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.

     

Evidence for the retention of genetic variation in Erodium seed dormancy by variable rainfall

Summary The periodic occurrence of summer/early autumn precipitation in the California annual grassland can result in the formation of early and late emerging cohorts of Erodium botrys and E. brachycarpum. The occurrence of early rainfall and the timing of such rainfall are highly variable from year to year. A series of field watering experiments in 1980–81 were used to simulate early emergence conditions that would result from significant rainfall (1 cm) occurring in mid-July, late August, and mid-September. Net reproduction was used to estimate fitness differentials between Erodium cohorts emerging in response to a watering treatment (early emerging cohorts) and Erodium cohorts emerging with the onset of winter rains in mid-October (late emerging cohorts). Survival was lower and gross reproduction was higher among early emerging cohorts than late emerging cohorts. For both species, net reproduction of the early cohort was lower than that of the late cohort under the July watering treatment and higher than that of the late cohort under the August watering treatment.Early cohorts, formed in response to rainfall in mid-September, 1982, were also compared demographically to later cohorts emerging in October. Compared to late cohorts, net reproduction, gross reproduction and survival were higher for the early cohorts.Common garden experiments indicate that differences in the duration of seed dormancy between the progenies of early and late emerging plants reflect a significant genetic component. Progency produced by early cohorts of E. brachycarpum from all three watering treatments possessed more extended seed dormancy than progeny of late cohorts. In E. botrys, progeny from early cohorts emerging in response to the July watering treatment were also more dormant than late progeny. In contrast, early cohorts of E. botrys emerging in response to the September watering treatment produced seed less dormant than seed produced by late cohorts. When combined with demographic data, indicating that fitness differentials between early and late cohorts varied with changes in the date of early emergence, genetic results suggest that year to year variation in early rainfall may act to retain genetic variation in the duration of seed dormancy.     

白皮松种子休眠特性的种源变异

对７个具有代表性种源的白皮松种子的休眠特性研究表明,白皮松种子种皮的透水性在种源间差异明显,表现出随地理纬度的增高而逐渐增大的趋势,分布越是偏北的种源透水性趋于加强。根据种皮的透水性,７个种源可被分为差异明显的２组,组间以秦岭为界。不同种源种子种皮的透气性以外种皮造成的差异为最大,内种皮次之。种子的透气性与透水性表现出的变化趋势相同。不同种源种子浸提液的抑制能力显著不同,而且这种差异随浸提液浓度的     

Variation in seed dormancy and germination among populations of Silybum marianum (Asteraceae)

Milk thistle (Silybum marianum) is a medicinal plant; however, lack of consistency in past dormancy studies has hindered propagation of this species from seeds. We tested the germination responses of freshly harvested and after-ripened (stored for 2 and 7 months; 25°C at 50% relative humidity) seeds from three populations (P1, P2 and P3) in Iran at varying constant or alternating temperatures, with or without GA₃ and in light and continuous darkness. No germination occurred in freshly harvested seeds incubated at any condition without GA₃ application, indicating that all the seeds were dormant. Seeds from P1 and P2, which developed under relatively dry, warm conditions, germinated over a wider range of temperatures after 2 months of dry storage, indicating type 6 of non-deep physiological dormancy (PD). Seeds from P3, which developed under relatively wet, cool conditions, incubated at constant temperatures (especially on GA₃), exhibited an increase in maximum temperature for germination, indicating type 1 of non-deep PD. Light improved germination of after-ripened seeds, and GA₃ application substituted for the light requirement for germination. This is the first report that environmental conditions during seed development may be correlated with differences in the type of non-deep PD. We conclude that milk thistle seeds are positively photoblastic and photodormant and the germination responses of after-ripened seeds from different populations are different under darkness. Therefore, the impacts of genetic differences and maternal effects on the induction of dormancy during seed development should be considered in attempts to domesticate this medicinal plant.     

横断山区干旱河谷川滇蔷薇种子休眠与萌发的地理空间差异

通过测定横断山区干旱河谷18个川滇蔷薇种群新采集种子以及低温层积8周种子的发芽率和发芽速度,分析发芽率和发芽速度与种子特征以及环境因子之间的关系,阐明川滇蔷薇种子休眠与萌发的地理空间差异及其影响因素。结果表明,18个川滇蔷薇种群种子具有不同程度的休眠,新采集种子发芽率变化幅度大,为15.8%±5.0%至82.7%±2.3%,发芽速度指数范围：2.3%±0.2%至5.3%±0.5%。不同种群种子发芽率和发芽速度差异显著。新采集种子的发芽率在流域间存在显著差异,表现为金沙江流域 > 雅砻江流域 > 大渡河流域和岷江流域。新采集种子的发芽率和发芽速度随着采集地点海拔的升高而显著增加,即种子休眠程度随着海拔的升高呈现降低趋势。低温层积8周显著提高了种子发芽率和发芽速度,但减弱了种子发芽率和发芽速度在流域以及海拔间的差异。偏相关分析表明：瘦果皮厚度与新采集种子萌发速度成显著负相关关系;环境因子中年蒸发量与发芽率以及发芽速度之间关系最为密切,成极显著正相关关系;其次为年降水量,与发芽率和发芽速度之间具有显著的负相关关系。综合分析表明,川滇蔷薇种子休眠与萌发在横断山区干旱河谷存在较强地理空间差异。环境因子中年蒸发量和年降水量以及植物自身特征瘦果皮厚度是引起种子休眠与萌发地理空间差异的主要因素。     

中国境内稗(Echinochloa crus-galli)形态变异及其遗传和地理背景分析

采集了浙江、福建、江苏、湖南、湖北、四川、重庆、黑龙江、河南9个省的稗（Echinochloa crus-galli（L.）P.Beauv.）及其变种的33份种子,分别播种在相同的环境下,获得33个种群,测定了种群的16个形态性状,筛选出重复性好的9条ISSR引物,从33个种群中扩增出了109个位点。基于这些形态性状和ISSR位点信息,对33个种群先进行主成分分析,在此基础上再进行模糊均值聚类分析,探讨了它们的形态和遗传变化特点,及其与形态-遗传-地理背景三者之间的关系。主要结论如下：（1）33个种群可以鉴别出形态性状相对一致的4组,能够识别出西来稗（E.crus-galli var.zelayensis（Kunth）Farw.）、无芒稗（E.crus-galli var.mitis（Pursh）Peterm.）、细叶旱稗（E.crus-galli var.praticola Ohwi）;（2）基于109个位点信息对33个种群进行聚类分析得到了6组,部分组与形态聚类分组有一定的对应性;（3）33个稗草种群的遗传分化受地理背景因素的影响（r=0.684,n=33,P<0.001）;形态变异也有较明显的遗传背景因素（r=0.425,n=33,P<0.02）。在相对一致的稻田生境中,可能存在着形态上的趋同适应,使遗传上分化的组间在形态上又往往有交叉过渡,致使稗原变种（E.crus-galli var.crus-galli）、西来稗、无芒稗、短芒稗（E.crus-galli var.breviseta（Döll）Podp.）在形态上难以区别;（4）基于遗传和形态数据分析,发现细叶旱稗无论在形态上,还是遗传上,均形成了明显的一组,推测与该种长期适应于干旱生境有关,建议将细叶旱稗提升为种的水平,并将其命名为Echinochloa praticola（Ohwi）Guo S L,Lu Y L,Yin L P&Zou M Y。     

Genomic regions affecting seed shattering and seed dormancy in rice

Non-shattering of the seeds and reduced seed dormancy were selected consciously and unconsciously during the domestication of rice, as in other cereals. Both traits are quantitative and their genetic bases are not fully elucidated, though several genes with relatively large effects have been identified. In the present study, we attempted to detect genomic regions associated with shattering and dormancy using 125 recombinant inbred lines obtained from a cross between cultivated and wild rice strains. A total of 147 markers were mapped on 12 rice chromosomes, and QTL analysis was performed by simple interval mapping and composite interval mapping. For seed shattering, two methods revealed the same four QTLs. On the other hand, for seed dormancy a number of QTLs were estimated by the two methods. Based on the results obtained with the intact and de-hulled seeds, QTLs affecting hull-imposed dormancy and kernel dormancy, respectively, were estimated. Some QTLs detected by simple interval mapping were not significant by composite interval mapping, which reduces the effects of residual variation due to the genetic background. Several chromosomal regions where shattering QTLs and dormancy QTLs are linked with each other were found. This redundancy of QTL associations was explained by ”multifactorial linkages” followed by natural selection favoring these two co-adapted traits. Received: 23 November 1998 / Accepted: 27 August 1999     

An ecological approach to study the physical and chemical effects of rye cover crop residues on Amaranthus retroflexus, Echinochloa crus-galli and maize

Weeds can be suppressed in the field by cover crop residues, extracts of which have been demonstrated to exert chemical inhibition of crop and weed germination and early growth in bioassays. In this study, two complementary bioassays were developed with soil and mulch material originating from a long‐term maize–cover crop experiment to determine the relative physical and chemical effect of rye cover crop residues on weed and maize germination and early growth. This was compared with the effect exerted by residue material from the natural vegetation that developed in the crop stubble during the winter before maize sowing. Germination percentage and early growth of maize and two maize weeds, Amaranthus retroflexus and Echinochloa crus‐galli, were assessed in a seed incubator in tilled (green manured) and nontilled (surface mulched) soil, with and without N fertilisation, at various dates after cover crop destruction. Responses were compared to those of the same species in a standard soil without mulch or with an inert poplar mulch. A second bioassay was set up in a glasshouse to determine the effect of different quantities of fresh residue material and additional N fertilisation on emergence speed and percentage and on plant vigour during the first 22 days after cover crop destruction. These results were compared with no‐mulch controls and poplar mulch controls. Results of these trials were compared with weed density and biomass that developed in the maize crop sown after cover crop destruction. Soil and mulch chemical and biological properties were determined for material collected in the field at different times after cover crop destruction. Chemical properties of the mulch differed only occasionally between the treatments, but variation in cover crop biomass production led to significantly different soil chemical properties. Although soil total phenolic acid content did not always correlate to weed and maize germination and early growth inhibition, soil microbial activity did. In suboptimal conditions, as is often the case in the field, plant residue material exerted both a physical and a chemical effect on maize and weed emergence and early growth. Nitrogen fertilisation and application timing can give the maize crop a competitive advantage with respect to the weeds, but the final response and the practical consequences depended largely on the weed species involved.     

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.     

植物种子休眠的原因及休眠的多形性

概述了植物种子休眠的原因及种子休眠的多形性。种壳障碍、胚形态发育不全和生理后熟以及种子中含有化学抑制剂都可导致种子休眠。根据不同的分类标准可将种子休眠可分成不同类型,但通常将休眠分为外源休眠、内源休眠和综合休眠。影响休眠的因素是复杂的,植物种类不同,休眠特性不同;同种植物的种子,来源于不同居群和植株,在不同时期采集,位于母株不同位置,其休眠有可能不同;甚至同一果实中的不同种子,休眠特性都会有差异。影响休眠性状表达的基因既有核基因,又有质基因,休眠通常表现为一种受多基因控制的数量性状。种子休眠的多形性有利于调节种子萌发的时空分布。     

Latitudinal variation in seed weight and flower number in Prunella vulgaris

Studies of seed-weight variation across altitudinal and latitudinal gradients have led to conflicting hypotheses regarding the selective value of this traint in relation to the length of the growing season. Growing-season length may also influence the evolution of seed number, and population differentiation in seed weight may be constrained by a negative genetic correlation between seed weight and seed number within populations. We examined variation in seed weight and an estimate of seed number (flower number) and the covariance of these traits among populations of Prunella vulgaris at five latitudes between northern Michigan and South Carolina. We measured seed weight and flower number in native habitats and in a common environment to determine the extent to which patterns observed in the field reflect genetic differentiation. We observed no genetically based variation in seed weight across the latitudinal gradient, although genetic variation among populations within a latitude was observed. In contrast to the lack of variation in seed weight, flower number increased clinally from northern Michigan to Tennessee in a common environment. Population mean flowering date in a common environment was successively later from north to south. Later-flowering individuals appear to achieve a larger size before flowering and consequently possess more resources for seed production. This difference may account for the greater flower production of late-flowering, southern populations. Independence of population mean seed weight and flower number across the latitudinal gradient suggests that population differentiation in seed weight has not been constrained by a trade-off between seed size and number within populations.     

长白落叶松种群地理变异研究

根据17年生长白落叶松种源试验资料,从生态遗传学的观点出发,采用典型相关分析等方法,揭示了长白落叶松地理变异规律可以认为,以海拔垂直梯度为主、纬向渐变为辅的连续型变异;低海拔、低等效纬度的小北湖种源是长白落叶松优良基因资源中心,生长较快、生长期较长;水热因子的综合选择作用是长白落叶松地理变异的重要因素,其中温度是主导因子;在地理种群的遗传分化中,生长性状的分化最为明显,可以作为种源区区划的主要性状;从长白落叶松分布的低等效纬度地区向高等效纬度地区调种(一般不超过2个等效纬度),可望获得较大的遗传增益.     

Some reflections on the relationship between endogenous hormones and light-mediated seed dormancy

The relationships between phytochrome and endogenous hormones in the light-mediated control of seed dormancy are discussed. It is concluded that gibberellins are primarily involved in post-dormancy metabolic processes leading to embryo growth and radicle emergence, such as food reserve mobilisation and endosperm softening. Evidence is considered that germination inhibitors, particularly abscisic acid, are involved in the establishment and maintenance of primary dormancy. The role of cytokinins not fully elucidated but there is considerable evidence to suggest that phytochrome control may involve cytokinin effects on transmembrane ion fluxes. In terms of hormonal control, phytochrome mediated dormancy is a complex phenomenon. There is a need for molecular studies of processes controlled by phytochrome, GAs, CKs and ABA during dormancy and germination to unravel the complexities of the dormancy mechanisms. Such studies would be facilitated by the availability of CK-deficient mutants of classical light-sensitive species.     

Differential effects of chilling on the activity of C4 enzymes in two ecotypes of Echinochloa crus-galli from sites of contrasting climates

Five-week-old plants of Echinochloa crusgalli (L.) Beauv. from Mississippi and from Québec grown under controlled conditions were subjected to dark chilling for 10 h at 5°C or light chilling treatments for 14 h at 7°C under hight light (1 000 μmol m⁻² s⁻¹). The activities of four C₄ enzymes of Québec plants, measured 4 h after the completion of the cold treatment, were not affected by the chilling treatment in the dark. The activities of pyruvate, P_i dikinase (PPDK; EC 2.7.9.1) and NADP⁺-malic enzyme (NADP⁺-ME; EC 1.1.1.40), were significantly reduced in dark-chilled Mississippi plants. Chilling under high light conditions elicited significant levels of reduction in the activities of the four enzymes from both ecotypes but the reductions were significantly less severe for Québec plants. The recovery of activities of phosphoenolpyruvate carboxylase (PEPC; EC 4.1.1.31) and PPDK for both ecotypes was completed within 36 to 60 hours following the chilling treatment, but NADP⁺-malate dehydro-genase (NADP⁺-MDH; EC 1.1.1.82) and NADP⁺-ME activities of chilled Mississippi plants remained below that of control plants at the end of the 5-day monitoring period. PPDK was inactivated in vitro at 0 and 10°C and the rates of cold inactivation were significantly higher for PPDK extracted from Mississippi plants. The activity of PEPC of Mississippi extracts was slightly, but significantly reduced by a 60 min treatment at 0°C.     

Genes controlling seed dormancy and pre-harvest sprouting in a rice-wheat-barley comparison

Pre-harvest sprouting results in significant economic loss for the grain industry around the world. Lack of adequate seed dormancy is the major reason for pre-harvest sprouting in the field under wet weather conditions. Although this trait is governed by multiple genes it is also highly heritable. A major QTL controlling both pre-harvest sprouting and seed dormancy has been identified on the long arm of barley chromosome 5H, and it explains over 70% of the phenotypic variation. Comparative genomics approaches among barley, wheat and rice were used to identify candidate gene(s) controlling seed dormancy and hence one aspect of pre-harvest sprouting. The barley seed dormancy/pre-harvest sprouting QTL was located in a region that showed good synteny with the terminal end of the long arm of rice chromosome 3. The rice DNA sequences were annotated and a gene encoding GA20-oxidase was identified as a candidate gene controlling the seed dormancy/pre-harvest sprouting QTL on 5HL. This chromosomal region also shared synteny with the telomere region of wheat chromosome 4AL, but was located outside of the QTL reported for seed dormancy in wheat. The wheat chromosome 4AL QTL region for seed dormancy was syntenic to both rice chromosome 3 and 11. In both cases, corresponding QTLs for seed dormancy have been mapped in rice.C. Li and P. Ni contributed equally to this work     

Control of seed dormancy in Nicotiana plumbaginifolia: post-imbibition abscisic acid synthesis imposes dormancy maintenance

The physiological characteristics of seed dormancy in Nicotiana plumbaginifolia Viv. are described. The level of seed dormancy is defined by the delay in seed germination (i.e the time required prior to germination) under favourable environmental conditions. A wild-type line shows a clear primary dormancy, which is suppressed by afterripening, whereas an abscisic acid (ABA)-deficient mutant shows a non-dormant phenotype. We have investigated the role of ABA and gibberellic acid (GA₃) in the control of dormancy maintenance or breakage during imbibition in suitable conditions. It was found that fluridone, a carotenoid biosynthesis inhibitor, is almost as efficient as GA₃ in breaking dormancy. Dry dormant seeds contained more ABA than dry afterripened seeds and, during early imbibition, there was an accumulation of ABA in dormant seeds, but not in afterripened seeds. In addition, fluridone and exogenous GA₃ inhibited the accumulation of ABA in imbibed dormant seeds. This reveals an important role for ABA synthesis in dormancy maintenance in imbibed seeds. Received: 31 December 1998 / Accepted: 9 July 1999     

Ecotypic variation of summer dormancy relaxation associated with rainfall gradient in the geophytic grass Poa bulbosa

Background and Aims

Summer dormancy is an adaptive trait in geophytes inhabiting regions with a Mediterranean climate, allowing their survival through the hot and dry summers. Summer dormancy in Poa bulbosa is induced by increasing day-length and temperature and decreasing water availability during spring. Populations from arid habitats became dormant earlier than those from mesic habitats. Relaxation of dormancy was promoted by the hot, dry summer conditions. Here we test the hypothesis that dormancy relaxation is also delayed in ecotypes of P. bulbosa inhabiting arid regions, as a cautious strategy related to the greater unpredictability of autumn rains associated with decreasing precipitation.

Methods

Ecotypes collected across a precipitation gradient (100–1200 mm year⁻¹) in the Mediterranean climate region were grown under similar conditions in a net-house in Israel. Differences among ecotypes in dormancy induction and dormancy relaxation were determined by measuring time to dormancy onset in spring, and time to sprouting after the first effective rain in autumn. Seasonal and ecotype variation in dormancy relaxation were assessed by measuring time to sprouting initiation, rate of sprouting and maximal sprouting of resting dry bulbs sampled in the net-house during late spring, and mid- and late summer, and planted in a wet substrate at temperatures promoting (10 °C) or limiting (20 °C) sprouting.

Key Results

Earlier dormancy in the spring and delayed sprouting in autumn were correlated with decreasing mean annual rainfall at the site of ecotype origin. Seasonal and ecotype differences in dormancy relaxation were expressed in bulbs planted at 20 °C. During the summer, time to sprouting decreased while rate of sprouting and maximal sprouting increased, indicating dormancy relaxation. Ecotypes from more arid sites across the rainfall gradient showed delayed onset of sprouting and lower maximal sprouting, but did not differ in rate of sprouting. Planting at 10 °C promoted sprouting and cancelled differences among ecotypes in dormancy relaxation.

Conclusions

Both the induction and the relaxation of summer dormancy in P. bulbosa are correlated with mean annual precipitation at the site of population origin. Ecotypes from arid habitats have earlier dormancy induction and delayed dormancy relaxation, compared with those from mesic habitats.