Interspecific variation in persistence of buried weed seeds follows trade‐offs among physiological,chemical, and physical seed defenses

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Different requirements for physical dormancy release in two populations of <Emphasis Type="Italic">Sophora alopecuroides</Emphasis> relation to burial depth

Most species of Fabaceae produce seeds with physical dormancy which was broken by some environmental factors in the field, yet for the mechanism of physical dormancy release in the natural condition is still poorly understood. In present study, seeds of Sophora alopecuroides from two populations were placed on the soil surface or buried at depths of 2 and 7 cm in the field and were exhumed at time intervals to assess the changes of dormancy. Also, microenvironments were simulated in the laboratory to determine the possible components (temperature, temperature fluctuation, and moisture) involved in physical dormancy release. The laboratory work indicated that wet heat is an important dormancy release mechanism for both populations; and dry heat for one of them, but showed no effects on the other one. Consistent with that, burial experiment showed seeds of two populations had two contrasting-dormancy release patterns in the field. Most seeds of the dry heat sensitive population released dormancy but the other one remained dormant when seeds placed on the soil surface for 11 months (where seeds were exposed to dry heat but not wet heat). The difference of physical dormancy release pattern response to burial depth and simulated condition may attribute to seed coat color and hilum thickness which showed significantly different between two populations. This conclusion may improve understanding of physical dormancy release mechanism among populations and have a practical use for grassland management and weed control.     

Predicted global warming scenarios impact on the mother plant to alter seed dormancy and germination behaviour in Arabidopsis

Abstract Seed characteristics are key components of plant fitness that are influenced by temperature in their maternal environment, and temperature will change with global warming. To study the effect of such temperature changes, Arabidopsis thaliana plants were grown to produce seeds along a uniquely designed polyethylene tunnel having a thermal gradient reflecting local global warming predictions. Plants therefore experienced the same variations in temperature and light conditions but different mean temperatures. A range of seed‐related plant fitness estimates were measured. There were dramatic non‐linear temperature effects on the germination behaviour in two contrasting ecotypes. Maternal temperatures lower than 15–16 °C resulted in significantly greater primary dormancy. In addition, the impact of nitrate in the growing media on dormancy was shown only by seeds produced below 15–16 °C. However, there were no consistent effects on seed yield, number, or size. Effects on germination behaviour were shown to be a species characteristic responding to temperature and not time of year. Elevating temperature above this critical value during seed development has the potential to dramatically alter the timing of subsequent seed germination and the proportion entering the soil seed bank. This has potential consequences for the whole plant life cycle and species fitness.     

ENVIRONMENTAL CONTROL OF SEED GERMINATION IN THLASPI ARVENSE (CRUCIFERAE)

The effect of environmental conditions during storage and imbibition on germination was investigated in field pennycress (Thlaspi arvense L.), a weed species that can behave as a winter or a summer annual. Freshly harvested seeds of an inbred line with a cold requirement for flowering exhibited primary dormancy that was rapidly lost following 1 month of afterripening in a dry state. Nondormant seeds were positively photoblastic. The light effect was mediated through phytochrome since germination was promoted by red light and inhibited by far red light. Seedling emergence was also inhibited by light filtered through a canopy of wheat leaves. Germination of field pennycress seeds was considerably more sensitive to moisture stress than two sympatric species, wild oat (Avena fatua L.) and wheat (Triticum aestivum L., cv. ERA). Seeds of the latter two species were chosen in order to compare the effect of water potential on germination in field pennycress with that in sympatric species. It was concluded that the major environmental factor limiting nondormant field pennycress seeds on the soil surface was water availability. Imbibition of fully afterripened seeds at low temperatures (6 C) induced a deep secondary dormancy. In contrast to primary dormancy, cold-induced dormancy was not alleviated by red light, alternating temperatures (21/5 C), or 2 months of dry storage at 6, 15, or 35 C. However, exogenous gibberellin A₃ or 24 weeks of dry storage resulted in germination in cold-induced dormant seeds. Secondary dormancy was not observed in fully afterripened seeds that were preincubated at 21 C for 1 or 2 days prior to the cold treatment. These results may explain the failure in field experiments to observe the cold-induced secondary dormancy that limits spring emergence in other winter annuals (J. Baskin, C. Baskin, Weed Res. 1979 19: 285–292).     

Effect of maternal nitrogen and drought stress on seed dormancy and germinability of Amaranthus retroflexus

Amaranthus retroflexus L. is an importunate annual weed in many cropping systems of different countries. The main aim of this study was to investigate the effects of maternal nitrogen and drought stress on the seed dormancy and germinability of A. retroflexus. Field experiment was carried out in a factorial based on randomized complete block design, with four potential levels of soil water (–2, ?6, ?8 and ?10 bar) and three levels of nitrogen (0, 100 and 200 kg/ha). The germination characteristics of the seeds were measured at three different times (1 month, 6 months and 1 year after harvesting). Results showed that drought stress had positive effects on breaking of A. retroflexus seed dormancy until 6 months after seed harvesting. Seeds that were developed under severe water stress exhibited the highest germination percentage and germination rate. The results obtained from this study revealed that application of 100 kg/ha nitrogen during seed development increases germinability of A. retroflexus, whereas application of 200 kg/ha nitrogen induced seed dormancy. Furthermore, 100 kg/ha nitrogen application in the field along with 200 ppm gibberellic‐acid treatment during seed after‐ripening showed the highest germination percentage and germination rate for seeds after 6 months harvesting. Results also indicated that after‐ripening significantly increased seed germination and germination rate of A. retroflexus. These findings indicate that long‐term management of the soil seed bank in this species requires more stringent control due to the changes in germination timing, as detected in this study.     

Annual dormancy cycles in buried seeds of shrub species: germination ecology of Sideritis serrata (Labiatae)

The germination ecology of Sideritis serrata was investigated in order to improve ex‐situ propagation techniques and management of their habitat. Specifically, we analysed: (i) influence of temperature, light conditions and seed age on germination patterns; (ii) phenology of germination; (iii) germinative response of buried seeds to seasonal temperature changes; (iv) temperature requirements for induction and breaking of secondary dormancy; (v) ability to form persistent soil seed banks; and (vi) seed bank dynamics. Freshly matured seeds showed conditional physiological dormancy, germinating at low and cool temperatures but not at high ones (28/14 and 32/18 °C). Germination ability increased with time of dry storage, suggesting the existence of non‐deep physiological dormancy. Under unheated shade‐house conditions, germination was concentrated in the first autumn. S. serrata seeds buried and exposed to natural seasonal temperature variations in the shade‐house, exhibited an annual conditional dormancy/non‐dormancy cycle, coming out of conditional dormancy in summer and re‐entering it in winter. Non‐dormant seeds were clearly induced into dormancy when stratified at 5 or 15/4 °C for 8 weeks. Dormant seeds, stratified at 28/14 or 32/18 °C for 16 weeks, became non‐dormant if they were subsequently incubated over a temperature range from 15/4 to 32/18 °C. S. serrata is able to form small persistent soil seed banks. The maximum seed life span in the soil was 4 years, decreasing with burial depth. This is the second report of an annual conditional dormancy/non‐dormancy cycle in seeds of shrub species.     

The impact of global warming on germination and seedling emergence in Alliaria petiolata,a woodland species with dormancy loss dependent on low temperature

• The impact of global warming on seed dormancy loss and germination was investigated in Alliaria petiolata (garlic mustard), a common woodland/hedgerow plant in Eurasia, considered invasive in North America. Increased temperature may have serious implications, since seeds of this species germinate and emerge at low temperatures early in spring to establish and grow before canopy development of competing species.
• Dormancy was evaluated in seeds buried in field soils. Seedling emergence was also investigated in the field, and in a thermogradient tunnel under global warming scenarios representing predicted UK air temperatures through to 2080.
• Dormancy was simple, and its relief required the accumulation of low temperature chilling time. Under a global warming scenario, dormancy relief and seedling emergence declined and seed mortality increased as soil temperature increased along a thermal gradient. Seedling emergence advanced with soil temperature, peaking 8 days earlier under 2080 conditions.
• The results indicate that as mean temperature increases due to global warming, the chilling requirement for dormancy relief may not be fully satisfied, but seedling emergence will continue from low dormancy seeds in the population. Adaptation resulting from selection of this low dormancy proportion is likely to reduce the overall population chilling requirement. Seedling emergence is also likely to keep pace with the advancement of biological spring, enabling A. petiolata to maintain its strategy of establishment before the woodland canopy closes. However, this potential for adaptation may be countered by increased seed mortality in the seed bank as soils warm.

     

Echinochloa crus‐galli seed physiological dormancy and germination responses to hypoxic floodwaters

• Hypoxic floodwaters can seriously damage seedlings. Seed dormancy could be an effective trait to avoid lethal underwater germination. This research aimed to discover novel adaptive dormancy responses to hypoxic floodwaters in seeds of Echinochloa crus‐galli, a noxious weed from rice fields and lowland croplands.
• Echinochloa crus‐galli dormant seeds were subjected to a series of sequential treatments. Seeds were: (i) submerged under hypoxic floodwater (simulated with hypoxic flasks) at different temperatures for 15 or 30 days, and germination tested under drained conditions while exposing seeds to dormancy‐breaking signals (alternating temperatures, nitrate (KNO₃), light); or (ii) exposed to dormancy‐breaking signals during hypoxic submergence, and germination monitored during incubation and after transfer to drained conditions.
• Echinochloa crus‐galli seed primary dormancy was attenuated under hypoxic submergence but to a lesser extent than under drained conditions. Hypoxic floodwater did not reinforced dormancy but hindered secondary dormancy induction in warm temperatures. Seeds did not germinate under hypoxic submergence even when subjected to dormancy‐breaking signals; however, these signals broke dormancy in seeds submerged under normoxic water. Seeds submerged in hypoxic water could sense light through phytochrome signals and germinated when normoxic conditions were regained.
• Hypoxic floodwaters interfere with E. crus‐galli seed seasonal dormancy changes. Dormancy‐breaking signals are overridden during hypoxic floods, drastically decreasing underwater germination. In addition, results indicate that a fraction of E. crus‐galli seeds perceive dormancy‐breaking signals under hypoxic water and germinate immediately after aerobic conditions are regained, a hazardous yet less competitive environment for establishment.

     

The relationship between seed dormancy,seed size and weediness,in Crepis tectorum (Asteraceae)

Summary I examined the germination characteristics of weed and outcrop populations of Crepis tectorum to test the hypothesis that the presumably more ephemeral weed habitat favors the highest levels of seed dormancy. The winter annual habit characterizing most plants of this species was reflected in a rapid germination of seeds sown in late summer. A slightly higher fraction of surface-sown seeds of weed plants delayed germination. Buried seeds of weed plants also survived better than seeds produced by plants in most outcrop populations, supporting the idea that weediness favors seed dormancy and a persistent seed bank. However, the differences in seed dormancy between the two ecotypes were small and not entirely consistent. Furthermore, high levels of seed dormancy were induced during burial in the outcrop group, suggesting that there is a potential for a dormant seed population in this habitat as well. Demographic data from one of the outcrop populations verified the presence of a large between-year seed bank. Possible environmental factors favoring seed dormancy in outcrop populations are discussed. The unusually large seeds of weedy Crepis contrasts with the relatively small difference in seed dormancy between the two ecotypes.     

Seed development and germination ecophysiology of the invasive tree <Emphasis Type="Italic">Prunus serotina</Emphasis> (Rosaceae) in a temperate forest in Western Europe

Seed development, dormancy and germination of the American invasive tree species, Prunus serotina, are described for plants growing in a large forest in Belgium. Seeds of P. serotina were collected following anthesis in the first week of July and thereafter at fortnightly intervals. Seed dormancy, temperature requirements for germination and the soil seed bank were investigated. At maturation (about 105 days after anthesis), seed moisture content had decreased to around 13.7%, and 44% of the seeds had attained the capacity to germinate. Mature seeds of P. serotina exhibited physiological dormancy, germinating only after a long cold, moist stratification period. Highest germination percentage occurred in seeds treated with gibberellic acid (GA₃), at 10°C. We found no evidence that P. serotina forms a persistent seed bank but noticed a persistent seedling bank in the field.     

Ecological longevity of Polaskia chende (Cactaceae) seeds in the soil seed bank,seedling emergence and survival

• Soil seed banks are essential elements of plant population dynamics, enabling species to maintain genetic variability, withstand periods of adversity and persist over time, including for cactus species. However knowledge of the soil seed bank in cacti is scanty. In this study, over a 5‐year period we studied the seed bank dynamics, seedling emergence and nurse plant facilitation of Polaskia chende, an endemic columnar cactus of central Mexico.
• P. chende seeds were collected for a wild population in Puebla, Mexico. Freshly collected seeds were sown at 25 °C and 12‐h photoperiod under white light, far‐red light and darkness. The collected seeds were divided in two lots, the first was stored in the laboratory and the second was use to bury seeds in open areas and beneath a shrub canopy. Seeds were exhumed periodically over 5 years. At the same time seeds were sown in open areas and beneath shrub canopies; seedling emergence and survival were recorded over different periods of time for 5 years.
• The species forms long‐term persistent soil seed banks. The timing of seedling emergence via germination in the field was regulated by interaction between light, temperature and soil moisture. Seeds entered secondary dormancy at specific times according to the expression of environmental factors, demonstrating irregular dormancy cycling.
• Seedling survival of P. chende was improved under Acacia constricta nurse plants. Finally, plant facilitation affected the soil seed bank dynamics as it promoted the formation of a soil seed bank, but not its persistence.

     

Modelling field emergence patterns in arable weeds

A model was developed to simulate weed emergence patterns after soil cultivation. In the model, the consecutive processes of dormancy release, germination and pre-emergence growth were modelled in separate modules. Input variables of the model were: date of soil cultivation, soil temperature and soil penetration resistance. Output variables of the model were: seedling density and timing of seedling emergence. The model was parameterized for Polygonum persicaria , Chenopodium album and Spergula arvensis with data from previous field and laboratory experiments. The model was evaluated with data from an experiment, in which emergence of P. persicaria , C. album and S. arvensis was monitored in field plots that were cultivated once only, at one of five dates in the spring. At the same time as the field observations on seedling emergence, seasonal changes in seed dormancy of the buried weed seeds were assessed by testing the germination of seed lots that were buried in envelopes. From a comparison between field observations and simulated data, it appeared that the model overestimated the rate of dormancy release in spring, whereas germination and pre-emergence growth were simulated well. In general, therefore, both the numbers of emerging seedlings and the timing of emergence could be predicted accurately, when dormancy was not simulated but introduced from experimental data. Improvement of predictions of field emergence of weeds should mainly focus on increasing the precision of the simulation of dormancy release. Close correlations were found between seedbed temperature and both the extent and rate of seedling emergence, but analysis with the simulation model revealed that they were only partly based on causal relationships, so that they have limited predictive value.     

Note on organic dormancy of estuarine Ruppia maritima L. seeds

Information on seed dormancy is one of the primary requirements for successful seedling propagation of submerged aquatics and seagrass. Studies on Ruppia maritima seed germination have been done, but the presence, requirements, and the types of dormancy have not been well understood. A laboratory study was conducted to understand presence and types of organic dormancy of the seeds of estuarine R. maritima collected from Lake Pontchartrain, Louisiana, USA. Our study results indicate that the brackish estuarine R. maritima population produces seeds that do not have any noticeable initial morphological, physical, and physiological dormancy. Although dry stratification reduced seed viability and final germination rates, drying seems to induce an earlier germination in R. maritima. Desiccation also appears to induce an environmental dormancy that can be disrupted by exposure to water. Further study on environmental dormancy is needed to provide information to develop methods for long-term seed storage that can be employed in greenhouse seedling propagation. Handling editor: P. Viaroli     

Fate of the soil seed bank of giant ragweed and its significance in preventing and controlling its invasion in grasslands

Giant ragweed (Ambrosia trifida, L. henceforth referred to as GR), an annual non‐native invasive weed, may cause health problems and can reduce agricultural productivity. Chemical control of GR in grasslands may have irreversible side effects on herbs and livestock. In an attempt to propose a solution to the harmful effects of GR on grasslands, this study explores the fate of its soil seed bank (SSB) and considers the physical control of its SSB reduction. By studying GR distributed in grasslands of the Yili Valley, Xinjiang, China, we measured the spatial and temporal changes in seed density, seed germination, dormancy, and death. We analyzed seed germination, dormancy, and death following different storage periods. The study analyzed population characteristics over time, including seed fate, and examined physical control methods for reducing the SSB density. The SSB of GR occurs in the upper 0–15 cm of soil in grasslands. Seed density in the SSB decreased by 68.1% to 82.01% from the reproductive growth period to the senescence period. More than 98.7% of the seeds were rotten, eaten, germinated, dispersed, or died within one year after being produced. The seed germination rate of the SSB decreased with the number of years after invasion. When stored for 0.5 or 3.5 years, seed germination rates fell by 40%, during which time seed death rate increased by almost 40%. When GR was completely eradicated for two consecutive years, the SSB and population densities decreased by >99%. The vast majority of GR seeds germinated or died within one year; the germination rate decreased significantly if the seeds were stored dry at room temperature for a long time. Newly produced seeds are the main source of seeds in the SSB. Therefore, thoroughly eradicating GR plants for several years before the seeds can mature provides an effective control method in grasslands.     

Diaspore dispersal season influences root growth and seed traits,but not germination responses,of sympatric Piper species in a seasonal forest

• Seeds may differ in terms of dormancy, longevity, sensitivity to desiccation and dry mass, according to the timing (dry season/rainy season) of diaspore dispersal. In addition, seasonal variations in temperature and water availability can act as signals of the season during seed development, influencing germination responses and root growth. We evaluated the effects of temperature variations and water availability on germination parameters, root growth and seed traits of four coexisting Piper species in seasonal vegetation that differed in diaspore dispersal timing.
• Eight temperature treatments (15, 20, 23, 25, 28, 30, 35 °C, and alternate 30 °C–20 °C) and four induced water potentials (0, −0.3, −0.6 and −1.2 MPa) were used. The parameters germination onset, germination percentage (G%), mean germination time (MGT), root elongation, seed longevity during ex situ storage and dry mass of seeds were evaluated.
• Germination responses observed were independent of the diaspore dispersal timing, such as variations in germination onset, G% and MGT, both in temperature and water availability treatments. In contrast, root elongation, longevity and dry mass of seeds varied according to the time of diaspore dispersal.
• Our results corroborate the hypothesis that the timing of diaspore dispersal is an important factor in controlling the initial development of seedlings in seasonal vegetation, but not in germination responses. The predominance of negative effects of temperature increases and water deficit on root growth shows that the initial stages of plant development can be strongly impacted by these environmental factors.

     

Ecological role of physical dormancy in seeds of Oxytropis racemosa in a semiarid sandland with unpredictable rainfall

Aims Seed dormancy and the soil seed bank are crucial to plant regeneration strategy, especially in semiarid ecosystems with unpredictable precipitation. The aim of this study was to investigate how seed dormancy is controlled by environmental factors and how it is correlated with the soil seed bank and regeneration of the perennial legume Oxytropis racemosa, a dominant perennial herb in Mu Us Sandland of semiarid China.Methods Germination and imbibition experiments on fresh intact and scarified seeds of O. racemosa were used to identify physical dormancy (PY) in seeds of this species. Soil seed bank dynamics, timing of seedling emergence and the fate of buried seeds in the natural habitat were investigated.Important findings PY was broken by mechanical scarification or wet heat/ice water cycles but not solely by dry heat or wet heat treatment. The soil seed bank exhibited seasonal changes in the number of seeds, which was highest in September and lowest in July. Seeds buried at different sand depths gradually lost dormancy; 20–42% of the seeds remained dormant after 20 months of burial. Dormancy break occurs gradually throughout the year. Our results indicate that O. racemosa exhibits hardcoatedness heterogeneity that spreads germination of a seed cohort between seasons and years in the semiarid environment, where the amount of precipitation during the growing season is highly variable.     

Soil warming increases plant species richness but decreases germination from the alpine soil seed bank

Global warming is occurring more rapidly above the treeline than at lower elevations and alpine areas are predicted to experience above average warming in the future. Temperature is a primary factor in stimulating seed germination and regulating changes in seed dormancy status. Thus, plant regeneration from seed will be crucial to the persistence, migration and post disturbance recruitment of alpine plants in future climates. Here, we present the first assessment of the impact of soil warming on germination from the persistent alpine soil seed bank. Contrary to expectations, soil warming lead to reduced overall germination from the soil seed bank. However, germination response to soil temperature was species specific such that total species richness actually increased by nine with soil warming. We further explored the system by assessing the prevalence of seed dormancy and germination response to soil disturbance, the frequency of which is predicted to increase under climate change. Seeds of a significant proportion of species demonstrated physiological dormancy mechanisms and germination of several species appeared to be intrinsically linked to soil disturbance. In addition, we found no evidence of subalpine species and little evidence of exotic weed species in the soil, suggesting that the soil seed bank will not facilitate their invasion of the alpine zone. In conclusion, changes in recruitment via the alpine soil seed bank can be expected under climate change, as a result of altered dormancy alleviation and germination cues. Furthermore, the alpine soil seed bank, and the species richness therein, has the potential to help maintain local species diversity, support species range shift and moderate species dominance. Implications for alpine management and areas for further study are also discussed.     

Aquaporins influence seed dormancy and germination in response to stress

Aquaporins influence water flow in plants, yet little is known of their involvement in the water‐driven process of seed germination. We therefore investigated their role in seeds in the laboratory and under field and global warming conditions. We mapped the expression of tonoplast intrinsic proteins (TIPs) during dormancy cycling and during germination under normal and water stress conditions. We found that the two key tonoplast aquaporins, TIP3;1 and TIP3;2, which have previously been implicated in water or solute transport, respectively, act antagonistically to modulate the response to abscisic acid, with TIP3;1 being a positive and TIP3;2 a negative regulator. A third isoform, TIP4;1, which is normally expressed upon completion of germination, was found to play an earlier role during water stress. Seed TIPs also contribute to the regulation of depth of primary dormancy and differences in the induction of secondary dormancy during dormancy cycling. Protein and gene expression during annual cycling under field conditions and a global warming scenario further illustrate this role. We propose that the different responses of the seed TIP contribute to mechanisms that influence dormancy status and the timing of germination under variable soil conditions.     

Fire and flood: Soil‐stored seed bank and germination ecology in the endangered Carrington Falls Grevillea (Grevillea rivularis,Proteaceae)

Abstract Seed germination is dependent on the interaction between the dormancy state of a seed and the presence of favourable environmental conditions. Thus, the spectacular pulse of seedling recruitment in many Australian vegetation communities following disturbances such as fire can be attributed to changes in microsite conditions and/or the dormancy‐breaking effect of the disturbance on accumulated seed banks. Grevillea rivularis is a threatened species endemic to the area immediately above Carrington Falls in the NSW Southern Highlands. Most of the population is confined to the riparian vegetation zone in woodland and heath, and is therefore subject to periodic disturbance from fire and flood. For this species, a pulse of seedling recruitment has been recorded after fire, flood and mechanical soil disturbance. The aims of this study were to examine the density and vertical distribution of the soil‐stored seed bank and to investigate the role of heat and scarification as cues for germination of fresh and soil‐stored seed. There was a large seed bank under the canopies of established individuals (194 ± 73 seeds m^?2) and most seeds were found in the 0–2 cm and leaf‐litter layers of the soil profile. The germination response of soil‐stored and fresh seed was examined using a hierarchical series of laboratory experiments. Seeds of G. rivularis showed marked dormancy polymorphism. Thirty‐six percent of soil‐stored seed germinated without treatment, whereas no untreated fresh seeds germinated. Scarification or heating caused significant germination of dormant soil‐stored seed, but only scarification resulted in germination of dormant fresh seeds. These results highlight important differences in the dormancy state of soil‐stored and fresh seed. Thus, being a riparian species in a fire‐prone environment, the dormancy mechanisms in seeds of G. rivularis suit this species to disturbance by both fire and flood.