International principles and standards for native seeds in ecological restoration

The growing demand for native seeds in ecological restoration and rehabilitation, whether for mining, forest, or ecosystem restoration, has resulted in a major global industry in the sourcing, supply, and sale of native seeds. However, there are no international guidance documents for ensuring that native seeds have the same standards of quality assurance that are regular practice in the crop and horticultural industries. Using the International Principles and Standards for the Practice of Ecological Restoration as a foundation document, we provide for the first time a synthesis of general practices in the native seed supply chain to derive the Principles and Standards for Native Seeds in Ecological Restoration (“Standards”). These practices and the underpinning science provide the basis for developing quality measures and guidance statements that are adaptable at the local, biome, or national scale. Importantly, these Standards define what is considered native seed in ecological restoration and highlight the differences between native seeds versus seeds of improved genetics. Seed testing approaches are provided within a logical framework that outline the many different dormancy states in native seed that can confound restoration outcomes. A “pro‐forma” template for a production label is included as a practical tool that can be customized for local needs and to standardize reporting to end‐users on the level of seed quality and germinability to be expected in a native seed batch. These Standards are not intended to be mandatory; however, the guidance statements provide the foundation upon which regulatory approaches can be developed by constituencies and jurisdictions.     

含水量对种子贮藏寿命的影响

建立以收集种子为主体的基因库乃是当今保护植物种质资源最为普遍且可靠易行的方式,在世界库存约 61 0 0 0 0 0份种质资源中,近 90 %是以种子形式保存于约 1 30 0个基因库中。低温贮藏仍是目前基因库中种子种质保存的主要方法。种子含水量和贮藏温度是影响种子在贮藏期间生活力和活力保持的关键因素。传统的经验认为控制温度比控制水分来得安全有效,因而趋向于向低温或超低温的贮藏方向发展。国际植物遗传资源研究所（ＩＰＧＲＩ）曾推荐 5%～ 6%的含水量和 - 1 8～- 2 0℃低温作为各国长期保存种子的理想条件。目前,世界各国都把更多的…     

Lack of adequate seed supply is a major bottleneck for effective ecosystem restoration in Chile: friendly amendment to Bannister et al. (2018)

We argue that the need for a quality seed supply chain is a major bottleneck for the restoration of Chile's native ecosystems, thus supplementing the list of bottlenecks proposed by Bannister et al. in 2018. Specifically, there is a need for defining seed transfer zones, developing standards and capacities for properly collecting and storing seeds, reducing information gaps on seed physiology and longevity, and implementing an efficient seed supply chain with certification of seed origin and quality. Without such capacities, countries are unlikely to meet their restoration commitments. Although we focus on bottlenecks in Chile, the issues we raise are relevant to other countries and thus the global agenda for ecological restoration.     

Latitude, seed predation and seed mass

Aim We set out to test the hypothesis that rates of pre‐ and post‐dispersal seed predation would be higher towards the tropics, across a broad range of species from around the world. We also aimed to quantify the slope and predictive power of the relationship between seed mass and latitude both within and across species. Methods Seed mass, pre‐dispersal seed predation and post‐dispersal seed removal data were compiled from the literature. Wherever possible, these data were combined with information regarding the latitude at which the data were collected. Analyses were performed using both cross‐species and phylogenetic regressions. Results Contrary to expectations, we found no significant relationship between seed predation and latitude (log₁₀ proportion of seeds surviving predispersal seed predation vs. latitude, P = 0.63; R² = 0.02; n = 122 species: log₁₀ proportion of seeds remaining after postdispersal seed removal vs. latitude, P = 0.54; R² = 0.02; n = 205 species). These relationships remained non‐significant after variation because of seed mass was accounted for. We also found a very substantial (R² = 0.21) relationship between seed mass and latitude across 2706 species, with seed mass being significantly higher towards the tropics. Within‐species seed mass decline with latitude was significant, but only about two‐sevenths, as rapid as the cross‐species decline with latitude. Results of phylogenetic analyses were very similar to cross‐species analyses. We also demonstrated a positive relationship between seed mass and development time across ten species from dry sclerophyll woodland in Sydney (P < 0.001; R² = 0.77; Standardized Major Axis slope = 0.14). These data lend support to the hypothesis that growing period might affect the maximum attainable seed mass in a given environment. Main conclusions There was no evidence that seed predation is higher towards the tropics. The strong relationship between seed mass and latitude shown here had been observed in previous studies, but had not previously been quantified at a global scale. There was a tenfold reduction in mean seed mass for every c. 23° moved towards the poles, despite a wide range of seed mass within each latitude.     

Seed survival of the tropical tree Cryptocarya aschersoniana (Lauraceae): Consequences of habitat disturbance

Seed survival in soil could be strongly influenced by habitat characteristics, but little is known about the behaviour of seeds sensitive to desiccation in seed banks installed in natural or disturbed habitats. Cryptocarya aschersoniana seeds disperse at the end of the rainy season but do not germinate immediately; thus, they may form seed banks in soil. This study evaluated the behaviour of C. aschersoniana seed banks induced in the natural environment of the species and in a disturbed area. Recently harvested C. aschersoniana germination units were characterized according to their water content, germination and viability. In 2011 and 2012, seed banks were established by burying samples of seeds in the understory of a semi‐deciduous forest. In 2012, samples were also buried in a disturbed area. The seed banks were sampled at certain time intervals, and the samples were characterized as described above. Precipitation and air temperature data were collected. As a result, seeds in the seed bank established in the natural environment form a transient seed bank and showed the same behaviour in both years studied. A germination peak was observed starting 210 days after burial (coinciding with the onset of the rainy season) and reached germination percentages higher than 80% at the end of the experiment for both years. Seed mortality did not exceeded 28% in the natural environment. However, in the disturbed environment, the seeds lost their viability more rapidly, with 90% of the seeds becoming unviable 240 days after burial. Germinated seeds in the disturbed environment (maximum 21%) were not able to establish seedlings. These results underscore the importance of maintaining a natural, undisturbed forest for the conservation of this species.     

Seed storage at elevated partial pressure of oxygen, a fast method for analysing seed ageing under dry conditions

Background and Aims

Despite differences in physiology between dry and relative moist seeds, seed ageing tests most often use a temperature and seed moisture level that are higher than during dry storage used in commercial practice and gene banks. This study aimed to test whether seed ageing under dry conditions can be accelerated by storing under high-pressure oxygen.

Methods

Dry barley (Hordeum vulgare), cabbage (Brassica oleracea), lettuce (Lactuca sativa) and soybean (Glycine max) seeds were stored between 2 and 7 weeks in steel tanks under 18 MPa partial pressure of oxygen. Storage under high-pressure nitrogen gas or under ambient air pressure served as controls. The method was compared with storage at 45 °C after equilibration at 85 % relative humidity and long-term storage at the laboratory bench. Germination behaviour, seedling morphology and tocopherol levels were assessed.

Key Results

The ageing of the dry seeds was indeed accelerated by storing under high-pressure oxygen. The morphological ageing symptoms of the stored seeds resembled those observed after ageing under long-term dry storage conditions. Barley appeared more tolerant of this storage treatment compared with lettuce and soybean. Less-mature harvested cabbage seeds were more sensitive, as was the case for primed compared with non-primed lettuce seeds. Under high-pressure oxygen storage the tocopherol levels of dry seeds decreased, in a linear way with the decline in seed germination, but remained unchanged in seeds deteriorated during storage at 45 °C after equilibration at 85 % RH.

Conclusions

Seed storage under high-pressure oxygen offers a novel and relatively fast method to study the physiology and biochemistry of seed ageing at different seed moisture levels and temperatures, including those that are representative of the dry storage conditions as used in gene banks and commercial practice.     

Seed storage: maintaining seed viability and vigor for restoration use

Effective seed storage after sourcing (harvesting or purchasing) is critical to restoration practitioners and native seed producers, as it is key to maintaining seed viability. Inadequate seed storage can lead to a waste of both natural and economic resources when seeds of poor quality are sown. When working with native species with unknown storage behavior, general assumptions can be made based on studies on related species, and standard practices may be applied with caution; however, an investigation should be conducted to understand if specific storage requirements are needed and for how long seeds can be stored before they lose significant viability. In this paper of the Special Issue Standards for Native Seeds in Ecological Restoration, we provide an overview of the key concepts in seed storage and the steps to take for effective storage of native seeds for restoration use.     

A comparative study of seed yield parameters in Arabidopsis thaliana mutants and transgenics

Because seed yield is the major factor determining the commercial success of grain crop cultivars, there is a large interest to obtain more understanding of the genetic factors underlying this trait. Despite many studies, mainly in the model plant Arabidopsis thaliana, have reported transgenes and mutants with effects on seed number and/or seed size, knowledge about seed yield parameters remains fragmented. This study investigated the effect of 46 genes, either in gain- and/or loss-of-function situations, with a total of 64 Arabidopsis lines being examined for seed phenotypes such as seed size, seed number per silique, number of inflorescences, number of branches on the main inflorescence and number of siliques. Sixteen of the 46 genes, examined in 14 Arabidopsis lines, were reported earlier to directly affect in seed size and/or seed number or to indirectly affect seed yield by their involvement in biomass production. Other genes involved in vegetative growth, flower or inflorescence development or cell division were hypothesized to potentially affect the final seed size and seed number. Analysis of this comprehensive data set shows that of the 14 lines previously described to be affected in seed size or seed number, only nine showed a comparable effect. Overall, this study provides the community with a useful resource for identifying genes with effects on seed yield and candidate genes underlying seed QTL. In addition, this study highlights the need for more thorough analysis of genes affecting seed yield.     

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。     

Seed dynamics of resprouting shrubs in grassy woodlands: Seed rain,predators and seed loss constrain recruitment potential

Abstract Measuring the fate of seeds between seed production and seedling establishment is critical in understanding mechanisms of recruitment limitation of plants. We examined seed fates to better understand the recruitment dynamics of four resprouting shrubs from two families (Fabaceae and Epacridaceae) in temperate grassy woodlands. We tested whether: (i) pre‐dispersal seed predation affected seed rain; (ii) post‐dispersal seed predation limited seed bank accumulation; (iii) the size of the seed bank was related to seed size; and (iv) viable seeds accumulated in the soil after seed rain. There was a distinct difference in seed production per plant between plant families with the legumes producing significantly more seeds per individual than the epacrids. Seed viability ranged from 43% to 81% and all viable had seed or fruit coat dormancy broken by heat or scarification. Pre‐dispersal predation by Lepidopteran larvae removed a large proportion of seed from the legume seed rain but not the epacrids. Four species of ants (Notoncus ectatomoides, Pheidole sp., Rhytidoponera tasmaniensis and Iridomyrmex purpureus) were major post‐dispersal seed removers. Overall, a greater percentage of Hardenbergia (38%) and Pultenaea (59%) seeds were removed than the fleshy fruits of Lissanthe (14%) or Melichrus (0%). Seed bank sizes were small (<15 seeds m^?2) relative to the seed rain and no significant accumulation of seed in the soil was detected. Lack of accumulation was attributed to seed predation as seed decay was considered unlikely and no seed germination was observed in our study sites. Our study suggests that seed predation is a key factor contributing to seed‐limited recruitment in grassy woodland shrubs by reducing the number of seeds stored in the soil.     

The ecophysiology of seed persistence: a mechanistic view of the journey to germination or demise

Seed persistence is the survival of seeds in the environment once they have reached maturity. Seed persistence allows a species, population or genotype to survive long after the death of parent plants, thus distributing genetic diversity through time. The ability to predict seed persistence accurately is critical to inform long‐term weed management and flora rehabilitation programs, as well as to allow a greater understanding of plant community dynamics. Indeed, each of the 420000 seed‐bearing plant species has a unique set of seed characteristics that determine its propensity to develop a persistent soil seed bank. The duration of seed persistence varies among species and populations, and depends on the physical and physiological characteristics of seeds and how they are affected by the biotic and abiotic environment. An integrated understanding of the ecophysiological mechanisms of seed persistence is essential if we are to improve our ability to predict how long seeds can survive in soils, both now and under future climatic conditions. In this review we present an holistic overview of the seed, species, climate, soil, and other site factors that contribute mechanistically to seed persistence, incorporating physiological, biochemical and ecological perspectives. We focus on current knowledge of the seed and species traits that influence seed longevity under ex situ controlled storage conditions, and explore how this inherent longevity is moderated by changeable biotic and abiotic conditions in situ, both before and after seeds are dispersed. We argue that the persistence of a given seed population in any environment depends on its resistance to exiting the seed bank via germination or death, and on its exposure to environmental conditions that are conducive to those fates. By synthesising knowledge of how the environment affects seeds to determine when and how they leave the soil seed bank into a resistance–exposure model, we provide a new framework for developing experimental and modelling approaches to predict how long seeds will persist in a range of environments.     

Comparing methods of weed seed exposure to predators

Several methods of seed exposure are used in seed predation studies, but how these methods influence the results remains poorly studied. In this article, two commonly used methods of seed exposure – seed cards and plasticine trays – were compared in the field and in the laboratory using three species of weed seeds. In the field, cards or trays with seeds were exposed either with or without cages to keep vertebrates out and either with or without impermeable roofs to provide protection from the weather. The overall seed retrieval from the control stands varied significantly between the methods of seed exposure, roof treatment and the species of seeds, and the scatter in the seed retrieval increased or decreased monotonically with the temperature or precipitation for some species of seeds. This indicates that the controls were more or less relevant depending on the weather conditions and species. The seed removal from cards varied between the species of seeds. The effect of exclosure cages indicated that invertebrates were the dominant seed predators of Capsella bursa‐pastoris, while in Poa annua and Stellaria media both vertebrates and invertebrates were important. Higher seed removal from plasticine trays compared to seed cards was found for all three species of seeds, and placing roofs over the seeds affected seed removal in C. bursa‐pastoris and P. annua. In the laboratory, seed consumption varied with the method of exposure in 10 out of 12 combinations of seed species and predator (two species of carabids and two species of isopods). The overall tendency was the opposite of the field observations: seed consumption was higher for seeds on filter paper and cards compared to seeds on tin trays. We conclude that seed cards are more convenient for short term studies in the field, while tin trays are useful in laboratory multi‐choice experiments. To measure the realistic consumption of invertebrate predators in the laboratory, filter paper seems to be the best option as it does not present an obstacle to predators eating the seed. Using roofs introduced further bias regarding the estimates of seed removal in the field and thus should be avoided.     

香果树(Emmenopterys henryi)种群种子雨、种子库及 实生苗数量的海拔梯度变化

香果树为我国Ⅱ级野生保护植物,由于原生境中香果树种群的数量迅速减少,目前已濒临灭绝,种群亟待恢复。研究了武夷山不同海拔的香果树种群种子雨、种子库及其幼苗存活现状,目的是阐明不同海拔地区的香果树种群自然更新的动态过程,寻找其更新脆弱的环节及其影响因素,为香果树自然种群的恢复提供理论依据。以分布于不同海拔高度（819、980、1 140及1 301 m）的香果树母树为中心,在其周围（东、南、西、北四个方位）布设种子雨收集框和土壤种子库样方,连续观测种子雨和种子库的动态,并对不同地表覆盖物中种子萌发及幼苗数量进行观测和记录。结果表明,武夷山香果树种群种子雨持续时间近2个月,尤其是11月底至12月中旬为种子雨高峰期;随着海拔的上升,香果树种子总密度以及虫蛀种子密度均显著降低,但千粒重以及饱满种子密度则有所增加;香果树土壤种子库为瞬时种子库,其水平分布以母树南侧和西侧种子密度较大,母树北侧种子密度最小。香果树种子的垂直分布主要集中于枯落物和苔藓层;3月时香果树土壤种子库中存留的饱满种子仅占种子散布后1.80%,而1年实生苗存活率为6.18%,土壤种子库的损耗是其种群自然更新困难的主要原因;不同地表覆盖物对香果树幼苗的存活产生显著影响,枯落物及苔藓层的幼苗死亡率显著高于土壤表面。建议清理林内枯落物及苔藓,增加林内裸露土壤的面积,减小地上草本的盖度,以减少种子霉烂、虫蛀的几率,增加香果树实生苗胚根与土壤接触的几率以及改善光照条件,促进香果树自然更新。     

壳斗科三种植物种子大小对昆虫寄生及种子存活率的影响

种子内的寄生昆虫可以严重影响种子的发育、损害种子活力。种子足余策略理论认为大种子有利于抵御和适应昆虫寄生取食,但动物最优觅食理论推测,大种子更易遭受昆虫寄生。为对这两种对立观点进行验证,本实验以青冈、苦槠和麻栎各2个种群的种子为材料,对昆虫寄生与完好种子间的体积和萌发率进行比较,并对寄生种子萌发率与种子体积的关系进行了分析。结果显示:(1)在6个种群的种子中,只有松阳麻栎和青冈种群的寄生种子体积大于完好种子,其余4个种群的寄生种子体积小于完好种子,但这种差异不显著;(2)所有寄生种子的整体萌发率(18%)显著低于完好种子(45.66%)(P<0.001),在不同种群内,寄生种子的萌发率也分别显著低于完好种子。(3)比较同种植物体积差异显著的寄生种子的萌发率发现,大种子总比小种子具有更高的萌发率,但差异不显著;在不同植物的寄生种子间比较时,体积最大的麻栎种子萌发率显著高于体积较小的青冈和苦槠种子。研究结果表明,象虫在种子上产卵时对大种子没有选择偏好,在昆虫寄生取食严重损害种子活力的压力下,大种子比小种子具有更强的耐受力。     

Foreword: International Standards for Native Seeds in Ecological Restoration

Restoration practitioners must increasingly incorporate seed procurement models and seed use planning early in project development, despite insufficient guidance about what are reasonable expectations for the sourcing and use of native seeds. This special issue presents a series of articles examining each key step in the native seed supply chain, and provides a framework for the “standards” that need to be applied to native seed batches if the native seed supply chain is to achieve the levels of reliability and transparency required. These Standards provide seed buyers, end users, and funding bodies with a level of confidence and reliability in the sourcing of quality native seeds, and a pathway toward global best practice in native seed use.     

江西三清山濒危植物天女花种子特性研究

研究了江西三清山天女花种子的特性,结果表明：(1)三清山天女花种群种子雨自10月初开始,10月末结束,10月中旬强度达高峰,玉京峰地段天女花种子雨持续时间较短;(2)产种率受母树冠层影响,即上层>中层>下层;种子败育率高(49.4%～100.0%);(3)主成分分析表明,乔木层盖度、大气湿度、光照因子以及土壤含水量对天女花种子发育产生显著影响。     

Spatial distribution and population dynamics of Striga hermonthica seeds in naturally infested farm soils

Densities and spatial distribution in soil of seeds of Striga hermonthica were analysed for four naturally infested farm fields in Western Kenya. A revised method for extraction of Striga seeds from soil was used, combining centrifugation with existing techniques based on flotation. Tests showed that 85% of Striga seeds were retrieved from soil samples. In all fields the majority of seeds were found in the plough layer (0 – 20 cm). New seeds entering the soil from the surface after seed shedding created a strong gradient with depth. Downward penetration from the soil surface was larger in sandy soil than in clay soil. In tilled soils no significant vertical density gradient was found within the plough layer. At a fine scale (0.2 m) seed densities showed little horizontal variation, but significant differences in seed densities in the horizontal plane were found at larger scale distances (several m) between locations in all fields. At 125 days after sowing the estimated average number of seeds produced per emerged Striga shoot was 4,827, excluding an approximately similar amount of seeds present in maturing capsules. The estimated average number of seeds produced per mature Striga seed capsule was 1188. Large seasonal fluctuations in the Striga seedbank were measured. An average net increase of 88,825 Striga seeds m^-2 (equivalent to 340%) was calculated from seedbank analyses in 16 sorghum plots. The level of Striga infestation in one field had decreased by 62% from 34,250 seeds m^-2 to 13,125 seeds m^-2 after keeping it fallow for a year. A sharp decline in Striga seed density was found in samples taken at increasing distances from highly infested fields, irrespective of wind direction or slope, suggesting very limited dispersal of Striga seeds by wind or water. Parasite emergence was non-linearly related to initial Striga seed densities in the soil, but this relationship was only observable at the scale of individual plant holes. Seed production was also non-linearly related to numbers of emerged parasites, when measured at plot scale (25 m²), but not at the scale of individual plant holes. In the fields we studied, seed densities below levels of 13,000 Striga seeds m^-2 could be considered to suppress the number of emerging parasites. However, if two or three emerged Striga plants per m^-2 were left to seed, enough seeds would be produced to keep the seedbank in balance.     

Prediction of desiccation sensitivity in seeds of woody species: a probabilistic model based on two seed traits and 104 species

BACKGROUND AND AIMS: Seed desiccation sensitivity limits the ex situ conservation of up to 47 % of plant species, dependent on habitat. Whilst desirable, empirically determining desiccation tolerance levels in seeds of all species is unrealistic. A probabilistic model for the rapid identification of woody species at high risk of displaying seed desiccation sensitivity is presented. METHODS: The model was developed using binary logistic regression on seed trait data [seed mass, moisture content, seed coat ratio (SCR) and rainfall in the month of seed dispersal] for 104 species from 37 families from a semi-deciduous tropical forest in Panamá. KEY RESULTS: For the Panamanian species, only seed mass and SCR were significantly related to the response to desiccation, with the desiccation-sensitive seeds being large and having a relatively low SCR (i.e. thin 'seed' coats). Application of this model to a further 38 species, of known seed storage behaviour, from two additional continents and differing vegetation types (dryland Africa and temperate Europe) correctly predicted the response to desiccation in all cases, and resolved conflicting published data for two species (Acer pseudoplatanus and Azadirachta indica). CONCLUSIONS: This model may have application as a decision-making tool in the handling of species of unknown seed storage behaviour in species from three disparate habitats.     

植冠与土壤种子库储存种子的萌发特性及策略

为明确黄土丘陵沟壑区植物种子库如何调控种子萌发来提高个体适合度,选择研究区7种具有种子库的主要物种为研究对象,以刚成熟和室内储存种子为对照,比较植冠宿存(5个宿存期)和土壤埋藏(5a埋藏期)对植物种子萌发特性的影响,探讨植冠种子库与土壤种子库储存下的种子萌发策略。结果表明:7种植物种子经过不同种子库储存后萌发特性表现出明显的种间差异,黄刺玫(Rosa xanthina)和水栒子(Cotoneaster multiflorus)种子萌发力表现为植冠宿存不变型、土壤储存增强型,土壤储存明显提高水栒子种子萌发速率;达乌里胡枝子(Lespedeza davurica)和狼牙刺(Sophora davidii)种子萌发力表现为植冠宿存增强型、土壤储存减弱型,种子萌发历时表现为植冠宿存延长型,土壤种子库储存还可加快达乌里胡枝子萌发速率、缩短萌发历时;茭蒿(Artemisia giralaii)和铁杆蒿(Artemisia gmelinii),种子萌发率随植冠宿存时间先升高后降低,随土壤储存时间先降低后升高,土壤储存可推迟其萌发,铁杆蒿种子萌发速率在植冠与土壤储存后均加快;紫丁香(Syringa oblata)种子萌发率随植冠宿存先升高后降低,土壤储存明显加快其种子启动萌发与速率。在黄土丘陵沟壑区,植物种子经过植冠或土壤种子库储存,或增加、加快、提早萌发充分利用有利条件提高占据性,或减少、减缓、推迟萌发分摊不利条件的风险;而且该区植物植冠与土壤储存后种子萌发特性间的关系,体现各自适应环境与应对干扰的分工与协作策略,主要表现为:单一主导型和相辅相成型。