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。     

Additive and nonadditive effects of herbivory and competition on tree seedling mortality, growth, and allocation

The interaction between simulated cotyledon herbivory and interspecific competition was studied in a greenhouse experiment using two species of trees, Acer rubrum and Quercus palustris, which commonly invade abandoned agricultural fields. Herbivory treatments were applied as a gradient of cotyledon removal for A. rubrum with 0, 25, 50, 75, and 100% of cotyledon tissue removed. Cotyledons from Q. palustris were clipped and removed (control, early, and late removal) to create a gradient of seed reserve availability. The competition treatment consisted of plugs of old-field vegetation that filled the pots with perennial cover. Mortality of seedlings was higher with competition. There was a significant interaction between herbivory and competition with the highest mortality occurring with competition at the highest intensity of herbivory in both species. Herbivory reduced biomass for Q. palustris only, while competition reduced biomass in both species. Neither species showed an interaction between herbivory and competition for growth. There was a significant interaction between herbivory and competition on allocation patterns for both species, with greater allocation to roots with competition at the highest intensity of herbivory. This study demonstrates the potential for cotyledon herbivory and competition to interact, altering the invasion of tree seedlings into abandoned agricultural land.     

Mechanisms of seedling mortality by subterranean insect herbivores

Summary The nature of the mortality of germinating seeds of Vicia sativa, the common vetch, was investigated in a pot trial under controlled conditions. Chafer and tipulid larvae were restricted by nylon mesh partitions to enable radicle and/or hypocotyl herbivory to occur. The effects of the two insects were very similar. In control situations, an average of 88% of the viable seed sown recruited successfully. Hypocotyl and radicle herbivory had similar effects on seedling mortality, with recruitment of viable seed sown being reduced to 52%. When both modes of attack occurred together, successful recruitment was only 34%. Feeding on both plant parts resulted in an average post-emergence mortality of 14% of the seed sown, but the effect on pre-emergence mortality was of greater importance, amounting to 41%. The mechanisms by which soil-dwelling herbivores may cause seedling mortality in the field are discussed.     

The ecology of predispersal insect herbivory on tree reproductive structures in natural forest ecosystems

Plant-insect interactions are key model systems to assess how some species affect the distribution, the abundance, and the evolution of others. Tree reproductive structures represent a critical resource for many insect species, which can be likely drivers of demography, spatial distribution, and trait diversification of plants. In this review, we present the ecological implications of predispersal herbivory on tree reproductive structures by insects (PIHR) in forest ecosystems. Both insect's and tree's perspectives are addressed with an emphasis on how spatiotemporal variation and unpredictability in seed availability can shape such particular plant-animal interactions. Reproductive structure insects show strong trophic specialization and guild diversification. Insects evolved host selection and spatiotemporal dispersal strategies in response to variable and unpredictable abundance of reproductive structures in both space and time. If PIHR patterns have been well documented in numerous systems, evidences of the subsequent demographic and evolutionary impacts on tree populations are still constrained by time-scale challenges of experimenting on such long-lived organisms, and modeling approaches of tree dynamics rarely consider PIHR when including biotic interactions in their processes. We suggest that spatially explicit and mechanistic approaches of the interactions between individual tree fecundity and in sect dynamics will clarify predictions of the demogenetic implications of PIHR in tree populations. In a global change context, further experimental and theoretical contributions to the likelihood of life-cycle disruptions between plants and their specialized herbivores, and to how these changes may gen erate novel dynamic patterns in each partner of the interaction are increasingly critical.     

Dispersion pattern and mortality of seeds and seedlings ofFagus crenata Blume in a cool temperate forest in western Japan

The seed and seedling mortality ofFagus crenata Blume after a mast year (1993) was examined in relation to density and distance from the nearest conspecific adult tree in a mixed conifer-hardwood forest in Ohdaigahara, western Japan. The mortality of fallen seeds during winter amounted to 93.7%, and 79.2% of the current-year seedlings died in the first growing season. The most important factor of death for both seeds and seedlings was predation by vertebrates. The mortality of seeds during winter was positively correlated with sound seed density. The mortality of seedlings was positively correlated with density but not significantly related to the distance from the nearest crown edge of a conspecific adult tree. Mortality patterns varied with stages and spatial scales due to the behavior of predators; it is thus important to investigate the spatial pattern of seeds and seedling mortality at various temporal and spatial scales. After the first growing season, the difference in seedling density between distance classes was not significant at <4m from the nearest adult trees due to density-dependent mortality. However, seedling density was significantly lower in the ≥4 m class than in the <4 m classes.     

Tree seedling establishment under insect herbivory: edge effects and inter- annual variation

As the density and species composition of insects may change in relation to distance from the forest edge, the role of herbivory in tree establishment may also change across edges. To determine the importance of insect herbivory in tree establishment, insect densities were experimentally altered at different distances from the forest edge. Plots were established at three distances from the edge, with plots located in forest, edge, and field habitats. In half of each plot, insect densities were reduced by insecticide application. Seeds of two tree species, Acer rubrum and Fraxinus americana, were planted into each plot in 1995. The experiment was repeated in 1996 with the addition of Quercus palustris and Quercus rubra.Distance from the forest edge was the most important factor in determining seedling emergence and mortality. Overall seedling performance increased from field to edge to woods, although responses varied among species. In 1995, a drought year, insect removal increased emergence and decreased mortality of tree seedlings. In 1996, a year with normal precipitation, insect removal had much less effect on A. rubrum and F. americana. For the two Quercus species, mortality was reduced by insect removal. The tree species differed in their susceptibility to insect herbivory, with Acer rubrum the most susceptible and Fraxinus americana the least. Herbivory by insects was shown to have the potential to affect both the composition and spatial pattern of tree invasions. Herbivore importance differed greatly between the two years of the study, making the interaction between insects and tree seedlings variable both in space and time.     

High mortality,fluctuation in numbers,and heavy subterranean insect herbivory in bush lupine,Lupinus arboreus

Sporadic patchy die-off of bush lupine, Lupinus arboreus, has long been known. We describe in detail a series of these incidents on the central California coast, based upon observational and comparative evidence. Stands of thousands of plants die, while nearby mature plants live on. In some sites, repeated die-off followed by regeneration from the seed bank has led to the cover and density of this woody, perennial plant fluctuating widely over the 40 year period for which records exist. Root damage by caterpillars of the ghost moth or swift Hepialus californicus (Lepidoptera, Hepialidae) is a major cause of individual bush death and a probable cause of die-off of stands of lupine. Hidden from view underground, a few of these insects readily kill a juvenile or young mature plant by girdling and reaming-out roots. The mass mortality of L. arboreus that we observed involved heavy root damage by these caterpillars in evenaged stands of plants in their first (1.5-year-old) or second (2.5-year-old) flowering season. The injured plants set seed before dying. Older, larger bush lupines better withstood root damage. In plants aged 3 or more years, damage and mortality were correlated with the intensity of ghost moth caterpillars in the roots. At the highest intensity (mean = 37.5, maximum = 62 caterpillars/root), a stand of large, old L. arboreus suffered 41% mortality; 45% of root cambium (median value) was destroyed by feeding caterpillars. Mass death of mature L. arboreus was not correlated with folivory, and leaf damage ranged from nil to moderate in instances of die-off. The western tussock moth, Orgyia vetusta, accounted for the highest levels of folivory, but this insect was rare when die-offs occurred. The lowest lupine mortality rates in our study occurred where tussock caterpillar intensities were high and where plants were repeatedly defoliated by this insect. However, experimental defoliation by high, but realistic, intensities of tussock moth caterpillars resulted in some mortality of mature bushes, and the combined effects of leaf and root herbivory have yet to be assessed. In its natural range on the California coast, bush lupine has several additional species of insect herbivores that can be locally abundant and injurious to the plant, although none is associated with die-off. Subterranean natural enemies of ghost moth caterpillars may play a role in the patchy waxing and waning of this shrub. Locally, a new species of entomophagous nematode (Heterorhabditis sp.) cause high mortality in the soil, before ghost moth caterpillars have entered the root. This natural enemy may thus afford lupines protection from heavy underground herbivory.     

生物土壤结皮对种子萌发和幼苗建植的影响研究进展

生物土壤结皮（BSC）在陆地生态系统中具有重要的生态地位,尤其是旱地生态系统中,BSC占据了种子植物之间的广阔地面。因此,BSC的发展必然影响种子植物更新过程与植被空间格局;但其作用方式、影响程度等因相关研究涉及多气候要素、土壤类型、BSC组成物种和种子植物物种的差异及其不同组合,导致目前的研究结论存在广泛争议。研究综合论述了BSC改变地表微形态对种子传播过程的影响;BSC改变土壤特性（物理、化学、生物学）对种子萌发和幼苗存活与建植等关键环节的影响;并结合种子形态特征及种子萌发、幼苗建植的性状等,综合分析了BSC对种子传播、种子萌发与幼苗建植等更新过程的潜在影响机理;探讨了目前研究矛盾性结论产生的原因。总体来说,深入研究并全面揭示BSC对种子植物更新过程的影响,应加强学科交叉,将分子生物学、植物生理学、生物化学等微观研究,与遥感、野外生态因子过程监测、控制实验等宏观、中观研究结合,从机理到过程方面动态研究BSC对种子植物更新过程的影响,并引入水文模型、气候模型、种群动态模型等模型预测方法,研究气候变化、各类干扰频发的情景下,BSC对种子传播、萌发及幼苗建植过程的潜在影响,以期促进对BSC与种子植物间相互关系的研究,加深对干旱脆弱生态系统植被发展规律的认识。     

Reproductive ecology of Thymelaea velutina (Thymelaeaceae)- Factors contributing to the maintenance of heterocarpy

Thymelaea velutina (Thymelaeaceae) is a dioecious shrub that presents a unique type of heterocarpy which consists of the simultaneous production of dry and fleshy fruits. It is endemic to the Balearic Islands (Western Mediterranean) and is found both in dunes and mountain areas. The goal of this study was to identify which factors influence the production of both fruit types, examining the variation of their effects at a spatio-temporal scale (comparing two localities in different years). Specifically, we investigated (1) whether pollen limitation influences the type of fruit produced, and (2) the possible differences in seed size, mass, dispersal capacity, seed predation, germination patterns and seedling survival between fruit types. We also examined if the production of fleshy fruits was modified with the application of gibberellins to reproductive branches. Although fleshy fruits were consistently more abundant than dry ones at both populations, their proportion was significantly higher at the site with greater precipitation. The addition of either pollen or gibberellins did not affect the proportion of each fruit type. Seeds in fleshy fruits are consistently larger, heavier and more likely to be dispersed than seeds in dry fruits, but germinability, germination rate and seedling survival was similar among fruit types. Heterocarpy in this species is currently maintained as there is no apparent factor that exerts any strong selective pressure on either fruit type. The two fruit types might even have different `functions', one serving especially for dispersal and population expansion and the other for producing a seed bank that ensures an eventual germination.     

Effect of tetcyclacis and chlormequat applied to seed on seedling growth of triticale cv. Lasko

The growth of triticale seedlings from seeds treated with three concentrations of the plant growth regulators (PGRs), tetcyclacis and chlormequat, with or without drying after soaking was studied. Both tetcyclacis and chlormequat inhibited shoot growth. They reduced shoot:root ratios, first by restricting shoot growth (one week after treatment) and later by boosting root growth (eight weeks after transplanting). At the concentrations used tetcyclacis was a more active PGR than chlormequat and promoted tiller production. Drying, after soaking, promoted root growth, retarded the elongation growth of seedlings and enhanced some of the effects of the PGRs. Analyses of regressions between those growth characteristics significantly influenced by PGRs and the concentration revealed a quadratic relationship.     

六种阔叶树种叶片水浸提液对云南松种子萌发生长的化感作用

为了筛选营建云南松抗虫混交林的树种,该研究观察了云南切梢小蠹非寄主植物川滇桤木、缅桂、滇朴、樟树、麻栎和山茶六种阔叶树种叶片不同浓度的水浸提液对云南松种子萌发及幼苗生长的化感作用.结果表明:(1)川滇桤木、滇朴、麻栎和山茶四种阔叶树种叶片的水浸提液对云南松种子萌发和幼苗生长表现出低浓度促进,高浓度抑制效应.(2)在测试...     

Spatial patterns of tree seedling establishment and their relationship to environmental variables in a cold-temperate deciduous forest of eastern North America

Patterns of seedling recruitment may have persistent effects on population and community processes. Assuming seed availability is not limiting, the environmental sieve (i.e., the suite of factors influencing seed germination and seedling emergence and survival) determines how many seedlings establish and, most importantly, where they do so. In this study, we identify the spatial structure of some resources and abiotic conditions known to be significant for tree seedling emergence and survival and determine how these environmental factors influence the establishment of Fagus grandifolia, Acer saccharum, Fraxinus americana, and Ostrya virginiana in a deciduous forest of southern Québec (Canada). We expect an increase from Fagus, through Acer and Fraxinus, to Ostrya in the control of environmental variables on seedling emergence and survival, because of differences in the seed size of these species. Density of newly-emerged seedlings of all four species showed positive spatial autocorrelation at distances of up to ca. 10 m. Environmental variables were also structured at the same spatial scale, except for soil moisture. Acer seedling emergence pattern was positively correlated to photosynthetic photon flux density (PPFD), and the pattern of Fraxinus to soil N and moisture. Seedling survival was not spatially autocorrelated for any of the four species, although it was positively density-dependent in Acer and Fagus. In only Ostrya was seedling survival correlated (positively) to one of the environmental variables studied, i.e., PPFD. Overall, environmental variables were spatially less heterogeneous than seedling emergence and survival. Either seed availability was not saturating or factors not considered here, such as competition and predation (the intensity of which often varies with resources and/or abiotic conditions), modified the influence that the physical environment had on patterns of seedling establishment. Our prediction of a greater environmental control on seedling emergence and survival in small-seed species was not totally confirmed.     

不同处理对银沙槐种子休眠破除和幼苗生长的影响

对银沙槐种子破除休眠的适宜方法及其幼苗生长进行研究。结果表明:破口和沙磨处理休眠破除效果最好,但容易引起种子腐烂死亡;分别用65、80和95℃热水处理种子,以95℃1 min破除种子休眠效果最好,且幼苗长势好,65℃效果最差,80℃可破除部分种子休眠,除80℃外延长热水处理时间可导致死种子数增加,95℃则引致大部分种子死亡;液氮浸种处理效果均显著,其中以液氮处理10 min效果最好,但幼苗多为不正常苗而致死;硫酸处理以浸种120 min效果最好,但处理时间过长可导致死种子数增多,且造成少数幼苗根部腐蚀死亡。综上所述,打破银沙槐种子休眠的较优处理方法是硫酸120 min浸种处理和95℃1 min热水处理。     

Factors affecting seedling recruitment and survivorship of the Japanese subalpine stone pine, <Emphasis Type="Italic">Pinus pumila</Emphasis>, after seed dispersal by nutcrackers

To clarify the factors affecting the seedling establishment of the subalpine stone pine (Pinus pumila Regl.) from nutcracker-cached seeds, subsequently emerged seedlings were examined in two plots on Mt Akitakomagadake, northern Japan. The survivorship of older seedlings (2–20years old) was also monitored during six seasons at nine different sites. In one plot with 18 caches, seed germination occurred between June and mid-July. During this period, nutcrackers retrieved the stored pine seeds from 16 caches and ate some seeds immediately. Simultaneously, this nutcracker behavior caused mechanical damage to newly emerging seedlings (e.g. uprooting and tearing off cotyledons). Such initial loss to in situ harvesting and mechanical damage accounted for 75% of the total seeds remaining in the caches. The number of established current seedlings sharply declined during the first 2years, and the survival rate was 4.4% over four winters. Two major mortality factors were identified: uprooting by frost-heave soil disturbance in the spring of the second year and standing death by drought or other physiological stresses in early summer. In another plot with 13 caches, survivorship of newly emerged seedlings was also low (4.2%), but mortality was mostly due to summer drought, indicating that the frost-heave event was a site-specific disturbance factor. For older seedlings, survival rates reached approximately 90% even after six seasons and summer drought stress was a major mortality factor. My findings suggest that seedling recruitment of P.pumila was largely limited by both nutcracker disturbance and external disturbance (and/or stresses) at an early stage. However, after the critical first few years, pine seedlings were highly likely to survive and grow to the sapling stage despite the harsh environment of the high mountains.     

模拟干旱胁迫对巴东木莲种子萌发和芽苗生长的影响

巴东木莲原生种群天然更新差,林下鲜见更新幼苗或幼树.为探究其种子萌发和芽苗在生长期对干旱胁迫的耐受性,该文以巴东木莲种子为材料,采用不同质量浓度的聚乙二醇(PEG-6000)模拟干旱胁迫,分析干旱胁迫对其种子萌发、芽苗生长及其相关的生理生化指标的影响.结果表明:(1)不同质量浓度的PEG溶液对巴东木莲种子萌发和芽苗生长...     

埋深和播种密度对豚草种子出苗及幼苗生长的影响

豚草是一种全球性的恶性入侵杂草,给我国生物多样性及经济发展造成了巨大威胁。为研究豚草种子出苗和幼苗生长的影响因素,采用盆栽实验分析了种子大小（L、M、S）与埋深（2、4 cm和6 cm）或播种密度（2、4粒/盆和8粒/盆）对豚草（Ambrosia artemisiifolia）种子的出苗和幼苗生长的影响。研究结果表明,埋深对豚草种子出苗的影响高于播种密度及种子大小的影响,较浅的埋深有利于豚草种子出苗;出苗率在不同播种密度下均具有较高水平,达到67.9%-100%,这种高出苗率是豚草在不同生境成功定植的原因之一。豚草的幼苗生长受埋深或播种密度的影响大于种子大小的影响,且小种子更易受埋深或播种密度的影响。种子大小显著影响豚草幼苗的株高和基径（P<0.05）,总体上较大种子的株高和基径高于小种子,此外更小的种子会将生物量更多的分配给根以促进幼苗的生长。幼苗的株高和基径在较浅的埋深下更高,而豚草幼苗的单株生物量、单株地上和地下生物量随埋深的增加而增加。播种密度的增加会加剧豚草幼苗之间对水分、营养、光照等资源的竞争,导致其单株地上、地下及总生物量显著降低（P<0.05）。研究发现豚草在浅埋深、低密度生境中更容易入侵成功,因此可以通过对土壤进行深翻,采取替代控制,种植竞争能力强的本地植物等手段有效管理和防治豚草。     

辽东栎幼苗生长对种子大小和子叶去除处理的响应

该文在温室内盆播条件下,设置不去除子叶(对照)、去除1/4子叶(轻度子叶去除)、1/2子叶(中度子叶去除)和3/4子叶(重度子叶去除)4个处理,研究了种子大小和子叶去除处理对辽东栎幼苗生长及其形态学可塑性的影响.结果表明:(1)大种子幼苗株高、基茎、叶片数、单株叶面积、总干质量和根冠比在所有处理均大于小种子幼苗,除株高...     

不同种源五角枫幼苗叶片N、P化学计量学特征对氮添加的响应

不同土壤氮环境下植物对氮沉降的响应趋势不同,因此研究不同种源地的五角枫(Acer mono)对氮沉降的响应机理具有重要的意义。为了深入了解不同种源五角枫对氮沉降的适应和响应机制,主要通过3个种源的五角枫幼苗氮添加控制实验,分析不同种源的五角枫幼苗比叶面积(SLA)和叶片干物质含量(LMDC)以及叶片N、P含量和N∶P对氮添加的响应规律。结果表明:(1)不同氮添加处理下,内蒙古种源五角枫幼苗(NW)和山西种源五角枫幼苗(SW)的LDMC均显著低于对照,NW和SW在N4条件下表现为显著的促进作用。随着氮添加浓度的增加,NW与北京种源的五角枫幼苗(BW)的SLA在N4条件下显著增加;SW的SLA在N2的水平下显著增加。(2)氮添加下,只有SW的P_(mass)在N1水平下显著增加,N_(mass)在N1和N3水平下表现为显著增加。说明在低氮水平下,能够促进SW对P的吸收,从而增加植物体内对P的积累。同时在低氮和中高氮条件下,能够促进SW对N的吸收,说明SW能较好的适应低氮和高氮环境,使其生存能力更强。(3)分析变异来源及各指标的相关性得出,氮添加条件下,五角枫的SLA、LDMC、P_(area)及N∶P与氮添加处理呈显著的相关关系,而SLA、LDMC、N_(mass)、P_(mass)、Narea、P_(area)和N∶P对氮添加处理和种源的交互作用均未表现出相关关系,说明引起五角枫各性状主要受氮添加处理影响。(4)综合各指标的变化,氮添加条件下,NW的LDMC显著降低,SLA显著增大,同时叶N含量、叶P含量以及N∶P比较高;SW的LDMC显著降低,SLA增大,同时叶N含量、叶P含量表现为显著增大,结合叶经济谱理论分析得出,SW和NW属于"快速投资-收益"型,而BW倾向于"缓慢投资-收益"型。因此,不同种源的五角枫对于氮沉降的响应存在差异,并且持续的氮添加已经改变了五角枫的适应性。