms of recruitment limitation of plants. We examined seed fates to better understand the recruitment dynamics of four resprouting shrubs from two families (Fabaceae and Epacridaceae) in temperate grassy woodlands. We tested whether: (i) pre‐dispersal seed predation affected seed rain; (ii) post‐dispersal seed predation limited seed bank accumulation; (iii) the size of the seed bank was related to seed size; and (iv) viable seeds accumulated in the soil after seed rain. There was a distinct difference in seed production per plant between plant families with the legumes producing significantly more seeds per individual than the epacrids. Seed viability ranged from 43% to 81% and all viable had seed or fruit coat dormancy broken by heat or scarification. Pre‐dispersal predation by Lepidopteran larvae removed a large proportion of seed from the legume seed rain but not the epacrids. Four species of ants (Notoncus ectatomoides, Pheidole sp., Rhytidoponera tasmaniensis and Iridomyrmex purpureus) were major post‐dispersal seed removers. Overall, a greater percentage of Hardenbergia (38%) and Pultenaea (59%) seeds were removed than the fleshy fruits of Lissanthe (14%) or Melichrus (0%). Seed bank sizes were small (<15 seeds m^?2) relative to the seed rain and no significant accumulation of seed in the soil was detected. Lack of accumulation was attributed to seed predation as seed decay was considered unlikely and no seed germination was observed in our study sites. Our study suggests that seed predation is a key factor contributing to seed‐limited recruitment in grassy woodland shrubs by reducing the number of seeds stored in the soil.     

Effect of Bat Exclusion on Patterns of Seed Rain in Tropical Rain Forest in French Guiana

To assess the impact of bats on seed dispersal in a tropical mature forest (Nouragues, French Guiana), we conducted a bat exclusion experiment and tested the hypotheses that an artificial reduction in the abundance of bats would result in: (1) a decrease in seed species diversity, at the community level; and (2) an increase in seed limitation (a failure of seeds to reach all suitable sites for germination) at the species level. Seed rain was sampled in two contiguous forest plots for a total of 120 d, using 49 seed traps (1 m²) arranged in 7 × 7 grids and spaced at 5-m intervals. Using mist nets, bat activity was reduced in one forest plot for a total of 60 nights. Thirty-nine plant species, or species groups, likely to be consumed and dispersed by bats, were collected within a total sample of 50,063 seeds. The overall seed rain was dominated by epiphytic Araceae and Cyclanthaceae (83.3%) and tree species within the genera Cecropia and Ficus (16.0%). Seeds from bat-dispersed shrubs and treelets ( Piper , Solanum , and Vismia ) were relatively rare (0.7%). The bat exclusion resulted in a 30.5 percent reduction in seed species richness and increased seed limitation for most of the species sampled. Seed limitation was caused mainly by a reduced seed rain (seed source limitation) rather than a decrease in seed dispersal uniformity (seed dispersal limitation). Therefore, bat-dispersed plants with low seed production are likely to be particularly affected by a decline in bat abundance, as a result of anthropogenic change.     

Reproductive potential and seedling establishment of the invasive alien tree Schinus molle (Anacardiaceae) in South Africa

Schinus molle (Peruvian pepper tree) was introduced to South Africa more than 150 years ago and was widely planted, mainly along roads. Only in the last two decades has the species become naturalized and invasive in some parts of its new range, notably in semi‐arid savannas. Research is being undertaken to predict its potential for further invasion in South Africa. We studied production, dispersal and predation of seeds, seed banks, and seedling establishment in relation to land uses at three sites, namely ungrazed savanna once used as a military training ground; a savanna grazed by native game; and an ungrazed mine dump. We found that seed production and seed rain density of S. molle varied greatly between study sites, but was high at all sites (384 864–1 233 690 seeds per tree per year; 3877–9477 seeds per square metre per year). We found seeds dispersed to distances of up to 320 m from female trees, and most seeds were deposited within 50 m of putative source trees. Annual seed rain density below canopies of Acacia tortillis, the dominant native tree at all sites, was significantly lower in grazed savanna. The quality of seed rain was much reduced by endophagous predators. Seed survival in the soil was low, with no survival recorded beyond 1 year. Propagule pressure to drive the rate of recruitment: densities of seedlings and sapling densities were higher in ungrazed savanna and the ungrazed mine dump than in grazed savanna, as reflected by large numbers of young individuals, but adult : seedling ratios did not differ between savanna sites. Frequent and abundant seed production, together with effective dispersal of viable S. molle seed by birds to suitable establishment sites below trees of other species to overcome predation effects, facilitates invasion. Disturbance enhances invasion, probably by reducing competition from native plants.     

Vertebrate‐mediated seed rain and artificial perches contribute to overcome seed dispersal limitation in a Mediterranean old field

Natural regeneration of vegetation is a frequent outcome of land abandonment, although the rate and diversity of such regeneration may be severely restricted by seed dispersal limitation, among other factors. In spite of this, studies aiming to quantify seed rain and test methods to enhance it, such as artificial perches, are still underrepresented in the Mediterranean. In our study, we quantified seed rain density and richness and tested the effects of artificial perches on such rain over a distance gradient on seven Mediterranean island old fields. In each of the seven sites, we positioned three sampling stations, each consisting of 1 seed trap under an artificial perch and 1 as a control on the ground, distributed at 30, 60, and 90 m from natural vegetation remnant. All traps received seeds, suggesting no overall dispersal limitation. Of the 11 seed species found, 10 were fleshy‐fruited and dispersed by vertebrates. Seed traps under perches received significantly higher seed rain of fleshy‐fruited species dispersed by birds, while ground traps received significantly more seeds of the species also dispersed by mammals, especially Rubus ulmifolius. The distance from the seed source was nonsignificant in all cases. Our study demonstrates the key role of vertebrate‐mediated seed dispersal services to overcome dispersal limitation in old fields, as well as the effective contribution of even small artificial perches in contrasting such limitation. The lack of differences over the distance gradient reveal that the upper spatial limit of dispersal limitation was not achieved.     

种子雨研究进展

种子雨是指在特定的时间和特定的空间从母株上散落的种子量。种子雨的组成和大小具有时空异质性。种子雨的空间异质性表现在种子雨的组成和大小因群落而异,种群间的种子雨因种群而异,种群内部的种子雨因个体而异;种子雨的时间异质性表现在不管是群落、种群还是种群内部的个体,其种子雨既具有季节动态,又具有年际变化。种子雨、种子库、幼苗库和地上植被相互联系、相互作用。种子雨的研究对更好地了解种群和群落动态等具有重要意义。应用现代的分子遗传标记技术、同位素标记法和荧光染料法等研究种子雨的散布过程和种子命运将是未来种子雨研究的热点,种子雨和种子库的结合研究及其与动植物关系的研究尚需加强。     

Seed Rain and Seed Limitation in a Planted Gallery Forest in Brazil

With seeds collected monthly during one year from 53 1‐m² seed traps, we investigated the seed rain and seed limitation in a gallery forest planted in 1994 in SE Brazil. Contrasting animal‐ (zoochorous) and wind‐dispersed (anemochorous) plants we investigated (1) which aspects of the composition and structure of the vegetation influence the abundance and species richness of the seed rain; (2) if such influences differ between zoochorous and anemochorous seeds; (3) if the abundance and richness of the seed rain sampled under zoochorous and nonzoochorous plant species differ; and (4) if seed limitation (given by the proportion of sites to which seeds were not dispersed) differs between zoochorous and anemochorous plant species, and also between species that have been planted and those that further colonized the area (colonists). Seed rain was intense and dominated by anemochorous species. The overall seed rain was not influenced by the vegetation parameters we analyzed (canopy height and cover, plant size, abundance, and richness) or by the plant species above the seed trap. The abundance and richness of zoochorous seeds in a given spot was influenced by the abundance and richness of zoochorous plants in its immediate vicinity. Seed limitation was higher for anemochorous than zoochorous species and higher for planted than for colonist species. We concluded with recommendations for the initial establishment of a planted forest, including the homogeneous distribution of zoochorous plants to permit a spatially homogeneous zoochorous seedfall, which will likely enhance the chances of survival and successful establishment of seeds.     

Dispersal and microsite limitation in Australian old fields

The recovery of native communities after cultivation may be constrained by (a) the failure of species to reach a site or (b) their failure to survive once there. Although seed addition is a common method to test for seed versus microsite limitation, most studies do not follow populations beyond seedling establishment, nor do they measure seed dispersal. We examined dispersal across native grassland/old field boundaries and investigated the relative importance of seed and microsite limitation across multiple life-history stages and generations. Seed trapping showed little movement of native seeds into old fields and that most species had extremely localized dispersal. Consequently, there was no pattern of seed density with distance from boundaries, and similarity between the seed rain and standing vegetation was moderate to high. Seed addition showed that two annual species were able to establish in all, and flower in most, subplots in the first year, and that seedling establishment increased with sowing density, consistent with seed limitation. However, the relative importance of microsite limitation increased over the lifespans of the species. Density dependence reduced the number of flowering plants, resulting in a large decline in seedling density in the following generation. This decline continued so that the initial positive effect of sowing density on seedling numbers disappeared by the fourth generation and hence the persistence of populations is uncertain. Thus, by monitoring seed dispersal and following experimental populations beyond seedling establishment, we showed that dispersal limits species distributions, but microsite plays an important role in limiting population growth and persistence.     

动物与植物种子更新的关系Ⅱ．动物对种子的捕食、扩散、贮藏及与幼苗建成的关系

植物的繁殖体总是面临来自各类生物（如昆虫、脊椎动物、真菌）的捕食风险。因动物捕食引起的种子死亡率影响植物的适合度、种群动态、群落结构和物种多样性的保持。种子被捕食的时间和强度成为植物生活史中发芽速度、地下种子库等特征的主要选择压力,而种子大小、生境类型等因素也影响动物对植物种子的捕食。捕食者饱和现象被认为是植物和种子捕食者之间的高度协同进化作用的结果,是限制动物破坏种子、提高被扩散种子存活率的一种选择压力。大部分群落中的大多数植物种子被动物扩散。种子扩散影响种子密度、种子被捕食率、病原体攻击率、种子与母树的距离、种子到达的生境类型以及建成的植株将与何种植物竞争,从而影响种子和幼苗的存活,最终影响母树及后代植物的适合度。种子被动物扩散后的分布一般遵循负指数分布曲线,大多数种子并没有扩散到离母树很远的地方。捕食风险、生境类型、植被盖度均影响动物对种子的扩散。植物结实的季节和果实损耗的过程也体现了其对扩散机会的适应。许多动物有贮藏植物种子的行为。动物贮藏植物繁殖体的行为,一方面调节食物的时空分布,提高了贮食动物在食物缺乏期的生存概率;另一方面也为种子萌发提供了适宜条件,促进了植物的扩散。于是,植物与贮食动物形成了一种协同进化关系,这种关系可能是自然界互惠关系（mutualism)的一种。影响幼苗存活和建成的因子包括种子贮蒇点的微生境、湿度、坡向、坡度、林冠盖度等。许多果食性动物吃掉果肉后,再将完好的种子反刍或排泄出来。种子经动物消化道处理后,发芽率常有所提高。     

Spatiotemporal patterns of seed dispersal in a wind-dispersed Mediterranean tree (Acer opalus subsp. granatense): implications for regeneration

Seed dispersal can severely limit the quantity of plant recruits and their spatial distribution. However, our understanding of the role of dispersal in regeneration dynamics is limited by the lack of knowledge of seed deposition patterns in space and time. In this paper, we analyse the spatiotemporal variability of seed dispersal patterns in the Mediterranean maple, Acer opalus subsp. granatense, by monitoring seed rain along two years at a broad spatial scale (2 mountain ranges, 2 populations per range, 4 microhabitats per population). We quantified seed limitation and its components (source and dispersal limitation), and explored dispersal limitation in space by analysing dispersal distances, seed aggregation, and microhabitat seed distribution. Acer opalus subsp. granatense was strongly seed‐limited throughout the gradients explored, being always dispersal limitation much higher than source limitation. The distribution of seeds with distance from adult individuals was leptokurtic and right‐skewed in all populations, being both kurtosis and skewness higher the year of the highest seed production. Dispersal distances were shorter than expected by random in the four populations, which suggests distance‐limited dispersal. Dispersal patterns were highly aggregated and showed a preferential direction around adults. At the microhabitat scale, most seeds accumulated under adult maples. However, there were no more seeds under trees and shrubs other than maple than in open interspaces, implying that established vegetation does not disrupt patterns of seed deposition by physically trapping seeds. When compared with patterns of seedling establishment, limited dispersal ability and inter‐annual spatial concordance in seed rain patterns suggest that several potentially safe sites for recruitment have a very low probability of receiving seeds in most maple populations. These findings are especially relevant for rare species such as Acer opalus subsp. granatense, and illustrate how dispersal studies are not only crucial for our understanding of plant population dynamics but also to provide conservation directions.