杂草种子传播研究进展

种子传播将母株生殖周期的末端与它们后代种群的建立连结了起来,广泛认为,其对植被结构具有深刻的影响。种子传播的整个过程称为种子传播循环。研究表明,杂草种子传播的因子多种多样,包括仅依赖自身来完成的主动传播,以及依赖风、水、动物、人类等外界媒介的被动传播。其中,人类传播杂草种子是影响最广泛的一种,对现代植物的分布格局产生了深刻的影响。杂草种子的传播,对杂草种子库的数量和空间动态影响很大。研究种子传播的主要方法有荧光染料标记法、放射性同位素标记法、稳定同位素分析、分子遗传标记等。结合近几年国内外的研究进展,作者就杂草种子传播对种子库数量和空间动态影响的精确直接研究、杂草种子传播的过程及传播后的命运、杂草种子适应传播的机理、生态控草措施研究、外来杂草入侵蔓延与其种子传播的关系等方面提出了展望。     

Long-distance seed dispersal in plant populations

Long-distance seed dispersal influences many key aspects of the biology of plants, including spread of invasive species, metapopulation dynamics, and diversity and dynamics in plant communities. However, because long-distance seed dispersal is inherently hard to measure, there are few data sets that characterize the tails of seed dispersal curves. This paper is structured around two lines of argument. First, we argue that long-distance seed dispersal is of critical importance and, hence, that we must collect better data from the tails of seed dispersal curves. To make the case for the importance of long-distance seed dispersal, we review existing data and models of long-distance seed dispersal, focusing on situations in which seeds that travel long distances have a critical impact (colonization of islands, Holocene migrations, response to global change, metapopulation biology). Second, we argue that genetic methods provide a broadly applicable way to monitor long-distance seed dispersal; to place this argument in context, we review genetic estimates of plant migration rates. At present, several promising genetic approaches for estimating long-distance seed dispersal are under active development, including assignment methods, likelihood methods, genealogical methods, and genealogical/demographic methods. We close the paper by discussing important but as yet largely unexplored areas for future research.     

Splash seed dispersal by raindrops

Splash seed dispersal by raindrops was investigated for plants in southern Japan. Nine families, 10 genera and 19 species were confirmed as raindrop-dispersed plants. The 10 genera were Gentiana, Gratiola, Chrysosplenium, Mazus, Mitella, Ophiorrhiza, Sagina, Sedum, Trigonotis and Veronica. The method of splash rain dispersal in these species was clarified. Raindrop-dispersed species were all small herbaceous plants with a vertical pedicel and an apically opening fresh capsule when the seeds mature. Open capsules were cup-shaped or boat-shaped and can accommodate raindrops easily. The raindrops splashed the seeds from the capsule. In general, the seeds weighed very little, but they were heavier than powder or dust seeds dispersed by wind. A strong negative correlation was found between seed weight and the number of seeds per capsule. In the case of Trigonotis brevipes (Maxim.) Maxim., raindrops were received into the cup-shaped calyx-tube and dispersed the fruitlets. Some species, such as Gentiana thunbergii (G. Don) Griseb., Gentiana zollingeri Fawcett and Ophiorrhiza japonica Blume, had hydroscopic movement capsules that opened widely only when wet. Raindrop-dispersed plants were found in various habitats. For example, some plants grew together on rocks along the mountain torrents where splash water could easily be caught. The results of the laboratory and field experiments indicated that the dispersal distance of seeds by raindrops was 1m or less. For small herbaceous plants, splash dispersal by rain might be an effective and advantageous method of seed dispersal because dispersal is not affected by plant height.     

Contagious dispersal of seeds of synchronously fruiting species beneath invasive and native fleshy‐fruited trees

In subtropical Australia, many native and invasive plant species rely on a shared suite of frugivores, largely birds, for seed dispersal. Many native plants fruit during summer in this region, whereas most invasive plants fruit during winter, thus providing the opportunity for contagious dispersal of seeds beneath synchronously fruiting species. We sampled invasive and native seed rain beneath the canopy of a native summer‐fruiting tree Guioa semiglauca and an invasive winter‐fruiting tree Cinnamomum camphora, in three study sites over the course of a year. In July, during peak fruiting season for C. camphora and other invasive species, seed rain of invasive species was higher beneath C. camphora than G. semiglauca. This was partly due to the invasive tree Ligustrum lucidum, whose seed rain was three times higher beneath C. camphora than beneath the native tree. In February, seed rain of native species was more abundant beneath the canopy of G. semiglauca than beneath C. camphora, despite the fact that C. camphora was also fruiting at this time. This was probably due to the larger fruit crop produced by G. semiglauca at this time of year. Our study provides evidence that the presence of invasive bird‐dispersed plants may facilitate contagious seed dispersal of other invaders, and likewise native species may facilitate seed spread of other native plants.     

Overseas seed dispersal by migratory birds

Long-distance dispersal (LDD) promotes the colonization of isolated and remote habitats, and thus it has been proposed as a mechanism for explaining the distributions of many species. Birds are key LDD vectors for many sessile organisms such as plants, yet LDD beyond local and regional scales has never been directly observed nor quantified. By sampling birds caught while in migratory flight by GPS-tracked wild falcons, we show that migratory birds transport seeds over hundreds of kilometres and mediate dispersal from mainland to oceanic islands. Up to 1.2% of birds that reached a small island of the Canary Archipelago (Alegranza) during their migration from Europe to Sub-Saharan Africa carried seeds in their guts. The billions of birds making seasonal migrations each year may then transport millions of seeds. None of the plant species transported by the birds occurs in Alegranza and most do not occur on nearby Canary Islands, providing a direct example of the importance of environmental filters in hampering successful colonization by immigrant species. The constant propagule pressure generated by these LDD events might, nevertheless, explain the colonization of some islands. Hence, migratory birds can mediate rapid range expansion or shifts of many plant taxa and determine their distribution.     

The experimental study of seed dispersal by fish (ichthyochory)

1. The last few years have seen an increased interest in the experimental study of seed dispersal by fish (ichthyochory). This paper reviews such experiments, aiming to determine what functional aspects of ichthyochory have been investigated, what experimental designs have been used and what the potential pitfalls are. 2. The process of seed dispersal by fish can be divided into six discrete stages, each with its own probability of occurrence and each individually quantifiable in seed feeding trials: (i) seed uptake, (ii) ingestion, (iii) retention time, (iv) survival, (v) germination probability and (vi) germination rate after gut passage. 3. Inter‐ and intraspecific variation in seed traits (e.g. size, coat hardness, coat morphology, colour, presence and chemical composition of fruit pulp) and characteristics of fish (e.g. gape width, jaw morphology, presence of teeth, length of the digestive tract and digestive capability) can significantly affect the probability of one or more of the six stages of ichthyochory, thereby affecting the probability and distance of seed dispersal by fish. 4. To date only seven studies, which together investigated a total of nine fish species and 25 plant species, have used feeding experiments to study one or more of these quantifiable stages in the ichthyochory process. There is a clear bias in the research questions towards assessing seed survival during passage through the gut and subsequent viability. Only a few studies focus on seed retention in the digestive tract and germination rate, and even fewer address seed ingestion. 5. There is also considerable variation in experimental design among studies: Some have used groups of fish, while others used fish that are individually housed; some have fed seeds to hungry fish, while others used sated fish; some studied germination of seeds dissected from the alimentary tract, rather than seeds recovered from the faeces. 6. I present a number of recommendations for a more standardised protocol for future experimental studies of zoochory in general, and ichthyochory in particular, and highlight areas of interest for future research.     

Primate seed dispersal: Coevolution and conservation implications

Early studies of primates have demonstrated that many species rely heavily on fruit, and that primates constitute a large component of the frugivore biomass in tropical forests. Consequently, primates have long been thought to be important seed dispersers. It is only recently that studies have been conducted that have illustrated the complex nature of the interactions between fruit-eating primates and their food trees. Such studies have raised questions as to the causes and conse-quences of the intriguing differences between primate communities, the importance of other animals in the interactions (such as dung beetles and rodents that secon-darily disperse seeds), how primate-plant interactions evolve, and the significance of primates in forest regeneration and conservation. Since subsistence and com-mercial hunting of primates has heavily impacted frugivore communities, but left the physical structure of the forest relatively unaltered, studies of primate seed dispersal have important implications for the future of forests where seed dispersers have declined or disappeared.     

鼠类扩散种子的几种标签标记法的比较

以四川省都江堰市亚热带常绿阔叶林内的优势鼠种——小泡巨鼠为实验动物,通过半自然状态围栏控制实验,检验了几种种子标签法对鼠类扩散植物种子的影响。结果表明:鱼线和细钢丝两种标记线在跟踪扩散种子命运上都是很有效的,但考虑到鱼线有时会被实验鼠咬断,细钢丝是更值得推广应用的一种标记线。3种标记牌(大塑料牌、小塑料牌和金属牌)在跟踪扩散种子命运上没有显著差异,但由于较大的尺寸和较强的可见度,大塑料牌更适合作为标签应用于野外种子扩散标记。     

Adapting mechanized vascular plant seed dispersal technologies to biocrust moss restoration

Biocrust restoration is an emerging field relevant to management of rangelands. Manual dispersal of biocrust is an effective approach, though there are few examples of biocrust restoration greater than a square meter, in part because specialized machinery has yet to be developed or adapted for dispersal across larger areas. Restoration with vascular plants is now conducted using a variety of equipment for work at small to large scales. We investigated the potential of two tractor draft implements, a rangeland seed drill and imprinter, to establish moss. We assessed three treatments: moss passed through a seed drill, manually broadcasted over imprinted soil, and manually broadcasted onto unaltered soil. We conducted these treatments at two sites that differed in management histories and vegetative cover. To use the seed drill to disperse moss we needed to amend moss materials to prevent jamming, which reduced application rate. Imprinted treatments established the most cover, a result we attribute to creation of favorable microsites for establishment. Broadcast methods established a comparable number of moss colonies to broadcast‐over‐imprinted soil, but less cover. Drill treatments were not effective by any metric, likely due to burial of moss. Final differences between our highest cover treatment and controls were positive, but small: 2.38% in imprinted versus 0.44% in controls. However, moss cover in imprinted areas increased during our two‐year experiment at this site, while controls did not. Future work should focus on integrating biocrust and vascular plant treatments and identifying equipment to efficiently achieve these goals.     

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。     

Non-redundancy in seed dispersal and germination by native and introduced frugivorous birds: implications of invasive bird impact on native plant communities

Seed dispersal by vertebrate animals is important for the establishment of many fleshy-fruited plant species. Different frugivorous species can provide different seed dispersal services according to their specific dietary preferences as well as behaviour and body traits (e.g. body size and beak size of birds). Our aim was to study redundancies and complementarities in seed dispersal and germination between the two main native seed disperser birds and the introduced silver pheasant Lophura nycthemera in the temperate Patagonian forests. For this, we collected fresh droppings from the studied species and analyzed seed content. We conducted germination trials for four plant species common in bird droppings; two native species (Aristotelia chilensis and Rhaphithamnus spinosus) and two invasive non-native species (Rubus ulmifolius and Rosa rubiginosa). Both native frugivorous birds and the silver pheasant dispersed fruits of non- native fleshy-fruited plants, but their roles were non-redundant in terms of species dispersed and effect on seed germination. The silver pheasant dispersed a proportionally high number of non-native seeds, while native birds dispersed a high number of native seeds. In addition, the effect of gut treatment in seed germination differed between seed dispersers. Native birds promoted the germination for the two native plant species studied, while the silver pheasant promoted the germination of one non-native plant. This suggests that seed dispersal by the silver pheasant may contribute to the spread of some invasive fleshy-fruited plants in the ecosystems that otherwise would not be dispersed by any other bird. The understanding of redundancies and complementarities on seed dispersal and germination between native and introduced birds will allow improving the management of fleshy-fruited non-native plants.     

种子特征和结实量对啮齿动物取食和扩散种子行为的影响

为了深入了解啮齿动物在不同种子丰富度条件下对不同大小和单宁含量种子的觅食行为策略及其与植物种群更新的关系,在宁夏六盘山区的华北落叶松人工林,研究了不同大小和单宁含量[0%Tannin(T)、2%T、8%T和15%T]的人工种子在模拟结实小年和结实大年对啮齿动物取食和扩散行为的影响.结果表明: 啮齿动物消耗种子速度在结实小年更快,结实大年的种子消耗速度相对缓慢. 种子就地取食率(ISPR)在不同结实年份间无显著差异,扩散后取食率(PRAD)在结实小年显著高于结实大年,但前者的扩散后贮藏率(HRAD)显著低于后者;种子扩散后的取食距离(PDAD)和贮藏距离(HDAD)在结实小年均显著大于结实大年.在结实小年,大种子的PDAD和HDAD均大于小种子,前者在不同大小种子间均差异显著,而后者仅在2%T和15%T的不同大小种子间差异显著;在结实大年,除0%T外的其他单宁含量种子的PDAD和HDAD在不同大小种子间均差异显著.ISPR在中等单宁含量种子最大,高单宁含量种子最小;PRAD分别在结实小年的高单宁含量种子和结实大年的无单宁种子最大;不论在结实大年还是结实小年,HRAD均在高单宁含量种子最大,中等单宁含量种子最小.这说明结实大年可延缓啮齿动物对种子的消耗速率,提高种子的HRAD,但种子扩散距离减小;啮齿动物在结实大年和小年均表现出对大种子的扩散偏好,且大种子被扩散的距离更远;啮齿动物在不同结实年份均偏好于就地取食中等单宁含量种子,而扩散高单宁含量种子.     

Pollen and seed dispersal among dispersed plants

The ecological significance of spacing among plants in contributing to the maintenance of species richness, particularly in tropical forests, has received considerable attention that has largely focussed on distance- and density-dependent seed and seedling mortality. More recently it has become apparent that plant spacing is also relevant to pollination, which often constrains seed production. While seed and seedling survival is reduced at high conspecific densities, pollination success, by contrast, is positively correlated to local conspecific density. Distance-dependent mechanisms acting on pollination and seed production have now been described for a variety of plants, with relatively isolated plants or fragmented populations generally suffering reduced fecundity due to pollen limitation. Yet there is considerable variability in the vulnerability of plant species to pollination failure, which may be a function of breeding system, life history, the pollination vector, the degree of specialisation among plants and their pollinators, and other indirect effects of habitat change acting on plants or pollinators. As reduced tree densities and population fragmentation are common outcomes of anthropogenically altered landscapes, understanding how pollination processes are affected in such degraded landscapes can inform effective conservation and management of remaining natural areas.     

Flow regulation affects temporal patterns of riverine plant seed dispersal: potential implications for plant recruitment

1. Changes to the natural flow regime of a river caused by flow regulation may affect waterborne seed dispersal (hydrochory), and this may be an important mechanism by which regulation affects riverine plant communities. We assessed the effect of altered timing of seasonal flow peaks on hydrochory and considered the potential implications for plant recruitment. 2. We sampled hydrochory within five lowland rivers of temperate Australia, three of which are regulated by large dams. These dams are operated to store winter and spring rains and release water in summer and autumn for agriculture. At three sites on each river, hydrochory was sampled monthly for 12 months using passive drift nets. The contents of the drift samples were determined using the seedling‐emergence method. 3. More than 33 000 seedlings from 142 taxa germinated from the samples. In general, more seeds and taxa were observed in the drift at higher flows. By altering the period of peak flows from winter–spring to summer–autumn, flow regulation similarly affected the period of peak seed dispersal. The effect of regulation on seed dispersal varied between taxa depending on their timing of seed release and whether or not they maintain a persistent soil seed bank. 4. Hydrochory in rivers is a product of flow regime and the life history of plants. By altering natural flow regimes and thus hydrochorous dispersal patterns, flow regulation is likely to affect adversely the recruitment of native plant species with dispersal phenologies adapted to natural flow regimes (such as many riparian trees and shrubs) and encourage the spread of non‐native (exotic) species. 5. Changes to hydrochorous dispersal patterns are an important mechanism by which altered flow timing affects riverine plant communities. Natural seasonal flow peaks (in this case spring) are likely to be important for the recruitment of many native riparian woody taxa.