Comparative effects of desiccation,heat shock and high temperatures on seed germination of savanna and forest tree species

Although forest and savanna biomes predominate in tropics regions, the factors that control their distribution remain unclear. South American savannas occur in regions that are considered warm and humid enough to support forests, indicating that agents other than climate determine the occurrence of one or the other physiognomy. Herbivory, fire and water deficit have been considered environmental filters that limit the forest species encroachment in savanna physiognomies, but the effects of these filters on the capability of these species to recruit from seeds remain poorly understood. In this study we investigated how stress factors characteristic of savanna environments, such as soil desiccation, heat shocks and high temperatures affect the survival and germination of seeds from savanna and forest tree species. We found that desiccation (to 5%) reduced the germination percentage of forest seeds, but had no effect on the germination of savanna seeds. Forest seeds were less tolerant to heat shocks of 140°C and 200°C, and showed lower germination percentage at temperatures of 35 and 40°C, when compared with savanna seeds. Savanna seeds presented longer germination times and higher germination variance than forest seeds, indicating a risk‐spreading germination strategy among savanna species. The low tolerance of forest seeds to desiccation, heat shock and high temperatures may explain the low recruitment of forest trees into savanna physiognomies. Climate change models predict lower soil moisture, higher temperatures and higher fires frequency for South America biomes. Our results suggest that savanna species are likely to be more capable of withstanding the effects of these changes than forest species.     

热激对大兴安岭三种松科树种种子萌发的影响

研究了热激对大兴安岭地区兴安落叶松、樟子松、红皮云杉种子萌发的影响,热激温度分别为80℃、100℃、120℃、150℃,每个温度下热激时间分别为1min、3min、5min、10min。研究结果表明:兴安落叶松种子在100℃热激10min,120℃热激5min,10min,150℃热激5min处理下萌发率较对照显著下降(P%0.05),除80℃热激1min、3min、5min、120℃热激1min、150℃热激1min外,其他处理萌发速率指数较对照均显著下降(P%0.05);樟子松种子在高于80℃的所有热激处理萌发率较对照均显著下降(P%0.05),甚至没有萌发,除80℃热激1min外,所有处理下萌发速率指数较对照均显著下降;红皮云杉种子萌发率在80℃热激5min和100℃热激1min处理下较对照显著提高(P%0.05),萌发速率指数在80℃热激1min、3min、5min,100℃热激1min处理下较对照显著提高(P%0.05)。兴安落叶松种子可以在短时间高强度(150℃)热激下维持萌发率,但不能承受长时间热激;樟子松种子对高于80℃的热激反应敏感,热激降低了樟子松种子的萌发能力;热激可以提高红皮云杉种子的萌发能力,短时间(1min、3min、5min)低强度(80℃)热激(包括100℃热激1min)提高红皮云杉种子萌发率和萌发速率指数,红皮云杉种子也可以承受高强度(150℃)瞬时(1min)热激。     

Interactive effects of heat shock and smoke on germination of nine species forming soil seed banks within the Sydney region

Abstract The germination response of seeds from fire‐prone vegetation to fire‐related cues such as heat shock and smoke has usually been studied by applying the cues singly. The few studies that have applied the cues in combination have shown that interactions between the cues are possible. Here, the response of seeds from a number of species to combined heat shock and smoke is reported. Heat shock (25, 50, 75 and 100°C) and aerosol smoke (0, 5, 10 and 20 min) were applied factorially to nine species that form soil seed banks in the Sydney region of south‐eastern Australia. These species were from Epacridaceae (four species), Myrtaceae (four species) and Cyperaceae (one species) and ranged from fire‐sensitive obligate seeders to fire‐tolerant facultative resprouters. Germination of Dracophyllum secundum R. Br and Sprengelia monticola (A. Cunn. ex DC.) Druce was low and did not respond to the germination cues. The positive response of Gahnia sieberiana Kunth and Kunzea ambigua (Sm.) Druce to heat shock and smoke was independent and additive. The positive response of Kunzea capitata Rchb. to the interaction between heat shock and smoke was synergistic, and the response of Baeckea diosmifolia Rudge and Baeckea imbricata (Gaertn.) Druce was unitive, with germination increase only occurring following combined heat and smoke application. Epacris coriacea A. Cunn. ex DC. and Epacris obtusifolia Sm. had low levels of dormancy and hence it was not possible to find a fire response. Gahnia sieberiana and K. capitata responded differently to the combination of heat shock and smoke than has previously been reported. Germination of species from habitats that are infrequently burnt was not affected by heat shock or smoke. Low‐intensity fire or patches within fire may be important for seedling recruitment as the 50°C heat shock stimulated germination in four of the five species that responded to the heat cue, and germination of Baeckea imbricata declined within the 100°C heat shock treatment. Germination of one species, Baeckea imbricata, was only stimulated by a specific combination of cues, indicating that regeneration niches may be narrow for some species and that the application of a range of heat and smoke doses is required to find such responses. Of the species positively responding to heat shock and smoke, a requirement for both cues was prevalent, therefore the response to these cues in isolation cannot be relied upon to give a true indication of the fire response of a species.     

Response surfaces for the combined effects of heat shock and smoke on germination of 16 species forming soil seed banks in south‐east Australia

Abstract There is limited understanding of how fire‐related cues such as heat shock and smoke can combine to affect the germination response of seeds from fire‐prone vegetation because combinations of multiple levels of both cues have rarely been investigated. Germination response surfaces were determined for the combination of heat shock and smoke by applying factorial combinations of temperature (up to 100°C) and aerosol smoke (0–20 min) to 16 species that form soil seed banks in the Sydney region of south‐eastern Australia. Duplicate populations of three species were also examined to assess the constancy of a species response surface. Of the 19 populations examined, 16 showed a germination response to both the fire cues, which combined interactively in 14 populations, and independently in two. No population responded only to a single cue; however, seeds of 11 populations responded to heat in the absence of smoke, and nine responded to smoke in the absence of heat. Heat applied in the absence of smoke negatively affected germination in seven populations, either progressively as temperature increased, or above a set temperature. Negative germination responses over part of the temperature range were fully reversed at higher temperatures for unsmoked seeds of four populations (curvilinear heat response). Smoke effects were most frequently positive over all or part of the range of durations used, and when combined with heat frequently fully or partially reversed negative heat effects. Three populations required the obligatory combination of smoke and heat. A novel response to the cues was observed for three species, with smoke reversing negative heat effects at 75°C, being supplanted by a positive heat response of unsmoked seed at 100°C. The response surface for duplicate populations of two of the three species examined was variable. Heat shock and smoke frequently combined to affect germination, in both positive and negative ways. Consequently, to gain an accurate assessment of the response of seeds to fires, an experimental design that samples within the potential response zones of germination cues is essential.     

Germinable soil seed banks in a tropical savanna: seasonal dynamics and effects of fire.

Abstract The germinable soil seed bank of a tropical eucalypt savanna of north‐eastern Australia was found to be dominated by grasses and forbs, with seed bank density ranging from 58 to 792 seeds per square metre, from a total of 53 species. Late dry season fires and the fire‐related cues, heat shock and smoke, broke the seed dormancy of a range of tropical savanna species. Heat shock promoted the germination of the species groups natives, exotics, subshrubs, ephemeral and twining perennial forbs, and the common species Indigofera hirsuta, Pycnospora lutescens and Triumfetta rhomboidea. Exposure to smoke at ambient temperature promoted germination from the soil seed bank of the species groups combined natives, upright perennial forbs and grasses, as well as the common grasses Digitaria breviglumis and Heteropogon triticeus. The germinable soil seed bank varied seasonally, increasing from the mid wet season (February) and early dry season (May) to a maximum in the late dry season (October). The effect of recent fire history on soil seed bank dynamics was limited to the immediate release of some seed from dormancy; a reduction in seed densities of subshrubs and monocots, other than grasses, in recently burnt savanna; and enhanced seed density of the ephemeral I. hirsuta in the year following fire. The seed banks of most savanna species were replenished in the year following burning.     

Fire‐related cues break seed dormancy of six legumes of tropical eucalypt savannas in north‐eastern Australia

Abstract This paper describes an assessment of the effect of exposure to fire‐related cues (heat shock, smoke and nitrate) and the interactions between the cues on seed dormancy release of tropical savanna legumes in north‐eastern Australia. Ten legume species were tested, comprising both native and exotic species. The ten species responded variously to the treatments. Brief exposure to temperatures between 80 and 100°C was found to break the seed dormancy of the native ephemeral herbs Chamaecrista mimosoides, Crotalaria calycina, Crotalaria montana, Indigofera hirsuta and Tephrosia juncea, as well as the exotic ephemeral herb Crotalaria lanceolata. Exposure to 80°C combined with treatment with a nitrate solution produced an additive effect on the germination of Chamaecrista mimosoides and Crotalaria lanceolata. However, the four species with the heaviest seeds, two exotic ephemeral herbs (Chamaecrista absus and Crotalaria pallida) and two native perennials (Galactia tenuiflora and Glycine tomentella) displayed no significant increase in germination with exposure to fire‐related cues. Exposure to 120°C for 5 min produced seed mortality in all species tested. Two of the largest seeded species, Crotalaria pallida and Galactia tenuiflora, displayed the lowest tolerance to heat shock, with seed mortality after exposure to 100°C for 5 min. These data indicate that fire can promote the germination of some tropical savanna legumes. As a proportion of seeds of each species displayed no innate dormancy, some germination may occur in the absence of fire, especially of exotic species.     

Distinct roles of savanna and forest tree species in regeneration under fire suppression in a Brazilian savanna

Questions: Has fire suppression relaxed barriers to the exchange of species between savanna and forest? Do all species or a subset of species participate in this exchange? Would current vegetation structure persist if fire suppression were to cease? Location: A gallery forest edge in the Cerrado region of central Brazil that burned only once in the past 35 years. Methods: Density of tree seedlings, saplings and adults, leaf area index (LAI), tree basal area and diameter were surveyed in 12, 10 m × 70 m transects centred on and perpendicular to the forest–savanna boundary. Community composition was assessed using non‐metric multi‐dimensional scaling (NMDS). Results: Basal area and LAI declined substantially from forest to savanna, with an associated shift in species composition. Savanna tree species were nearly absent in the forest, but accounted for the majority of stems in the savanna. In contrast, forest species comprised 14% of adults and more than one‐third of juveniles in the savanna. Despite the high diversity of trees (85 species) in the forest, five species play a particularly large role in this initial phase of forest expansion. Reintroduction of fire, however, would result in widespread topkill of juveniles and the majority of adult forest trees, thereby interrupting the succession towards forest. Conclusions: After 35 years during which the site burned only once, the savanna still remains dominated by savanna species. Nevertheless, the dominance of forest juveniles in border and savanna tree communities suggests that with a continued policy of fire suppression, the forest will continue to expand.     

Diversity of germination strategies and seed dormancy in herbaceous species of campo rupestre grasslands

The effects of fire on the vegetation vary across continents. However, in Neotropical fire‐prone grasslands, the relationship between fire and seed germination is still poorly understood, while their regeneration, especially after strong anthropogenic disturbance, is challenging for their conservation. In the present study, we assessed diversity of germination strategies in 15 dominant herbaceous species from Neotropical altitudinal grasslands (locally known as campos rupestres). We exposed seeds to several fire‐related treatments. We also compared germination between regularly and post‐fire fruiting species. Finally, we investigated the diversity of dormancy classes aiming at better understanding the biogeography and phylogeny of seed dormancy. Germination strategies varied among families. Velloziaceae and Xyridaceae produced non‐dormant, fast‐germinating seeds. Cyperaceae and Poaceae showed an extremely low or null germination due to a high proportion of unviable or embryo‐less seeds. The seeds of campo rupestre grasslands are fire resistant, but there is no evidence that fire triggers germination in this fire‐prone ecosystem. Although heat and charred wood did not promote germination, smoke enhanced germination in one grass species and decreased the mean germination time and improved synchrony in Xyridaceae and Velloziaceae. Fire had a positive effect on post‐fire regeneration by stimulating fruit set in some Cyperaceae and Poaceae species. These species produced faster germinating seeds with higher germination percentage and synchrony compared to regularly fruiting Cyperaceae and Poaceae species. This strategy of dispersion and regeneration seems to be an alternative to the production of seeds with germination triggered by fire. Physiological dormancy is reported for the first time in several clades of Neotropical plants. Our data help advance the knowledge on the role of fire in the regeneration of Neotropical grasslands.     

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。     

Ecophysiological effects of light quality and nitrate on seed germination in species from Western Australia

Germination occurs usually in response to multiple environmental cues. Seeds with the ecophysiological ability to simultaneously sense the previous presence of fire and appropriate levels of temperature, light and soil nitrate could restrict germination to postfire, winter and competition-free microhabitats, where the potential for seedling survival is enhanced. Germination responses of 16 species with a range of life forms, fire responses and seed weights were determined under controlled conditions of 15°C temperature, a 12 h light cycle, exposure to 1 g L^?1 nitrate solution, and six conditions of light quality (white, blue, yellow, red, far-red light and darkness). Germination in Oenothera stricta, a weedy naturalized ephemeral, and two small-seeded indigenous Asteraceae species of mulga woodlands, Leucochrysum fitzgibbonii and Craspedia sp., were enhanced by white, yellow or red light compared with germination achieved in the dark, or under far-red or blue light. In red light, KNO₃ further enhanced germination of these positively photoblastic species. The germination response of Trachyandra divaricata, a naturalized herb of sandy, seaside locations, and several native jarrah forest legumes (four Acacia species, Bossiaea aquifolium, Gompholobium marginatum and Sphaerolobium vimineum) proved to be negatively photoblastic. Of these seven negatively photoblastic herb and shrub species, exposure to KNO₃ overcame the inhibition of light only in the resprouter species, Acacia lateriticola. In the serotinous, negatively photoblastic tree species, Corymbia calophylla and Eucalyptus marginata, KNO₃ seemed to be required before the negative response to light exposure was recorded. A dose–curve experiment on two positively photoblastic and three negatively photoblastic species indicated that although KNO₃ exposure affected germination in all species, different concentrations of KNO₃ (0, 0.5, 1, 2, and 5 g L^?1) produced different levels of response. Detailed studies with additions of KNO₃ (1 g L^?1) and the growth hormone, gibberellic acid (GA₃; 50 mg L^?1), showed that increased germination percentages of the positively photoblastic species, Oenothera stricta, occurred in the light, but blocking endogenous gibberellic synthesis with paclobutrazol, or adding exogenous GA₃ or KNO₃ had no effect on the light-induced germination levels. In the negatively photoblastic species Trachyandra divaricata, additions of KNO₃ and GA₃ had no influence on the germination inhibition induced by exposure to light nor did blocking endogenous GA synthesis. The 16 species growing naturally in Western Australia, Australia show a range of germination responses to environmental conditions, but depending on their natural habitat, the ecophysiology of each species appears to be optimized for subsequent seedling survival.     

Direct seeding of Brazilian savanna trees: effects of plant cover and fertilization on seedling establishment and growth

Direct seeding is a promising technique for ecological restoration, but it has been poorly studied in neotropical savannas. Different types of plant cover (no cover, crops, or green manure) and fertilization (unfertilized, synthetic fertilizer, or poultry litter) were used to verify if survival and growth of different tree species after direct seeding could be enhanced by the use of any combination of these techniques. Seedling emergence, establishment, and growth were followed for 2 years for six savanna tree species sown in an agricultural field in Central Brazil. Germination was high (52%, on average) for Anacardium occidentale, Aspidosperma macrocarpon, Hymenaea stigonocarpa, Dipteryx alata, Eugenia dysenterica, and Magonia pubescens. Six additional species were planted, but less than 5% of these seeds germinated. Crops (60% shade) did not affect seedling survival and biomass compared with the control treatment, supporting the use of this strategy during the initial phase of restoration to involve farmers in the process. In contrast, the excessive shading (95%) from the green manure treatment decreased the survival of two species and the growth in biomass and diameter of five species, especially when combined with fertilization. Seedling growth was very slow throughout the experiment, requiring extended weed management. This study supports the use of direct seeding of the studied species for savanna restoration, but methods could be improved to include a larger number of species.     

Heat shock proteins in Sorghum bicolor and Pennisetum americanum I. genotypic and developmental variation during seed germination

Abstract The capacity to synthesize heat shock proteins (HSPs) during seed germination of sorghum (Sorghum bicolor) and pearl millet (Pennisetum americanum) has been examined. HSP synthesis is detectable in a thermotolerant genotype of sorghum during the first hour of imbibition of the seed under high temperature stress. A non-coordinate control of HSP synthesis during germination was revealed. Genotypic differences were manifest in the stage of germination at which the ability to synthesize HSPs was first apparent and this related to the thermosensitivity of that genotype.     

两种结缕草种子休眠及萌发特性

通过测定兰引Ⅲ号结缕草和青岛结缕草休眠种子和解除休眠种子的吸水率、呼吸强度和脱氢酶活性等萌发生理指标,探讨了结缕草种子的休眠类型。试验结果表明,解除休眠的结缕草种子的吸水率大于未处理的种子。解除休眠种子的呼吸强度、脱氢酶活性都有较大幅度的增长,显著高于未处理。结缕草种子萌发期需30～35℃高温和充足光照。两种结缕草种子的颖壳、种皮的透性障碍及种子内存在发芽抑制物质是导致种子休眠的主要原因,属于混合休眠类型。     

贮藏条件对河西走廊四种旱生灌木种子萌发的影响

不同贮藏条件下,探讨了河西走廊枸杞(Lycium barbarum)、黑果枸杞(L.rutheni-cum)、花棒(Hedysarum scorpaium)和泡果白刺(Nitraria sphaerocarpa)4种旱生灌木种子萌发情况。结果表明:枸杞、黑果枸杞和花棒种子萌发对贮藏条件有一致性的响应,即经过低温湿润(-5℃和4℃)贮藏后有较高的萌发率、较快的萌发速度和较早的萌发开始时间,贮藏条件对泡果白刺的种子萌发(<30%)的影响较小;冬季冷屋贮藏后(-5℃,湿润)的枸杞种子萌发率(68%)显著大于冰箱贮藏(4℃,湿润)和室温干燥贮藏(<20%);冰箱和冬季冷屋贮藏后黑果枸杞种子萌发率约80%,室温干燥贮藏后萌发率仅为5%;冰箱和冬季冷屋贮藏后的花棒种子萌发率达到88%和65%,干燥贮藏后为40%。低温层积(-5℃和4℃)是打破枸杞、花棒和黑果枸杞种子休眠的有效方法之一。     

Ethylene in seed dormancy and germination

The role of ethylene in the release of primary and secondary dormancy and the germination of non-dormant seeds under normal and stressed conditions is considered. In many species, exogenous ethylene, or ethephon – an ethylene-releasing compound - stimulates seed germination that may be inhibited because of embryo or coat dormancy, adverse environmental conditions or inhibitors (e.g. abscisic acid, jasmonate). Ethylene can either act alone, or synergistically or additively with other factors. The immediate precursor of ethylene biosynthesis, 1-aminocyclopropane-1-carboxylic acid (ACC), may also improve seed germination, but usually less effectively. Dormant or non-dormant inhibited seeds have a lower ethylene production ability, and ACC and ACC oxidase activity than non-dormant, uninhibited seeds. Aminoethoxyvinyl-glycine (AVG) partially or markedly inhibits ethylene biosynthesis in dormant or non-dormant seeds, but does not affect seed germination. Ethylene binding is required in seeds of many species for dormancy release or germination under optimal or adverse conditions. There are examples where induction of seed germination by some stimulators requires ethylene action. However, the mechanism of ethylene action is almost unknown.
The evidence presented here shows that ethylene performs a relatively vital role in dormancy release and seed germination of most plant species studied.     

Does fire drive fatty acid composition in seed coats of physically dormant species?

• Seed dormancy is the key driver regulating seed germination, hence is fundamental to the seedling recruitment life-history stage and population persistence. However, despite the importance of physical dormancy (PY) in timing post-fire germination, the mechanism driving dormancy-break within seed coats remains surprisingly unclear. We suggest that seed coat chemistry may play an important role in controlling dormancy in species with PY. In particular, seed coat fatty acids (FAs) are hydrophobic, and have melting points within the range of seed dormancy-breaking temperatures. Furthermore, melting points of saturated FAs increase with increasing carbon chain length. We investigated whether fire could influence seed coat FA profiles and discuss their potential influence on dormancy mechanisms.
• Seed coat FAs of 25 species within the Faboideae, from fire-prone and fire-free ecosystems, were identified and quantified through GC–MS. Fatty acid profiles were interpreted in the context of species habitat and interspecific variation.
• Fatty acid compositions were distinct between species from fire-prone and fire-free habitats. Fire-prone species tended to have longer saturated FA chains, a lower ratio of saturated to unsaturated FA, and a slightly higher relative amount of FAs compared to fire-free species.
• The specific FA composition of seed coats of fire-prone species indicated a potential role of FAs in dormancy mechanisms. Overall, the distinct FA composition between fire-prone and fire-free species suggests that chemistry of the seed coat may be under selection pressure in fire-prone ecosystems.

     

Soil temperature and depth of legume germination during early and late dry season fires in a tropical eucalypt savanna of north‐eastern Australia

Abstract Temperatures that significantly increase seed germination of some tropical legumes (i.e. 80–100°C) were documented in the topsoil during the passage of early (May) and late (October) dry season fires in a tropical eucalypt savanna of north‐eastern Australia. Elevated temperatures penetrated at least 30 mm into the soil during the higher‐intensity, late dry season fires, but were only detected at 10 mm during the early dry season fires. The depth from which germination of two native legume forbs Galactia tenuiflora and Indigofera hirsuta occurred was positively related to the temperature elevation in the topsoil and was greater after late compared with early dry season fires. A broader range in germination depth, resulting in higher seedling densities, was recorded for I hirsuta after late dry season fires. These results suggest that seedling emergence of native leguminous forbs is likely to occur at a greater density after late rather than early dry season fires in tropical eucalypt savannas of north‐eastern Australia. Therefore, the season of burning, as a result of its relationship to fire intensity, can influence species composition through its effect on seed germination.     

火烧信号对种子萌发影响的研究进展

火作为一个基础的生态因子, 对森林、草地等陆地生态系统的结构和功能有着重要的影响。种子萌发是种子植物的重要生活史阶段, 也是火后植被更新和恢复的主要途径。植被燃烧产生了烟、热以及与烟相关的一系列火烧信号, 在打破种子休眠, 促进种子萌发方面发挥重要作用。该文将火烧信号分为物理信号和化学信号, 物理信号主要是伴随火烧产生的高温, 化学信号主要包括气态烟以及近年来从烟水中提取的影响种子萌发的关键化学物质karrikins和glyceronitrile。该文围绕火烧的基本信息, 火烧信号对种子萌发的影响, 火烧信号在实践中的应用3个方面进行系统综述, 重点探讨了不同类别的火烧信号及其交互作用对种子萌发的影响。在系统总结火烧信号对种子萌发影响的研究进展的基础上, 提出未来的研究应与烟信号作用机理的探究以及全球变化等方面相结合, 旨在充分发挥火的生态服务功能, 为火的科学管理应用和退化生态系统恢复提供理论支撑。