Effect of seed passage through vertebrate frugivores'' guts on germination: a review

The capacity of seeds to germinate after ingestion by frugivores is important for the population dynamics of some plant species and significant for the evolution of plant-frugivore interactions. In this paper the effects of different vertebrates on seed germination of nearly 200 plant species are reviewed, searching for patterns that predict the circumstances in which germination of seeds is enhanced, inhibited, or unaffected by the passage through the digestive tract of a seed disperser. It was found that seed dispersers commonly have an effect on the germinability of seeds, or on the rate of germination, or both, in about 50% of the plants they consume, although the diversity of animal species tested so far is still rather low (42 bird species, 28 non-flying mammals, 10–15 bats, 12 reptiles, 2 fishes). Enhancement of germination occurred about twice as often as inhibition.

In spite of the morphological and physiological differences in their digestive tracts, the different animal groups tested have similar effects on seed germination, although non-flying mammals tend to influence germination slightly more often than the other groups. Data on fishes are still too scarce for any generalization. Seed retention time in the dispersers' digestive tract is one factor affecting germination, and helps to explain the variation in seed responses observed among plant species, and even within a species. However other factors are also important; for example, the type of food ingested along with the fruits may affect germination through its influence on chemical or mechanical abrasion of the seed coat. Seed traits such as coat structure or thickness may themselves be responsible for some of the variation in seed retention times. Seeds of different sizes, which usually have different transit times through frugivores, and seeds of either fleshy or dry fruits, show often similar germination response to gut passage.

Seeds of different plants species differ strongly in their germination response after ingestion, even by the same frugivore species. Congeneric plants often show little consistency in their response. Even within a species variation is found which can be related to factors such as the environmental conditions under which germination takes place, seed morphology, seed age, and the season when the seeds are produced.

The effect of gut passage on germination differs between tropical and temperate zones. Seed germination of both shrubs and trees (data on herbaceous species are still scarce) in the temperate zone is more frequently enhanced than in the tropics. This result supports the hypothesis that enhanced germination may be more advantageous in unpredictable or less constant environments. Significant differences in frugivore-mediated germination are also found among different life forms. In both tropical and temperate zones, trees appear to be consistently more affected than shrubs or herbs. This might be due to an overall higher thickness of the seed coats, or to a higher frequency of seed-coat dormancy in tree species.

The influence of frugivory upon the population dynamics of a species has to be evaluated relative to other factors that influence germination and seedling recruitment at a particular site. Whether seed ingestion by dispersers is really advantageous to a plant (as has commonly been assumed) can only be assessed if we also determine the fate of the ingested seeds under natural conditions, and compare it to the fate of seeds that have not been ingested.     

Effects of rabbit gut passage on seed retrieval and germination of three shrub species

It has been known for a few decades that European rabbits consume seeds and fleshy fruits of native woody plants, but relevant factors in the endozoochory processes such as seed predation (chewing and digestion), sexual differentiation, or the temporal pattern of seed recovery have been little evaluated until now. In this study, we examined seed dispersal of three Mediterranean shrub species by wild rabbits through monitoring of seed retrieval and germination after gut passage. Twelve adult wild rabbits (Oryctolagus cuniculus; six males and six females) of similar size and age were fed seeds of three shrub species with fleshy fruits (Crataegus monogyna, Myrtus communis and Pistacia lentiscus). After ingestion of fruits, seeds were retrieved from dung every 12 h for a day and a half. The viability and germination of retrieved seeds were tested along with that of uneaten seeds. Between 5% and 76% of ingested seeds were retrieved from dung, with significant differences between species and sex. Most M. communis seeds were retrieved with 12–24 h after ingestion; almost all C. monogyna seeds were recovered with 0–12 h after ingestion; no seeds of P. lentiscus were recovered. Only in the case of M. communis seed was the recovery rate greater in female than in male. Passage through the rabbit gut significantly increased seed germination in M. communis, and decreased it in C. monogyna. In conclusion, wild rabbits acted in this study as predators of C. monogyna and P. lentiscus seeds and potential dispersers of M. communis seeds.     

植物化感物质对种子萌发的影响

所有的化感物质都对一些植物的种子萌发产生影响.本文介绍了植物化感作用和化感物质的定义,化感物质的释放方式和种类,然后重点综述了化感物质影响种子萌发的作用机理、内外因素和生态学意义.化感物质通过影响细胞膜透性、细胞分裂生长和分化、呼吸作用、蛋白质合成、基因表达、激素合成和平衡,最终影响种子萌发.化感物质对种子萌发的影响与化感物质的种类、浓度、受体植物种类和环境条件等有关.     

镉对水稻种子萌发的影响

以不同类型的319个水稻品种为研究对象,研究了10 mg·L^-1 Cd²⁺处理对水稻种子萌发的影响.结果表明:Cd²⁺对种子发芽率影响较小,对根系生长的影响显著,且大于对芽生长的影响;不同类型水稻种子萌发对Cd的响应差异较大,敏感顺序为粳稻>籼稻>杂交稻;两系不育系根系长度和根系数量受Cd的抑制程度显著高于三系.应用快速聚类方法,可以将参试品种划分为耐受型、中间型和敏感型3种不同的敏感类型.     

盐胁迫对二色补血草种子萌发的影响

以二色补血草(Limonium bicolor)种子为试验材料,用不同浓度的NaCl、Na2SO4及二者质量比为1:1的混合盐在其种子萌发过程中进行胁迫,调查盐胁迫对种子萌发的影响,并测定萌发过程中种子蛋白质含量、淀粉酶活力及还原糖和总糖含量的变化.结果表明:盐胁迫使二色补血草种子的发芽势、发芽指数和发芽率明显降低,而且随着盐浓度的增鲻加,抑制作用增强;3种盐对种子萌发的抑制作用强度依次为NaCl>混合盐>Na2SO4;二色补血草种子耐盐适宜范围为Na2SO4 1.2%、混合盐0.8%、NaCl 0.5%,耐盐半致死浓度为Na2SO4 2.1%、混合盐1.4%、NaCl 1.1%,耐盐极限浓度为Na2SO4 3.5%、混合盐2.4%、NaCl 2.0%.盐处理后不萌发的种子转入蒸馏水后重新萌发的试验结果表明,盐分胁迫主要是通过渗透效应,抑制种子中蛋白质和总糖的分解,进而延缓或抑制了种子的萌发.     

草食动物对豆科植物种子萌发的消化道作用效应的Meta分析

本文收集了近年来70种豆科植物种子形态学和消化道作用后萌发行为数据,利用植物亲缘系谱和整合分析,以阐明豆科植物种子经过草食动物消化道作用后萌发行为的变化。结果表明,豆科植物种子经过消化后,萌发率受到抑制(OR=0.89),但抑制效应和促进效应差异不显著(OR95%=0.61~1.28,P0.05);以亲缘关系为依据将70种豆科植物分为7个类群,其中消化作用对金合欢属、含羞草属、银合欢属、牧豆树属和角豆树属的第1类群种子萌发有显著的促进效应;而对包括木豆属、毛蔓豆属、爪哇大豆属、扁豆属、胡枝子属和距瓣豆属的第4类群有显著的抑制效应。将第3、5类群再划分成3个亚组进一步分析,结果表明,消化作用对苜蓿属种子萌发有显著促进,笔花豆属则受到显著抑制,三叶草属种子的抑制和促进效应之间差异不显著。车轴草族25种植物种子表现出质量中等种子经消化后萌发率显著提高,而过小或过大的种子经消化道作用后萌发率受到明显抑制。其他45种植物的种子重量与消化后的萌发效应存在不显著的负相关关系(R2=0.035,P0.05)。     

不同处理对沙生针茅种子萌发的影响

沙生针茅是重要的荒漠植物,研究其种子萌发特性及其对主要影响因子的响应对于认识其生存适应性以及荒漠植被恢复具有重要意义。鉴于沙生针茅种子常处休眠状态、不易萌发,而现有研究尚未充分揭示各种因素和处理对沙生针茅种子萌发的影响,本论文报道了不同因素、不同处理对沙生针茅种子萌发的影响。结果表明:(1)60℃蒸馏水浸泡种子24 h,种子吸水率、发芽率、发芽势分别达到26%、38%和32%;(2)-20/20℃变温处理96 h,发芽率、发芽势分别达到48%和46%;(3)切除绝大部分胚乳(3/4)后,种子发芽率、发芽势分别可达到58%和54%;(4)用80 mg·L~(-1)赤霉素(GA3)水溶液浸泡种子24 h,发芽率、发芽势分别达到54%和48%;(5)用聚乙二醇(PEG-6000)处理,随浓度增加,种子发芽率、发芽势、发芽指数均呈下降趋势,种子萌发水分胁迫的临界值和极限值分别为15.9%和26.3%;(6)盐胁迫处理下,种子发芽率、发芽势、发芽指数均随盐碱浓度增加呈下降趋势,其中Na2CO3处理下的种子萌发率下降幅度明显大于Na Cl,表明碱性盐胁迫对沙生针茅种子萌发的抑制作用大于中性盐胁迫;(7)采用综合处理方法,种子发芽率、发芽势分别达到78%和76%。     

Effects of drought stress on seed germination of ornamental sunflowers

The response to drought stress on germination was investigated on three hybrids of ornamental sunflower, ‘Hadar’, ‘Pazit’, and ‘Zohar’. Different levels of water potential [Ψ: 0.0 (control), ?0.15, ?0.30, ?0.45, ?0.60, ?0.75, and ?0.90 MPa] were adopted using polyethylene glycol-6000 (PEG6000) at four germination temperatures (15, 20, 25, and 30 °C). Final germination percentage, mean germination time, germination index, germination rate index, and germination stress tolerance index were used to evaluate the genotype response to PEG-induced water stress. Shoot and root length and fresh and dry weight were measured on seeds germinated at 20 °C under the different levels of water potentials. During germination, the three ornamental sunflowers showed to be more sensitive to suboptimal temperature than to supraoptimal. Decreasing water potential of imbibition solution progressively inhibited and delayed seed germination. Among cultivars, ‘Hadar’ and ‘Pazit’ performed better at temperature lower than 30 °C. ‘Zohar’ showed a lower sensitivity to PEG-induced water stress at all temperature conditions. Water stress during seed germination depressed the following seedling growth under favourable conditions. As a result, shoot and root length and fresh and dry weight was significantly lower in seedlings from seed germinated at ψ ≤ 0.45 MPa.     

Effects of bird ingestion on seed germination of Sorbus commixta

To determine the effects of ingestion by birds on seed germination, we performed germination experiments in the field and laboratory with Sorbus commixta. The germination of four groups of seeds was compared: ingested seeds, seeds defecated in feces after feeding of fruits to birds; extracted seeds, seeds deliberately extracted from the fruit pulp; juiced seeds, seeds plus the juice of the pulp after seeds had been deliberately extracted from the pulp; intact seeds, seeds in untreated intact fruits. In the laboratory, intact and juiced seeds hardly germinated, but ingested and extracted seeds germinated. Thus, the pulp and its juice appeared to inhibit germination, but seeds could germinate without ingestion by birds once the seeds had been manually extracted from the pulp. In the field, intact fruits did not germinate in the first spring, because the seed was still covered with pulp. The pulp of intact seeds decomposed during the first summer, and thus, the seeds had the potential to germinate during the second spring. In fact, most intact seeds do not germinate during the second spring either, since they lose their viability during the first summer. Thus, under natural conditions, most seeds of Sorbus commixta cannot germinate without bird ingestion. Received: 5 July 1997 / Accepted: 7 November 1997     

Effects of flood-influenced factors on seed germination of Ambrosia tenuifolia

Strong fluctuations are exhibited by populations of the perennial herb Ambrosia tenuifolia in the grasslands of the Salado basin (Province of Buenos Aires, Argentina), an area frequently enduring prolonged floods. Flooding causes the death of most dicotyledon plants of the community, A. tenuifolia among them, opening numerous gaps of various sizes. After the recession of the flood the density of A. tenuifolia seedlings was higher in flooded than in non-flooded plots and it was larger in wider gaps. Canopy removal in non-flooded plots increased field seedling emergence of A. tenuifolia up to the levels found in flooded plots. Responses of the seeds in the soil to gap-associated environmental factors such as light quality and temperature regime were studied both in the field and under controlled were studied both in the field and under controlled conditions. Seedling emergence was significantly enhanced when the red:far-red ratio of natural light reaching the soil surface under the canopy of nonflooded plots was increased by means of copper sulfate filters. The influence of light quality and temperature on germination of the soil seed population was also tested using grassland soil monoliths or mesocosms, transported from the field to the laboratory, in which the canopy was clipped and the soil exposed to either red or far-red light and kept at constant or fluctuating temperatures. Significant seedling emergence was observed only when the soil samples were exposed to red light and incubated at alternating temperatures. No emergence was recorded in samples exposed to far-red light or incubated at a constant 25°C. Seeds stored dry in the laboratory were also stimulated to germinate by red light and alternating temperatures but only after dormancy was sufficiently decreased by low temperature stratification or by low temperature under immersion. The results are consistent with the hypothesis that primary dormancy of A. tenuifolia seeds is decreased by low temperatures in winter even if the seeds are submerged as happens when floods occur. The decrease in dormancy makes the seeds prone to be stimulated to germinate by the Pfr form of phytochrome in combination with alternating temperatures. These conditions are likely to be met in the gaps opened by the flood-caused death of dicotyledon plants.     

Effects of osmotic priming and ageing on onion seed germination

Onion seeds were osmotically primed in polyethylene glycol solution (342 g/kg water) either for 14 days before accelerated ageing at 40°C. 18% m.c. for 0, 24, 48, 72 or 96 h, or for 10, 14 or 17 days after ageing. Priming improved the rate of germination compared with non-primed seed. Priming before ageing delayed the loss of viability due to ageing, but priming after ageing had no effect on viability. Primed and dried onion seed was stored for 18 months at 10°C, 9% m.c. with no effect on viability; improvements in germination rate due to priming were maintained over the storage period. Conductivity measurements of seed leachates were not a consistently reliable indicator of germination performance.     

光照和温度对飞机草种子萌发的影响

通过在人工培养箱内模拟环境条件,探讨了不同光照和变温对飞机草种子萌发的影响。结果表明:在有光照状况下,飞机草种子在15℃/10℃～40℃/35℃条件下均能萌发,飞机草种子萌发的最适变温为30℃/25℃,萌发率达47.5%;而黑暗条件下,飞机草种子在15℃/10℃时不能萌发;在20℃/15℃～35℃/30℃范围内,温度越高,飞机草种子萌发高峰的出现时间越早;在15℃/10℃～30℃/25℃范围内,飞机草种子的萌发率随温度的升高而升高,超过30℃/25℃后,萌发率下降,而适当的光照有利于飞机草种子的萌发。飞机草成为入侵种并迅速扩散与其种子萌发对光照和温度的适应性密切相关。     

不同种子预处理对10种沙拐枣植物萌发的影响

为了确定沙拐枣植物种子的萌发特性及最优播前预处理方法,在实验室条件下,对10种沙拐枣植物的种子进行了磨砺、硫酸和热水浸泡、冷藏、种子浸出液处理,然后进行发芽实验研究。萌发实验的结果表明,10种沙拐枣植物对于不同的种子预处理,均表现出相似的萌发反应。磨砺、硫酸浸泡和冷藏处理对种子萌发有明显地促进作用。与对照相比,种子浸出液处理对种子的发芽率、发芽速度均具有明显地抑制作用,并能增强种子的休眠。冷藏处理具有打破有活力的种子休眠、促进种子萌发的作用,但它与热水浸泡处理一样,对有活力种子表现出一定的致死作用。沙拐枣植物的萌发模式在不同种子预处理问均表现出明显的差异性。机械磨擦和硫酸处理能够促进种子的萌发率及发芽势。泡果沙拐枣(Callingonum junceum)在本项实验中表现出很强的萌发能力。     

三类抗性种子萌发对酸雨胁迫响应

实验采用 p H2 .0、2 .5、3.0、3.5、4 .0、5 .0模拟酸雨处理培养皿中的水稻 (Oryza sativa)、小麦 (Triticum aestivum)、油菜(Brassica chinensis var. oleifera) 3类抗性种子 ,每皿 5 0粒 ,置恒温培养箱内萌发 (2 5℃ ) ,每天更换 1次酸雨 ,处理与对照均 3次重复。定时测定酸雨胁迫强度、胁迫时间对种子发芽率、发芽势、发芽指数、活力指数、异状发芽率 ,吸水值、呼吸速率、贮藏物质运转效率、贮藏物质消耗率、根长抑制指数与芽长抑制指数的影响。结果表明 ,当酸雨胁迫强度 p H=2 .0～ 2 .5时 ,因胁迫强度过高 ,3类抗性种子皆不萌发 ;胁迫强度 p H≥ 3.0时 ,3类抗性种子 5项萌发指标的变幅是水稻 <小麦 <油菜 ;酸雨伤害阈值是水稻 (p H3.0～ 3.5 ) <小麦 (p H3.5～ 4 .0 ) <油菜 (p H4 .0～ 5 .0 ) ;当胁迫强度 (p H)≥ 2 .5时 ,3类抗性种子 6项生理指标的变幅是水稻 <小麦 <油菜 ,其生理反应阈值 (p H≥ 2 .0 ) <萌发反应阈值 (p H≥ 3.0 ) ;3类抗性种子贮藏物质消耗率、贮藏物质运转率 ,根长抑制指数及芽长抑制指数对酸雨胁迫时间的响应 (组间达到差异显著水平 )时间是 :水稻 (7d、7d,3d、3d)≥小麦 (6 d、6 d,3d、3d)≥油菜 (3d、4 d,3d、3d) ;3类抗性种子对酸雨胁迫强度与胁迫时间的耐受性差异 (     

Effects of a warmer climate on seed germination in the subarctic

Background and Aims

In a future warmer subarctic climate, the soil temperatures experienced by dispersed seeds are likely to increase during summer but may decrease during winter due to expected changes in snow depth, duration and quality. Because little is known about the dormancy-breaking and germination requirements of subarctic species, how warming may influence the timing and level of germination in these species was examined.

Methods

Under controlled conditions, how colder winter and warmer summer soil temperatures influenced germination was tested in 23 subarctic species. The cold stratification and warm incubation temperatures were derived from real soil temperature measurements in subarctic tundra and the temperatures were gradually changed over time to simulate different months of the year.

Key Results

Moderate summer warming (+2·5 °C) substantially accelerated germination in all but four species but did not affect germination percentages. Optimum germination temperatures (20/10°C) further decreased germination time and increased germination percentages in three species. Colder winter soil temperatures delayed the germination in ten species and decreased the germination percentage in four species, whereas the opposite was found in Silene acaulis. In most species, the combined effect of a reduced snow cover and summer warming resulted in earlier germination and thus a longer first growing season, which improves the chance of seedling survival. In particular the recruitment of (dwarf) shrubs (Vaccinium myrtillus, V. vitis-idaea, Betula nana), trees (Alnus incana, Betula pubescens) and grasses (Calamagrostis lapponica, C. purpurea) is likely to benefit from a warmer subarctic climate.

Conclusions

Seedling establishment is expected to improve in a future warmer subarctic climate, mainly by considerably earlier germination. The magnitudes of the responses are species-specific, which should be taken into account when modelling population growth and migration of subarctic species.Key words: Climate change, cold stratification, dwarf shrubs, germination percentage, incubation temperature, mean germination time, seedling establishment, seeds, Silene, subarctic species, Vaccinium, warming     

混合盐碱胁迫对芹菜种子萌发的影响

NaCl、Na2SO4、NaHCO3和Na2CO3按不同比例混合,模拟盐度和pH变化规律与天然盐碱地相似的15种盐碱条件,探讨混合盐碱胁迫对芹菜(Apium graveolens)种子萌发的影响.结果表明:芹菜种子的发芽率、发芽势、发芽指数和活力指数均随盐浓度的升高,pH的增大呈下降趋势.芹菜种子的萌发主要受盐浓度的影响,不同盐浓度间的影响差异大;当盐浓度为200 mmol/L时,基本不萌发.     

三维土壤异质性对种子萌发影响的实验研究

三维土壤异质性对种子萌发影响的实验研究 种子萌发受其生长环境的影响,但土壤异质性对种子萌发的作用还不太清楚。本文通过控制实验研究了三维土壤异质性对草本植物种子萌发的影响,实验设置了两种水平的土壤异质性,即通过在花盆内的三维空间上相间填充营养丰富和贫瘠两种土壤来构建不同的土壤异质性水平。两种水平土壤异质性的斑块大小分别为7.5和15.0 cm。实验采用两种草本植物(黑麦草Lolium perenne和垂穗披碱草Elymus nutans),每种植物选择大小一致的种子50粒,这些种子要么放在异质性土壤中萌发,要么放在培养皿中萌发。实验过程中,每天统计这些种子的萌发率。实验发现,斑块小的花盆内种子萌发率较低,这个结果与我们的预期相符,我们认为斑块小的花盆内,两种土壤斑块之间的间距较小,植物遭遇不同土壤斑块资源变动的几率较大,植物为了减小这种变动带来的不利影响,会倾向于降低萌发率和延迟萌发。我们的研究还发现,斑块小的花盆内种子萌发的变异性也较大,即与大斑块内的种子萌发率相比,小斑块内的种子萌发率在各重复间的差异性较大。与二维土壤异质性相比,三维土壤异质性更接近于真实情况,这样的土壤设置能够促进我们更好地理解土壤空间异质性对种子萌发等动态过程的影响.     

镧浸种对水稻种子萌发及幼苗生长的影响

以水稻种子为材料,研究了稀土元素镧对水稻种子活力和幼苗生长的影响。结果表明,1~20mg.L-1La3+处理能提高水稻种子的活力,提高蛋白酶、淀粉酶和脂肪酶的活性,并能促进水稻茎、叶和根系的生长,其中10mg.L-1La3+处理效果最佳。当浓度超过30mg.L-1时,水稻种子萌发和幼苗生长受到抑制。此外,镧对水稻植株地下部分(根系)生长的影响比地上部分(茎叶)生长的影响要明显。镧对水稻种子萌发期间3种酶活性的影响程度表现为蛋白酶>脂肪酶>淀粉酶。