Seed dormancy and soil seedbank of the invasive weed Chenopodium hybridum in north‐western China

Seed dormancy and persistence in the soil seedbank play a key role in timing of germination and seedling emergence of weeds; thus, knowledge of these traits is required for effective weed management. We investigated seed dormancy and seed persistence on/in soil of Chenopodium hybridum, an annual invasive weed in north‐western China. Fresh seeds are physiologically dormant. Sulphuric acid scarification, mechanical scarification and cold stratification significantly increased germination percentages, whereas dry storage and treatments with plant growth regulators or nitrate had no effect. Dormancy was alleviated by piercing the seed coat but not the pericarp. Pre‐treatment of seeds collected in 2012 and 2013 with sulphuric acid for 30 min increased germination from 0% to 66% and 62% respectively. Effect of cold stratification on seed germination varied with soil moisture content (MC) and duration of treatment; seeds stratified in soil with 12% MC for 2 months germinated to 39%. Burial duration, burial depth and their interaction had significant effects on seed dormancy and seed viability. Dormancy in fresh seeds was released from October to February, and seeds re‐entered dormancy in April. Seed viability decreased with time for seeds on the soil surface and for those buried at a depth of 5 cm, and 39% and 10%, respectively, were viable after 22 months. Thus, C. hybridum can form at least a short‐lived persistent soil seedbank.     

Effect of soil moisture during stratification on dormancy release in seeds of five common weed species

Although the effects of cold stratification on the release of physiological dormancy in seeds have been studied extensively, knowledge of the role of soil moisture content on seed dormancy release during cold stratification is limited. Our study determined seed dormancy characteristics and the effect of soil moisture content on seed dormancy breakage during cold stratification in the five common weed species Amaranthus retroflexus, Chenopodium album, Chenopodium hybridum, Plantago lanceolata and Setaria glauca. Seeds of all five species were dormant at the time of harvest and their germination response to light and temperature varied. Soil moisture content had a significant effect on seed dormancy release of all species except P. lanceolata. Germination percentage of A. retroflexus, C. album, C. hybridum increased and then decreased as soil moisture content increased, regardless of germination test temperature. The optimal soil moisture content and seed moisture content for dormancy breakage of A. retroflexus, C. album, C. hybridum were 8%, 12%, 8% and 22.0%, 37.7%, 25.7% respectively. Dry storage (after‐ripening) significantly increased germination of S. glauca. Moreover, increasing soil moisture content first slowed and then increased dormancy breakage in S. glauca. These results suggest that data on soil moisture content should be incorporated into models that predict weed seed dormancy breakage and timing of seedling emergence as well as those for weed management.     

Germination rates of weedy radish populations (Raphanus spp.) altered by crop‐wild hybridisation,not human‐mediated changes to soil moisture

Cultivated plants are known to readily hybridise with their wild relatives, sometimes forming populations with weedier life‐history strategies than their progenitors. Due to altered precipitation patterns from human‐induced global climate change, crop‐wild hybrid populations may have new and unpredictable environmental tolerances relative to parental populations, which would further challenge farming and land‐management weed control strategies. To recognise the role of seed dormancy variation in weed invasion, we compared seedbank dynamics of two cross‐type populations (wild radish, Raphanus raphanistrum, and crop‐wild hybrid radish, R. raphanistrum × R. sativus) across a soil moisture gradient. In a seed‐burial experiment, we assessed relative rates of seed germination, dormancy and seed mortality over two years across cross types (crop‐wild hybrid or wild) and watering treatments (where water was withheld, equal to annual rainfall, or double annual rainfall). Weekly population censuses in 2012 and 2013 assessed the frequency and timing of seedling emergence within a growing season. Generally, germination rates were two times higher and seed dormancy was 58% lower in hybrid versus wild populations. Surprisingly, experimental soil moisture conditions did not determine seedbank dynamics over time. Yet, seed bank dynamics changed between years, potentially related to different amounts of annual rainfall. Thus, variation in seedbank dynamics may be driven by crop‐wild hybridisation rates and, potentially, annual variation in soil moisture conditions.     

Thermal and hormonal regulation of the dormancy–germination transition in Amaranthus tuberculatus seeds

The transition from seed dormancy to germination is a multi‐step process. However, distinguishing between physiological processes involved in seed dormancy alleviation and those involved in germination has been difficult. We studied the seed dormancy alleviation process in Amaranthus tuberculatus, an important weed species in midwestern USA. Using three A. tuberculatus biotypes that differ in dormancy level, it was determined that stratification reduced seed dormancy from a high to a low level. Temperature alternation alleviated low seed dormancy and triggered germination. Exogenously applied abscisic acid (ABA) and gibberellic acid (GA) had no effect on seeds with high dormancy. However, ABA and paclobutrazol (a GA biosynthesis inhibitor) significantly reduced germination of seeds with low dormancy. Hormones could not replace the effects of stratification or temperature alternation on dormancy alleviation. Based on our results, we propose a seed dormancy–germination transition model in which the dormancy of A. tuberculatus seeds is progressively reduced from a high to a low level; but environmental conditions (i.e. stratification) can accelerate the dormancy alleviation process. Under low dormancy levels, the seed is more sensitive to environmental cues that are responsible for removing dormancy and triggering germination (i.e. temperature alternation). Finally, ABA and GA regulation occurs primarily during the final transition from low dormancy to germination rather than the alleviation of high dormancy.     

Seed dormancy and germination in the summer annual Galeopsis speciosa

This study examined germination and dormancy in Galeopsis speciosa (Lamiaceae), a common summer annual weed in cold‐temperate areas. Seeds collected in southern Sweden were subjected to several experiments. The seeds were dormant at maturity. Seeds sown outdoors after collection produced a small number of seedlings that emerged early in the spring. After long cold stratification or stratification outdoors over two winters, the maximum germination was 40–50%; germination occurring over a wide range of temperatures. Warm stratification preceding cold stratification had no effect on germination, but repeated warm and cold periods seemed to promote germination. Gibberellic acid (GA) stimulated germination, but full germination was only achieved after more than 2 months of incubation at the most suitable temperature regime tested. Excised embryos grew and developed into normal seedlings. With these results, the species does not fit into the currently used system for seed dormancy classifications. The response to GA and the growth of excised embryos indicate non‐deep or intermediate physiological dormancy, but dormancy alleviation by stratification was not in line with the guiding principles for these classifications. Galeopsis speciosa has a strong dormancy that is sufficiently alleviated during the winter to allow germination of only part of a seed batch each year; hence a stepwise germination pattern occurs over a period of several years.     

Factors affecting the seed germination and seedling emergence of muskweed (Myagrum perfoliatum)

Myagrum perfoliatum is a noxious broad‐leaved weed in western Iranian farming systems. A better understanding of the timing of seedling emergence would facilitate the development of better control strategies for this weed. Therefore, the objective of this study was to examine the effects of different factors on muskweed seed germination. Only 2.8% of the seeds of this species, which are encapsulated in siliques, germinated by, while the seeds that had been removed from the siliques had a 50% germination rate. The immersion of muskweed fruits in concentrated sulfuric acid for 110 min was the best treatment for promoting germination. Gibberellic acid stimulated the germination of the naked seeds by 29.1%, potassium nitrate (40 mmol L^‐1) increased the germination rate to 71%, while higher concentrations of potassium nitrate inhibited germination. The optimum germination temperature for the naked seeds was 20/10°C (day/night) and light was not required for germination. No seedling emerged when the seeds were buried 6 cm deep. The seeds were sensitive to both osmotic and salinity stress, but they germinated to 46–49% over a pH range of 4–10. The results of this study revealed that the seeds of M . perfoliatum have physiological dormancy and that it is slowly broken via after‐ripening. However, the fruit wall can prevent germination after physiological dormancy is broken. Thus, this species has the potential to form a persistent seed bank because of the presence of the fruit wall.     

Modelling Chloris virgata germination and emergence under different temperature and light quality conditions

Chloris virgata is a problematic weed around the world. Prediction of weed germination rates could be a useful strategy to optimise timing of weed control actions. We studied the germination and emergence of C. virgata collected seeds under different after-ripening treatments and different exhumation dates after seed dispersal, to estimate seed dormancy level and predict weed emergence dynamics under field conditions. Three experiments were conducted under controlled conditions to determine base, optimum and maximum germination temperatures (T_b, T_o and T_m respectively) and comprised: (a) exposure of seeds to gradually increasing and decreasing temperatures between 5 and 35°C; (b) exposure of seeds to different constant temperatures; and (c) exposure of seeds to different light quality conditions (red – far red ratio) and temperature regimes (constant and alternating temperatures). To explore genuine environmental conditions, a field experiment was performed to determine weed emergence under different shading levels. Finally, with the data obtained, a thermal time model for dormancy release was used to predict C. virgata seedling emergence in the Argentine Pampas region. Seeds after-ripened in cold and wet conditions and constant 25°C showed the highest germination percentages. The values of T_b (7°C), T_o (28°C) and T_m (40°C) remained constant at all exhumation dates. Neither light quality nor thermal regime modified the final germination percentages. However, shading delayed seedling emergence under field conditions, even when it was adjusted by thermal time. These results may allow predicting C. virgata emergence in temperate regions and help to improve weed control in integrated weed management strategies.     

Dormancy,germination and emergence of Urochloa panicoides regulated by temperature

Urochloa panicoides is an annual weed of summer crops. In Argentina, in subhumid areas with monsoon rainfall, it germinates and establishes in a single flush. To (i) identify the environmental factors that modify its seed dormancy level and germination and (ii) quantify the parameters describing the thermal behaviour of the germination and emergence dynamics of this weed under non‐limiting water conditions, we established a set of germination experiments performed (i) under controlled conditions using seeds after ripened for 3 or 6 months in different thermal and hydric conditions and (ii) under field conditions, where the soil temperature was modified by applying different shading levels. Seed dormancy level remained high with 3 months after ripening in all treatments. After 6 months, seeds stored at 4°C in dry conditions did not germinate at any temperature, while seeds stored at 25°C in dry conditions and in situ germinated c. 20% and 60% respectively. Germination percentage was higher in seeds harvested before their natural dispersal. The base, optimum and maximum temperatures for seed germination were 6, 35 and 45°C respectively. Shading reduced the number of emerged seedlings, possibly by reducing the soil thermal amplitude. The results explained the dormancy‐breaking mechanism of U. panicoides that allows a high germination rate in the field when rainfall occurs.     

Seasonal changes in seed dormancy of Solanum nigrum and Solanum physalifolium

The seed dormancy cycle in Solanum nigrum and Solanum physalifolium was studied in relation to seasonal temperature. Seed lots of both species were buried in pots outdoors in a randomised complete block design with four replicates from November 2004 to November 2006. At regular intervals, samples of the seeds were randomly exhumed and tested for germination in incubators at three temperatures and light/darkness regimes. For both species, low winter temperature weakened dormancy and high temperature strengthened it. Dormancy induction mainly occurred from August to October in both species after experiencing warm temperatures. An exception from the general pattern of seed dormancy was however observed; seed germination percentages were temporarily reduced in early spring, followed by a peak in germination, before the main period of strong dormancy in S. nigrum . The same phenomenon was observed in S. physalifolium during June in the first year. This short-lived dormancy induction might explain the late emergence of the species. Seed dormancy enables the species to maximise its chance of survival by regulating germination timing to favourable conditions. Therefore, information on the dormancy cycle can be used to predict seedling emergence and optimise weed control operations.     

Variation in seed dormancy and light sensitivity in Alopecurus myosuroides and Apera spica-venti

Variation in seed dormancy and light sensitivity was studied in Alopecurus myosuroides and Apera spica-venti . Seeds were collected from different populations, at different dates and from plants emerging in autumn or spring, and used in four experiments. In the first experiment, initial dormancy was investigated in light and darkness. In Expt 2, buried seeds were exhumed on 16 occasions, from September 1997 to March 2000, and germinated in light, in darkness and after a 5-s light exposure. In Expt 3, emergence was recorded for seeds sown in pots outdoors. In Expt 4, stratified seeds of A. myosuroides only were exposed to photon irradiance ranging from 0.1 to 25 600 μmol m⁻². Variation was high among seed collections, but both species showed winter annual dormancy patterns. Apera spica-venti germinated to high percentages in autumn but negligibly in spring. Alopecurus myosuroides germinated less in spring when tested in darkness and after a short light exposure and emerged poorly in spring, which reflected photo-desensitisation during cold stratification. We conclude that the peak of emergence in A. myosuroides , and to some extent in A. spica-venti , is largely regulated by exposure to light interacting with low-level dormancy. This offers valuable information regarding optimal timing of weed control measures.     

Germination characteristics and emergence time of annual Bromus species of differing weediness in Sweden

The germination ecology of four annual Bromus species, which differ in weediness on arable land in southern Sweden, was investigated. The most problematic species is Bromus sterilis , while Bromus hordeaceus frequently occurs on arable land. In contrast, Bromus arvensis is a rare weed, and Bromus tectorum is found infrequently in fields despite being a widespread ruderal species. Five experiments were conducted to identify germination characteristics that could explain differences in habitat and abundance: (i) intraspecific variation in dormancy level; (ii) germination response to different light conditions; (iii) light and temperature interactions at germination; (iv) timing of seedling emergence; and (v) seed persistence in soil. Bromus sterilis and B. tectorum behaved similarly in all tests. For both these species, there were large differences in dormancy level among populations and strong inhibition of germination by light. In addition, emergence from seeds sown on the soil surface was both delayed and reduced compared with buried seeds. In contrast, B. hordeaceus and B. arvensis showed generally weak dormancy, and germination was only slightly inhibited by light. It was concluded that germination characteristics alone do not explain the differences in weediness between these four species.     

不同贮藏条件下龙葵种子的季节性休眠特性

龙葵是农田恶性杂草?为明确龙葵种子休眠与季节温度的关系, 研究了室内和室外不同贮藏条件下其种子萌发对温度的响应规律?结果表明, 室外贮藏条件下的龙葵种子萌发呈季节性变化, 从10月到翌年5月, 龙葵种子萌发率均在95%以上, 随着夏季温度的升高, 萌发率从6月开始下降, 9月达到最低值(25.4%), 由此进行年际间休眠和非休眠周期的循环?夏季6月-9月的高温可诱导龙葵种子进入休眠状态, 而秋冬季的相对低温有利于解除种子休眠, 使种子恢复萌发状态?龙葵种子休眠和非休眠状态之间的切换受季节性温度变化的影响?室内贮藏的种子, 由于环境温度较为稳定, 其萌发率年际变化较小, 在20%~50%之间?本文明确了龙葵种子休眠的周期性变化规律, 有助于精准预测其出苗时间, 研究结果可为阐明龙葵种子休眠萌发机制和制订基于萌发调控的绿色防控策略提供依据?     

Mycelial growth rate and toxin production in the seed pathogen Pyrenophora semeniperda: resource trade‐offs and temporally varying selection

Pyrenophora semeniperda, an important pathogen in Bromus tectorum seed banks in semi‐arid western North America, exhibits >4‐fold variation in mycelial growth rate. Host seeds exhibit seasonal changes in dormancy that affect the risk of pathogen‐caused mortality. The hypothesis tested is that contrasting seed dormancy phenotypes select for contrasting strategies for increasing pathogen fitness, and that increased fitness on nondormant seeds involves a resource trade‐off between toxin production and growth. The strategy for successfully attacking rapidly germinating nondormant seeds at high inoculum loads in autumn involves increased post‐infection aggressiveness to prevent seed escape through germination. An earlier study demonstrated that slow‐growing strains caused higher mortality than faster‐growing strains on nondormant host seeds at high inoculum loads. In this study, production of the toxin cytochalasin B was significantly higher in slower‐growing strains, and was induced only in seeds or in seed‐constituent‐containing media. Its production was reduced in vivo by Bromus tectorum seeds, suggesting direct involvement in pathogenesis on seeds. Fast‐growing strains caused significantly higher mortality than slow‐growing strains at low inoculum loads on dormant seeds, which apparently have resistance that is overcome at high loads or through rapid mycelial proliferation. In a co‐inoculation study, the fast‐growing isolate produced 3 × more stromata than the slow‐growing isolate on dormant seeds, whereas the slow‐growing isolate was twice as successful on nondormant seeds. These results provide evidence that mycelial growth rate variation and associated variation in cytochalasin B production represent a trade‐off maintained through temporally varying selection resulting from seasonal variation in host seed dormancy status.     

Overcoming seed dormancy in oilseed rape (Brassica napus L.) with exogenous compounds

Dormant seeds of oilseed rape (OSR) can persist in the soil and cause OSR volunteers in subsequent crops. Several approaches were tested in the laboratory and in the field to determine whether dormancy induction and seed persistence can be reduced by using dormancy‐breaking exogenous compounds. In a laboratory experiment, OSR seeds were coated with KNO₃, micronutrients, or gibberellic acid (GA) prior to a secondary dormancy test. In a field experiment, seeds were coated in a manner analogous to the laboratory experiment, and then buried 10 cm deep in the soil for 2.5 months. In a practical demonstration, OSR plants were sprayed with either urea ammonium nitrate (UAN) or a commercial product containing GA prior to seed maturity. Seed coating (laboratory and field experiments) reduced secondary dormancy and seed persistence in the field by up to 99%. The efficiency of the treatments for mitigating secondary dormancy (laboratory and field experiments) in decreasing order was GA > micronutrients > KNO₃ > control. With pre‐maturity spraying (practical demonstration), UAN reduced primary dormancy by up to 77% and the development of secondary dormancy by up to 38%; GA had no effect. Dormancy and seed persistence of OSR seeds may be reduced by a pre‐maturity UAN treatment of OSR mother plants, or by applying appropriate exogenous compounds to OSR seeds.     

Weed seed germination and the light environment: Implications for weed management

Light regulates dormancy termination and the subsequent germination in many weed species. Under field conditions, the light environment of the seeds, which is perceived mainly by photoreceptors of the phytochrome family, provides essential information for cueing germination in the proper environmental situation. The light environment's spectral composition and irradiance allow weed seeds to sense their position in the soil profile, the presence of a leaf canopy capturing light and other resources and the occurrence of soil cultivation. From an agronomical point of view, the control of germination by light represents a potentially useful step in the life cycle of weeds for developing effective control practices. The goal of this article is to place current knowledge regarding photoreceptors, physiological and molecular bases of seed responses to light and their ecological implications within the context of weed management in agricultural systems. With that final objective, the authors intend to show how a better understanding of the way in which the light environment regulates dormancy termination and the subsequent germination of weed seeds could be used to develop more accurate control practices and to improve weed management strategies.     

Longevity,dormancy and germination of Cyanus segetum

Cyanus segetum is an iconic, colourful weed in arable fields that provides ecological and societal services. To understand better both the infestation dynamics of C. segetum as an abundant, harmful weed and maintain sustainable populations where it provides beneficial services, we compared information on seed dormancy, seed longevity and germination conditions in two populations. Persistence of seeds buried in the soil was low, with <10% viable after 3 years. Periodic dormancy cycling was observed over the 4 years in the soil, with a maximum of dormant seeds in the spring and a minimum in the autumn; however, 20% of the seeds were non‐dormant all the time. Seeds of C. segetum were positive photosensitive, but light requirement varied among populations. Base water potential for germination was ?1 MPa. Base temperature ranged from 1 to 2°C. Optimum temperature for germination was about 10 to 15°C, but the mean thermal time varied greatly between populations, from 80 to 134 day °C. Photoperiod and temperature combinations had no effect on germination percentage, but both reduced the germination rate. Burial deeper than 2 cm greatly reduced germination and seedling emergence strongly decreased at depths >0.5 cm. No seeds buried deeper than 8 cm emerged. Low seed longevity and a wide range of germination conditions could partly explain the rapid disappearance of C. segetum populations after herbicide application began in western Europe. However, yearly sowing in restoration areas does not seem to be essential.     

Germination response of Phalaris paradoxa L. seed to different light qualities

The influence of different light regimes on the germination of Australian and English populations of Phalaris paradoxa L. (awned canary‐grass) seed was investigated to determine the impact of changing tillage practices on weed infestation. Seeds of all biotypes were highly viable, but differed in levels of innate dormancy (26–99%). In one experiment seed from a single Australian biotype, either enclosed in the spikelet glumes or having the spikelet glumes removed, were exposed to nine light treatments. Germination was stimulated by red and white light, but was inhibited by far‐red light. Time to 50% germination was less for seed enclosed in the spikelet glumes than for naked caryopses, although the final percentage of seed germinating when still enclosed in the spikelet glumes was significantly lower than for naked caryopses. In another experiment, six Australian and English biotypes with varying dormancy characteristics were exposed to eight light treatments. Red light did not stimulate germination in the deeply dormant biotype, however stimulated all other biotypes. Germination in darkness was below 20% in all biotypes except for one where germination was 51%. To overcome dormancy seeds were imbibed and placed in darkness at 16°C for either 7 or 14 days prior to exposure to red or white light for a single 15‐min period. Dormancy in all biotypes was overcome indicating that a period of burial may decrease the dormancy level and increase seed sensitivity to light. This increased light sensitivity suggests that exposure to light during tillage may stimulate germination in P. paradoxa seed.     

Factors affecting seed germination and emergence of Gomphrena perennis

Controlled growth chamber experiments were conducted to determine factors affecting seed germination and emergence of the troublesome weed Gomphrena perennis. The objective of this research was to examine the effects of temperature, light, moist chilling, osmotic potential, dry storage and depth of seed burial on G. perennis germination and emergence. The optimum temperature for germination was around 15–20°C. Seeds showed germination rates above 90% under 20/10 and 25/15°C temperature regimes. The minimum exposure to light needed to stimulate germination was 1 min. However, the light requirement was reduced after a long storage period. Furthermore, germination was high (>90%) in all moist‐chilling treatments tested. Germination was highly sensitive to increasing osmotic stress. The highest germination percentage (94%) was achieved at 0 MPa, and decreasing osmotic potential from 0 to ?0.3 MPa reduced germination to 11%. The highest seedling emergence occurred for seeds placed from 0 to 1 cm deep, and no seedlings emerged from a 5‐cm burial depth. Gomphrena perennis has a suitable environment in a no‐till soybean field, where seeds remaining on the surface have the required temperature, light and depth needed for germination.     

Weed seed decay in conventional and diversified cropping systems

Diversified cropping systems can have high soil microbial biomass and thus strong potential to reduce the weed seedbank through seed decay. This study, conducted in Iowa, USA, evaluated the hypothesis that weed seed decay is higher in a diversified 4‐year maize–soyabean–oat/lucerne–lucerne cropping system than in a conventional 2‐year maize–soyabean rotation. Mesh bags filled with either Setaria faberi or Abutilon theophrasti seeds and soil were buried at two depths in the maize phase of the two cropping systems and sampled over a 3‐year period. Setaria faberi seed decay was consistently greater at 2 cm than at 20 cm burial depth and was higher in the more diverse rotation than in the conventional rotation in 1 year. Abutilon theophrasti seeds decayed very little in comparison with seeds of S. faberi. Separate laboratory and field experiments confirmed differences in germination and seed decay among the seed lots evaluated each year. Fusarium, Pythium, Alternaria, Cladosporium and Trichoderma were the most abundant genera colonising seeds of both species. A glasshouse experiment determined a relationship between Pythium ultimum and S. faberi seed decay. Possible differences in seed susceptibility to decay indicate the need to evaluate weed seedbank dynamics in different cropping systems when evaluating overall population dynamics and formulating weed management strategies.     

A review of the relationship between primary and secondary dormancy,with reference to the volunteer crop weed oilseed rape (Brassica napus)

Fresh seeds of oilseed rape (Brassica napus) are reported to be nondormant and nonphotoblastic. However, a portion of the seeds can be induced into a light‐requiring state (secondary dormancy) for germination and also exhibit dormancy cycling. Thus, if seeds become buried in the soil they can form a persistent seedbank and become a serious volunteer weed in succeeding crops. The capacity of nondormant seeds of B. napus to be induced into secondary dormancy is contrary to results of studies on fresh nondormant seeds of some other species. A reanalysis of published and unpublished data shows that fresh seeds of this species have some degree of primary dormancy and that there is a significant relationship between primary dormancy and the capacity to enter secondary dormancy. However, most germination tests on B. napus have not been done in enough detail to detect primary dormancy (or not) in fresh seeds of this species. The usefulness of information on the relationship between primary dormancy and the capacity of the seeds to enter secondary dormancy is discussed in relation to management of weedy volunteers of this species.