种子重量的生态学研究进展

作为植物生活史中的一个关键性特征,种子重量与其它许多植物性状和生态因子有关,种子重量的分异与其它一些植物性状及环境的变化关系在进化生物学上已经成为一个非常有意义的研究内容,且具有一定的实践意义。种子重量被发现与下列的一些植物学和群落学性状有关:植物的生活型、种子的散布能力、种子的散布方式、植物的高度、植物的冠幅、植物的比叶面积、植物的寿命、动物的捕食、植被中植物的数量或多度、土壤中种子的数量或多度、种子的休眠、种子在土壤中的持久性和植物的净初级生产力等,另外生态因子如降雨、温度、坡向、海拔、经度、纬度、光强和干扰等都影响种子的重量。种子的重量被认为是在大量小种子和少量大种子之间的进化折衷,在一定的能量限度内,较大重量的种子一般具有较少的数量,而较小重量的种子一般数量较多,这是种子重量和数量方面具有的一种反向关系。与其它性状相比,很多研究都表明种子重量和植物的生活型的关系密切。没有散布结构或风散布的种子比以动物和水作为散布媒介的种子重量要小。种子重量与捕食的关系现发现有3种格局。种子重量和形状与种子在土壤中的持久性的关系有4种格局。在干旱和阴暗的环境条件下,种子有变大的趋势。大重量种子比小种子赋予幼苗较优势的竞争地位,其原理尚有争论,尚不清楚是否是幼苗阶段的竞争决定了世界上大部分植被类型的物种组成。未来的研究方向主要有以下几个方面:1) 种子重量与植物系统学相结合,探索种子重量的变化规律;2)调查群落三向(纬度、经度和海拔)性的种子重量谱变化规律;3) 群落演替与群落种子重量谱的变化;4) 种子重量与群落中植物个体和种子的数量的关系及机理研究;5) 微生境、微地形如坡向、坡位和林间隙等对种子重量的影响;6) 全球气候变化和种子重量变化的关系。     

Seeds photoblastism and its relationship with some plant traits in 136 cacti taxa

Seed germination triggered by light exposure (positive photoblastism) has been determined in quantitative studies for numerous plant families and species. For Cactaceae, positive photoblastism is thought to be associated with life form and seed mass, but this association has never been evaluated. To explore hypotheses on associations between seed mass, seed dispersal, seed dormancy, life form, taxa and plant height with Relative Light Germination (RLG) in Cactaceae, we evaluated the effect of light on seed germination of 136 taxa. The taxa studied are native to several countries: México, Chile, Argentina, Brazil, Perú, USA, and Venezuela. Seed traits contrasted with RLG were life form, seed mass, seed dispersal, seed dormancy, adult plant height and taxon. We found some differences between RLG among taxa; Cacteae, Pachycereeae and Trichocereeae had higher RLG than Notocacteae. RLG was lower for seeds from taller than for shorter taxa, and lower for taxa with heavier seeds than for taxa with lighter seeds. Dispersal syndrome groups varied with RLG. RLG did not differ between cylindrical and globose taxa. Trends found here were in agreement with expectations for small-seeded species to have a light requirement to germinate more often than large-seeded species. This is the first time that cactus height is related to photoblastism. It is possible that seeds from tall plants are larger and thus have the capacity to produce taller seedlings than those from small plants, and that seedlings from large seeds with more resources have the ability to emerge from greater soil depths than those from small seeds.     

Seed size selection by olive baboons

Seed size is an important plant fitness trait that can influence several steps between fruiting and the establishment of a plant’s offspring. Seed size varies considerably within many plant species, yet the relevance of the trait for intra-specific fruit choice by primates has received little attention. Primates may select certain seed sizes within a species for a number of reasons, e.g. to decrease indigestible seed load or increase pulp intake per fruit. Olive baboons (Papio anubis, Cercopithecidae) are known to select seed size in unripe and mature pods of Parkia biglobosa (Mimosaceae) differentially, so that pods with small seeds, and an intermediate seed number, contribute most to dispersal by baboons. We tested whether olive baboons likewise select for smaller ripe seeds within each of nine additional fruit species whose fruit pulp baboons commonly consume, and for larger seeds in one species in which baboons feed on the seeds. Species differed in fruit type and seed number per fruit. For five of these species, baboons dispersed seeds that were significantly smaller than seeds extracted manually from randomly collected fresh fruits. In contrast, for three species, baboons swallowed seeds that were significantly longer and/or wider than seeds from fresh fruits. In two species, sizes of ingested seeds and seeds from fresh fruits did not differ significantly. Baboons frequently spat out seeds of Drypetes floribunda (Euphorbiaceae) but not those of other plant species having seeds of equal size. Oral processing of D. floribunda seeds depended on seed size: seeds that were spat out were significantly larger and swallowed seeds smaller, than seeds from randomly collected fresh fruits. We argue that seed size selection in baboons is influenced, among other traits, by the amount of pulp rewarded per fruit relative to seed load, which is likely to vary with fruit and seed shape.     

Why are small seeds dispersed through animal guts: large numbers or seed size per se?

Long-distance dispersal of seeds is an important process in metapopulation dynamics and in plant migrations, but at the same time extremely difficult to observe or quantify directly. If seed dispersal ability were related to attributes of seeds or motherplants, long-distance seed dispersal would be predictable by indirect approximation using easy-to-measure traits. Seed size has been suggested to be such a key trait in seed dispersal ability. However, having smaller seeds also implies having more numerous seeds per plant individual (given equal reproductive effort), and consequently increases the probability of seeds being ingested accidentally. The question is whether small-seeded species are more abundant in herbivore dung because smaller seed size increases survival rate during gut passage or because they are produced (and ingested) in greater numbers than larger seeds. We investigated endozoochorous seed dispersal via cattle grazing a meadow, and related seed abundance in dung samples to seed attributes. We found that seeds were ingested and passed through the bovine intestinal tract in proportion to the numbers produced per unit area in the grazed vegetation. In contrast, no relationship could be found between endozoochorous dispersal potential (measured as abundance of seeds in dung samples corrected for seed output in the grazed vegetation) and seed attributes such as seed mass, seed shape (roundness), and thickness of the seed coat. This finding underlines the importance of seed number in plant dispersal ability. In addition, it shows that grazing mammals may constitute an important dispersal vector for many plant species conventionally classified as 'unspecialised'.     

Resource distribution and the trade-off between seed number and seed weight: a comparison across crop species

In grain crops, total sink capacity is usually analysed in terms of two components, seed number and individual seed weight. Seed number and potential individual seed weight are established at a similar timing, around the flowering period, and seed weight at maturity is highly correlated with the potential established earlier. It is known that, within a species, available resources during the seed set period are distributed between both yield components, resulting in a trade-off between seed number and seed weight. Here we tested if this concept could apply for interspecific comparisons, where combinations of numbers and size across species could be related to the total available resources being either allocated to more seed or larger potential individual seed weight during the seed set period. Based on this, species differences in seed weight should be related to resource availability per seed around the period when seed number is determined. Resource availability per seed was estimated as the rate of increase in aboveground biomass per seed around the period of seed set. Data from 15 crop species differing in plant growth, seed number, seed weight and seed composition were analysed from available literature. Because species differed in seed composition, seed weight was analysed following an energy requirement approach. There was an interspecific trade-off relationship between seed number per unit of land area and seed weight (r = 0.92; F(1, 13) = 32.9; n = 15; P < 0.001). Seed weight of different species was positively correlated (r = 0.90; F(1, 13) = 52.9; n = 15; P < 0.001) with resource availability per seed around the seed set period. This correlation included contrasting species like quinoa (Chenopodium quinoa; ∼100000 seeds m⁻², ∼4 mg equivalent-glucose seed⁻¹) or peanut (Arachis hypogaea; ∼800 seeds m⁻², ∼1000 mg equivalent-glucose seed⁻¹). Seed number and individual seed weight combinations across species were related and could be explained considering resource availability when plants are adjusting their seed number to the growth environment and seeds are establishing their storage capacity. Available resources around the seed set period are proportionally allocated to produce either many small seeds or few larger seeds depending on the particular species.     

Seed size and shape and persistence in the soil in the New Zealand flora

Seed size and shape predict seed persistence in the soil for British and Argentinian herbaceous plant species. Those species with small, rounded seeds tend to have persistent seeds while those with larger, more elongate or flattened seeds usually lack persistence. It has been suggested that the mechanism underlying this pattern may be ease of burial, as small, rounded seeds are incorporated into the soil more easily than large, elongate or flattened seeds and are therefore less likely to be eaten by seed predators. We tested whether seed size and shape were related to persistence in the soil for 47 species native to New Zealand forests. There was a tendency for species with persistent seeds to have smaller seeds than species with transient seeds. However, species with large and/or elongate or flattened persistent seeds were relatively common. This indicates that seed size and shape are not related to persistence in New Zealand in the same way as in Britain and Argentina. A similar negative result has been found in Australia. The underlying cause of the patterns observed is unlikely to be ease of burial, since incorporation of seeds into the soil is likely to operate in all countries in a similar manner on seeds without specialised seed burial mechanisms. Data from all four floras studied to date also suggest that species with small, rounded seeds that do not germinate immediately must have the ability to survive periods of burial.     

Hamilton's rule, imprinting and parent-offspring conflict over seed mass in partially selfing plants

When genes in the offspring control the provisioning of the seed, the optimal seed size can be calculated exactly by applying Hamilton's rule. When seed size is a compromise between mother and offspring, we predict that outcrossing plant species produce larger seeds than selfers. This trend was found in the British flora and in a number of well-studied plant families. The analysis was extended to imprinting, a conditional strategy in which a gene in the offspring takes more resources when derived from the father than from the mother. The conditions for imprinting to be selected were rather restrictive. The analysis is relevant for the current debate about the evolution of imprinting in Arabidopsis thaliana.     

Rules of the seed size game: contests between large‐seeded and small‐seeded species

The coexistence of multiple seed size strategies within plant communities have been considered puzzling, based on a theoretical expectation of the existence of an optimal seed size under each set of specific environmental conditions. A model aimed at explaining the coexistence of different seed sizes has been suggested, where a seed size – seed number tradeoff is connected to a tradeoff between competition and colonization, leading to a competitive advantage in larger‐seeded species and a colonization advantage in smaller‐seeded species. Recently an alternative model has been suggested, based on a tradeoff between stress tolerance and fecundity, associated with the variation from large to small seeds. Here, we examine the role of seed size for recruitment in two‐species contests subjected to various treatments. In a garden experiment seeds of 14 plant species were combined pair‐wise into seven pairs, each with one larger‐seeded species and one smaller‐seeded species. Each species‐pair was sown with sparse and dense seed densities and subjected to different treatments of shading and litter. Recruitment was recorded during two years. Our results showed a general advantage of larger‐seeded species over smaller‐seeded species. This seed size advantage increased in treatments with litter, whereas there were minor effects of shade, and no effect of seed density was found. We thus found little support for a density dependent seed size game as assumed in models of a competition‐colonization tradeoff, whereas our results fit well with a model based on a tradeoff between stress tolerance and fecundity. Our experiment provides novel empirical data to theoretical models on co‐existence between multiple seed size strategies.     

Seed Size,Fecundity and Postfire Regeneration Strategy Are Interdependent in Hakea

Seed size is a key functional trait that affects plant fitness at the seedling stage and may vary greatly with species fruit size, growth form and fecundity. Using structural equation modelling (SEM) and correlated trait evolution analysis, we investigated the interaction network between seed size and fecundity, postfire regeneration strategy, fruit size, plant height and serotiny (on-plant seed storage) among 82 species of the woody shrub genus, Hakea, with a wide spectrum of seed sizes (2–500 mg). Seed size is negatively correlated with fecundity, while fire-killed species (nonsprouters) produce more seeds than resprouters though they are of similar size. Seed size is unrelated to plant height and level of serotiny while it scales allometrically with fruit size. A strong phylogenetic signal in seed size revealed phylogenetic constraints on seed size variation in Hakea. Our analyses suggest a causal relationship between seed size, fecundity and postfire regeneration strategy in Hakea. These results demonstrate that fruit size, fecundity and evolutionary history have had most control over seed size variation among Hakea species.     

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。     

Geographic variation in seed traits within and among forty‐two species of Rhododendron (Ericaceae) on the Tibetan plateau: relationships with altitude,habitat, plant height,and phylogeny

Seed mass and morphology are plant life history traits that influence seed dispersal ability, seeding establishment success, and population distribution pattern. Southeastern Tibet is a diversity center for Rhododendron species, which are distributed from a few hundred meters to 5500 m above sea level. We examined intra‐ and interspecific variation in seed mass and morphology in relation to altitude, habitat, plant height, and phylogeny. Seed mass decreased significantly with the increasing altitude and increased significantly with increasing plant height among populations of the same species. Seed mass differed significantly among species and subsections, but not among sections and subgenera. Seed length, width, surface area, and wing length were significantly negative correlated with altitude and significantly positive correlated with plant height. Further, these traits differed significantly among habitats and varied among species and subsection, but not among sections and subgenera. Species at low elevation had larger seeds with larger wings, and seeds became smaller and the wings of seeds tended to be smaller with the increasing altitude. Morphology of the seed varied from flat round to long cylindrical with increasing altitude. We suggest that seed mass and morphology have evolved as a result of both long‐term adaptation and constraints of the taxonomic group over their long evolutionary history.     

Seed enhancement: getting seeds restoration‐ready

Seed enhancement technologies such as seed priming and seed coating, developed by the agricultural seed industry, are standard procedures for the majority of crop and horticultural seeds. However, such technologies are only just being evaluated for native plant seeds despite the potential benefits of such treatments for improving restoration effectiveness. Key approaches applicable to native seed include: (1) seed priming, where seeds are hydrated under controlled conditions, and (2) seed coating, in which external materials and compounds are applied onto seeds through a diversity of treatments. These technologies are commonly employed to accelerate and synchronize germination and to improve seed vigor, seedling emergence, establishment, and to facilitate mechanized seed delivery to site, through standardizing seed size and shape. Seed enhancement technologies have now been tested on native seeds to overcome logistical and ecological barriers in restoration. However, further research is needed to extend the application of seed enhancements to a broader array of species, ecosystems, and regions as well as to evaluate new and innovative approaches such as the incorporation of beneficial soil microorganisms and plant growth regulators in the coatings. As techniques in native seed enhancement develop, these approaches need to be capable of being scaled‐up to provide the tonnages of seed required for global restoration.     

Relationships between intra‐specific variation in seed size and recruitment in four species in two contrasting habitats

Large seeds contain more stored resources, and seedlings germinating from large seeds generally cope better with environmental stresses such as shading, competition and thick litter layers, than seedlings germinating from small seeds. A pattern with small‐seeded species being associated with open habitats and large‐seeded species being associated with closed (shaded) habitats has been suggested and supported by comparative studies. However, few studies have assessed the intra‐specific relationship between seed size and recruitment, comparing plant communities differing in canopy cover. Here, seeds from four plant species commonly occurring in ecotones between open and closed habitats (Convallaria majalis, Frangula alnus, Prunus padus and Prunus spinosa) were weighed and sown individually (3200 seeds per species) in open and closed‐canopy sites, and seedling emergence and survival recorded over 3 years. Our results show a generally positive, albeit weak, relationship between seed size and recruitment. In only one of the species, C. majalis, was there an association between closed canopy habitat and a positive seed size effect on recruitment. We conclude that there is a weak selection gradient favouring larger seeds, but that this selection gradient is not clearly related to habitat.     

Growth-form regulates the altitudinal variation of interspecific seed mass of woody plants in Mt. Dalaoling,the Three Gorges Region,China

Aims Seed size is one of the most important characteristics of plant seeds, and has significant implications in plant ecological functions. Exploring the altitudinal pattern of seed size would help to detect environmental constraints on species distribution and understand the linkage between plant ecological function traits Methods The present study measured the quantitative features of seed size, including weight of 1 000 grain seeds, lengths of longer and shorter axes, and analyzed the relationships between seed size and altitude, as well as the influence of growth form. Seed samples were all collected from 201 locally common woody plants (belonging to 59 families and 87 genera) in the Dalaoling Natural Reserve in Yichang City, Hubei Province.Important findings Measured values of the seed mass, the longer axis, and the shorter axis of the 201 woody plant species all follow the lognormal distribution. Measurements of seed mass vary across five orders of magnitude. Significant correlation was found between seed mass, seed length of the longer and shorter axes (R² = 0.755; 0.819; 0.630, p < 0.01). Moreover, seed mass of trees and small trees are significantly heavier than those of shrubs and woody vines. Seed mass values of evergreen broad leaved species are significantly heavier than those of deciduous broad leaved species and needle leaved species. Seed mass of all 201 species shows a slightly but statistically significant decreasing trend with the increase of altitude. In addition, altitudinal patterns of seed mass varied between species with different growth form. Our results indicated the variation of altitudinal trends of seed mass for different structural components of plant communities, implying the local community structure as a critical aspect of variation in macro-ecological patterns.     

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.     

Co-evolution of seed size and seed predation

Using the evolutionarily stable strategy (ESS) approach in a model for the co-evolution of seed size and seed predation, I show that seed size variation within individual plants is favoured if there is a trade-off in the predator's attack rate for different seed sizes. A single seed size is not evolutionarily stable because a predator that is optimally adapted to one particular seed size cannot prevent invasion by plants with a different seed size. The model generates the following predictions. The ESS consists of a continuous range of seed sizes. Small seeds tend to be attacked more frequently than big seeds. Plants with many resources and plants with low (frequency-independent) juvenile mortality have more variable seeds than plants with few resources and a high juvenile mortality. Seed size variation is higher in fluctuating populations regulated by seed predation alone than in stable populations (partially) regulated by seedling competition. Predator searching behaviour does not directly affect the ESS seed size range, but may have an indirect effect by affecting population stability or the significance of seedling competition as a population regulating mechanism. Moreover, seed size distributions are found to be more skewed in favour of small seeds if predation is spatially non-uniform than if predation is more even. Application of the model to systems of several co-evolving plant and predator species is discussed.     

Frugivory and Seed Dispersal by the Lowland Tapir Tapirus terrestris in the Peruvian Amazon

The lowland tapir Tapirus terrestris is the largest herbivore in the Neotropics and feeds on a large quantity of fruits, often ingesting the seeds and defecating them intact. Seed dispersal by the lowland tapir in the southwestern Amazon was studied by examining seeds from 135 dung samples collected between 2005 and 2007. Seeds of a total of 122 plant species were identified, representing 68 genera and 33 families. The species accumulation curve showed that more species can be expected with further sampling. Many species (45%) were only encountered once, and only 10 percent of all species were found in >10 samples, indicating that the lowland tapir is an opportunistic forager. Seed diversity showed a clear seasonal pattern and was highly correlated with fruit availability. Seed diameter ranged from <1 to 25 mm with 81 percent <10 mm diam. The size distribution of seeds found in lowland tapir dung generally followed that of seeds found in the forest, but had a lower proportion of seeds in the smallest size class (<2.5 mm) and a larger proportion found in the largest size class (20–25 mm). The diversity of seeds encountered in dung of the lowland tapir in this study was much higher than in previous studies. We conclude that the lowland tapir is a potential disperser for a large number of plant species, including many that previously have been thought to be dispersed only by large primates.     

Seed weight increases with altitude in the Swiss Alps between related species but not among populations of individual species

Seed weight is a crucial plant life history trait, determining establishment success and dispersal ability. Especially in stressful environments, larger seeds may be selected at the expense of seed number, because larger seeds have a better chance of giving rise to an established offspring. We tested the hypotheses that between related species-pairs and among populations of single species a similar trend for increasing seed weight with increasing altitude should be present. Firstly, we measured seed weights from 29 species-pairs, with one species occurring in lowland areas and a congeneric species from high altitudes. Seeds of the alpine species were 28±8% larger than seeds from lowland species (P<0.01). Compared to the related lowland species, 55% of the alpine species had heavier seeds, 3% (one species) had lighter, and 41% had seeds of approximately equal weight. Secondly, we compared seed weights among populations of four species from different habitats and with different life histories. Seeds from between 11 and 34 populations per species were sampled along altitudinal gradients of 800–1,500 m (ca. 800 m in Scabiosa lucida, ca. 1,000 m in Saxifraga oppositifolia, ca. 1,000 m in Epilobium fleischeri, and ca. 1,500 m in Carex flacca). In all the four species, we found no indication for heavier seeds at higher altitudes. Our results indicate a selection pressure for species with heavier seeds at higher altitude, but the trend does not seem to operate across all cases. Phylogenetic constraints may limit the correlation among altitude and seed weight, operating particularly against selection for larger seed size, the closer populations and species are related to each other.     

Why is there variation in mean seed size among plants within single populations? test of the fertilization efficiency hypothesis

We tested the fertilization efficiency hypothesis, which attempts to explain mean seed size variation among plants within single populations, by comparing the patterns of seed size variation between chasmogamous (CH) flowers and cleistogamous (CL) flowers in Impatiens noli-tangere and Viola grypoceras, respectively. The fertilization efficiency hypothesis predicts that larger plants produce larger seeds if the number of pollen grains captured by a plant increases with increased allocation of resources to its attractive structures (e.g., corolla and nectar), but with diminishing gains. Thus, seed size should depend on plant size in seeds from CH flowers because of the diminishing gains of capturing pollen in these flowers, whereas seed size should not depend on plant size in seeds from CL flowers because CL flowers need not capture outcross pollen. We found significant positive correlations between mean seed size per plant and plant size for seeds from CH flowers in both species. However, there was no significant positive correlation between these two factors for seeds from CL flowers of both species. The results of the present investigations were thus consistent with the fertilization efficiency hypothesis.