Annual cycles of germinability and differences between primary and secondary dormancy in buried seeds of Echinochloa crus-galli

The seasonal changes in percentage of dormant seeds of Echinochloa crus-galli in the field were recorded for 4 years. The lots of seeds were wrapped in nylon fabric, buried 20 cm under the grass sward and exhumed at monthly intervals. The proportion of seeds germinating under light conditions at a constant temperature of 25 °C fluctuated between 0% and 96%, with maxima in May–July and minima in September–November. Small between-year differences in the course of summer dormancy induction and its winter termination were probably caused by variation of weather conditions.
Attributes of dormancy innate to seeds after maturation (primary dormancy) and dormancy induced in buried seeds during the summer (secondary dormancy) were compared by investigating the rate of dormancy termination during storage of (a) dry seeds at 25 °C, (b) imbibed seeds at 5°C and (c) in seeds buried under field conditions during October–June. Percentage of germination increased faster in secondary than primary dormant seeds at both constant 25 °C and 5 °C. The seeds with primary and secondary dormancy also differed in the response to `germination pre-treatment', a 10-day exposure of imbibed seeds at 25 °C that causes germination of the non-dormant fraction of seed materials. After this treatment the time to resuming germination in primary dormant seeds was substantially increased, whereas the secondary dormant seeds were much less affected. Annual variation in the proportion of germinable seeds explains the low efficiency of autumn soil cultivation for decreasing reserves of E. crus-galli seeds in the soil.     

Non-deep simple morphophysiological dormancy in seeds of the weedy facultative winter annual Papaver rhoeas

Embryos in freshly matured seeds of the facultative winter annual Papaver rhoeas are underdeveloped and physiologically dormant; thus, seeds have morphophysiological dormancy (MPD). Seeds lost physiological dormancy during 12 weeks of burial in moist soil at 12 h/12 h daily alternating temperature regimes of 15/5°C, 20/10 °C and 25/15 °C but not at 1 °C. Physiological dormancy was not broken in seeds stored dry at room temperature for 12 weeks. After physiological dormancy was broken, seeds required light for embryo growth (i.e. for loss of morphological dormancy) and consequently for germination. After a 12-week period of burial in soil at 25/15 °C, seeds that matured in 1997 germinated to 100% in light at 25/15 °C, demonstrating that cold stragification temperatures (≈ 0.5–10 °C) are not required for embryo growth. Thus, seeds have non-deep simple MPD. During exposure to low winter temperatures (5/1 °C, 1 °C), 52% of the seeds with physiologically non-dormant embryos entered conditional dormancy and thus lost the ability to germinate at 25/15 °C but not at 15/5 °C or 20/10 °C. The peak of germination for seeds sown in southern Sweden was in autumn, but some also germinated in spring. A higher percentage of seeds that matured in a relatively warm, dry year (1997) came out of MPD and germinated than did those that matured in a relatively cool, wet year (1998) at the same site.     

Germination, emergence, and dormancy of Mimosa pudica

Mimosa pudica (common sensitive plant) is a problematic weed in many crops in tropical countries. Eight experiments were conducted to determine the effects of light, seed scarification, temperature, salt and osmotic stress, pH, burial depth, and rice residue on the germination, seedling emergence, and dormancy of M. pudica seeds. Scarification released the seeds from dormancy and stimulated germination, though the germination of the scarified seeds was not influenced by light. The scarification results indicate that a hard seed coat is the primary mechanism that restricts germination. The germination increased markedly with the exposure to high temperature "pretreatment" (e.g. 150°C), which was achieved by placing non-scarified seeds in an oven for 5 min followed by incubation at 35/25°C day/night temperatures for 14 days. The germination of the scarified seeds was tolerant of salt and osmotic stress, as some seeds germinated even at 250 mmol L⁻¹ NaCl (23%) and at an osmotic potential of −0.8 MPa (5%). The germination of the scarified seeds was >74% over a pH range of 5–10. The seedling emergence of the scarified seeds was 73–88% at depths of 0–2 cm and it gradually decreased with an increasing depth, with no seedling emergence at the 8 cm depth. The rice residue applied to the soil surface at rates of ≤6 t ha⁻¹ did not influence the seedling emergence and dry weight. The information gained from this study identifies some of the factors that facilitate M. pudica becoming a widespread weed in the humid tropics and might help in developing components of integrated weed management practises to control this weed.     

Role of temperature in regulating timing of germination in soil seed reserves of Thlaspi arvense L.

Summary. Most freshly-matured seeds of Thlaspi arvense L. (Brassicaceae) were dormant at maturity in May. Seeds sown on soil germinated in autumn and spring, but mostly in autumn. Buried seeds exhumed at monthly intervals and tested in light and darkness over a range of thermoperiods exhibited annual dormancy/non-dormancy cycles. However, the dormant period was short, usually only in April, but sometimes May, and in some years 1–6% of the seeds remained conditionally dormant. After-ripening occurred during summer, and seeds were non-dormant during autumn. Seeds entered conditional dormancy in winter and dormancy in late winter or early spring. When buried dormant seeds were kept at 25/15, 30/15 or 35/20°C for 12 weeks, they gained the ability to germinate to 95–100% at 15/6, 20/10, 25/15, 30/15 and 35/20°C. After burial for 12 weeks at 15/6 and 20/10°C, seeds germinated to 80–100% at 15/6, 20/10 and 25/15°C. but to only 11–64% at 30/15 and 35/20°C. After 4 weeks at 5°C, initially-dormant seeds germinated to 100% at all thermoperiods except 35/20°C, where only 15% of them germinated. However, after 18 weeks at 5°C, only 0–1% of the seeds germinated at all thermoperiods. Most non-dormant seeds exposed to 1, 5 and 15/6°C for 16 weeks were induced into dormancy; 1–15% entered conditional dormancy and thus germinated only at 15/6, 20/10 and 25/15°C. This study indicates that seeds of winter annual plants of T. arvense are non-dormant in autumn and enter dormancy in winter, while those from summer annuals are dormant in autumn and become non-dormant during winter.     

UNTERSUCHUNGEN ZUR BIOLOGIE UND ÖKOLOGIE DER HÜHNER-HIRSE ECHWOCHLOA CRUS-GALLI L. BEAUV.

Summary. 1. Echinochloa crus-galli is a typical grass weed of root crops in warmer regions, germinating late and thus covering the soil at a late stage. It originates in central and east Asia and is now a weed of world-wide importance.
2. High temperatures are necessary for germination which begins in spring, but not before the soil temperature reaches 15° C. Minimum, optimum and maximum germination temperatures of 13° C, 20–30° C and 40° C were found in S.W. Germany.
3. Seeds of E. crus-galli are dormant during the first 3–4 months after harvest. Those developing relatively early in the growing season require rather longer for after-ripening than seeds which mature later.
4. For optimum germination, water saturation of the soil of 70–90% is required.
5. Soil acidity has some influence, and there is an apparent germination optimum around neutrality. Light also induces germination.
6. Seeds can emerge from a relatively wide range of depths. Greatest emergence and the strongest plants resulted from seeds at 2–6 cm, but even from 10 cm a high percentage of seedlings is likely to emerge.
7. Further development proceeds rapidly. The first panicles arc already formed 6–7 weeks after emergence in favourable conditions, but full maturity is possible only if there are high temperatures in late summer.
8. E. crus-galli is a hygrophilous species, with best development on medium heavy soil, sandy loam or loamy sand with sufficient water supply.
9. E. crus-galli is indifferent to the lime content of the soil.
10. Best development occurs with high fertility, and a rich supply of nitrogen is especially important.
Recherches sur la biologie et écologie du panic Echinochloa crus-galli (L.) Beauv.     

Light, temperature and burial depth effects on Rumex obtusifolius seed germination and emergence

Trials were carried out to investigate the effects of light and temperature on germination of Rumex obtusifolius L. After several months of storage, seeds gradually lost dormancy and became photosensitive. Thermal optima for germination were between 20 °C and 25 °C in light or in darkness. At lower temperatures there was a greater demand for light, so that the greatest differences in germination percentage (between low and high temperatures) were found within the 10–15 °C temperature range. The calculated thermal minima ( x -intercept method) in light and darkness were 8.3 °C and 6.1 °C respectively. Daily temperature fluctuation increased germination even after seed irradiation with far-red light, suggesting a lower demand for the far-red-absorbing form of phytochrome. Seed burial inhibited germination in proportion to depth; however, germination inhibition was independent of seed phytochrome photo-equilibrium, which had been diversified by seed pretreatment with light. Seedlings did not emerge when seeds were buried >8 cm deep. Recovery of ungerminated seeds showed that excessive burial did not impede seedling emergence but rather prevented seed germination. However, this induction of dormancy was lost once germination processes were activated (24–48 h at 20 °C) that made germination irreversible. Temperature was also involved in inhibition, and low temperature (<15 °C) induced the least inhibition. This is discussed in terms of processes of respiration and fermentation in buried seeds.     

Effect of soil temperature on disease development in melon plants infected by Monosporascus cannonballus

The effect of soil temperature on melon collapse induced by Monosporascus cannonballus was studied in the laboratory and in the field. In the laboratory, ascospore germination and hyphal penetration into melon roots were enhanced by increasing the temperature from 20 to 32°C. The optimum temperature for mycelial growth of five isolates of M. cannonballus was 30°C. In the field, the effect of temperature was studied in experiments conducted during the winter and autumn cropping seasons from 1995 to 1998. Disease progress was much faster in the autumn than in the winter crop seasons. Disease incidence reached 100% in the three consecutive autumn seasons studied. In the winter seasons, however, planting date influenced disease incidence. Early planting, at the beginning of January, resulted in a low disease incidence (6–26%, 125 days after planting), whereas planting at the end of January resulted in higher disease incidence (72–88%, 95–119 days after planting). In plots in which the soil was artificially heated to 35°C during the winter season, disease incidence reached 85%, as in the autumn season. Plants grown during the winter in unheated soil, or in artificially heated soil disinfected with methyl bromide, did not collapse. Root colonization by the pathogen was higher in the autumn and in heated soil than in the winter season in nonheated soil. Fifty per cent of root segments were colonized 35, 42 and 67 days after planting in the winter-heated, autumn and winter-unheated plots, respectively. A high correlation was found between soil temperatures above 20°C during the first 30 days after planting and disease severity. It is suggested that soil temperature during the early stages of plant development is an important factor in disease development and the expression of melon collapse caused by M. cannonballus.     

Environmental control of dormancy and germination in the seeds of Cenchrus longispinus (Hack.) Fern.

Seed dormancy and germination in sand burr (Cenchrus longispinus (Hack,) Fern,) were investigated in laboratory and field studies. The burrs contain two types of seeds which differed in their innate dormancy. Primary seeds formed in the upper spikelet usually germinated within a year. Secondary seeds from lower spikelets germinated slowly and remained dormant for longer periods. Dormancy was enforced at low and high temperatures, and secondary seeds apparently developed an induced dormancy when continuously exposed to high temperatures. More than 94 % of the seedlings established during spring. Light suppressed germination, and secondary seeds also developed an induced dormancy when stored in the light. Burrs sown on the soil surface had an extended period of germination lasting for more than 3 years. However, over 96 % of the seeds sown below the surface of bare soil germinated within 2 years. Deep burial did not enforce dormancy, but germination was suppressed by the presence of live vegetation. It is concluded that treatments which disturb the soil and bury the burrs will stimulate the germination of dormant seeds.     

Solarization and natural heating of irrigated soil amended with cruciferous residues for improved control of Macrophomina phaseolina

The efficacy of summer irrigation and soil solarization combined with cruciferous residues was tested against the dry root rot pathogen Macrophomina phaseolina in an arid climate. In irrigated amended soil, polyethylene mulching during May increased the soil temperature to 57°C and 50°C at depths of 0–15 and 16–30 cm, respectively. As a result, within l5 days the population of M. phaseolina was almost eradicated (93–99%) at both soil depths. A considerable reduction (75–96%) was also achieved by natural heating of irrigated soil (46–53°C) for l5 days after amending with cruciferous residues. Mulching alone was only partially effective (69–89% reduction). These results suggest a new approach to controlling soil-borne pathogens in hot, arid regions by combining summer irrigation with soil amendment. Amendment with residues alone or in conjunction with soil solarization also increased the population of lytic bacteria against M. phaseolina .     

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.     

Seed dormancy dynamics and germination characteristics of Solanum nigrum

Four experiments were conducted to study seed dormancy and germination requirements in Solanum nigrum . In Expt 1, seeds were stratified at different constant and stepwise rising temperatures and their germinability was tested at three germination regimes at weekly intervals. In Expts 2–4, seeds dry stored at 4°C and stratified at 5 and 15°C were tested at constant temperatures, as well as fluctuating temperatures with constant and increasing amplitudes. Results suggest that the rate of dormancy release increased with increasing temperatures ranging from 4.5 to 18.6°C. However, prolonged stratification at higher temperatures caused subsequent induction of dormancy. When tested at constant temperatures, stratified seeds germinated between 18 and 34°C, with the optimum between 26 and 30°C, while dry-stored seeds showed no germination. Fluctuating temperatures, with amplitudes ranging from 5 to 15°C, promoted germination of seeds from all treatments. The dormancy dynamics and germination characteristics of the species will have implications for its survival and establishment. This information can be used to predict time of emergence and, thus, improve control of the species in weed management systems.     

Ecological aspects of seed dormancy-break and germination in Heteranthera limosa (Pontederiaceae), a summer annual weed of rice fields

Summary Heteranthera limosa seeds were buried in flooded and in non-flooded soil and exposed to natural seasonal temperature changes in Lexington, Kentucky, USA. Seeds exhumed after various periods of burial ranging from 2 to 36 months were tested for germination under both flooded and non-flooded conditions. Seeds were dormant at maturity in September and became non-dormant during winter. Seeds buried in non-flooded soil during winter germinated to higher percentages and over a wider range of temperatures when tested under flooded conditions (in light) during spring and summer, than did those buried in flooded soil during winter. Thus, the water regime associated with rice culture (non-flooded in winter and flooded in summer) is optimal for dormancy-break and germination of H. limosa seeds. A portion of the buried seeds exhibited an annual dormancy/non-dormancy cycle, whereas others had a conditional dormancy/non-dormancy cycle. Regardless of the type of cycle, seeds buried in non-flooded soil retained the ability to germinate in light at high temperatures under flooded conditions throughout the summer. Thus, seeds potentially can germinate at any time during the growing season, whenever rice fields are flooded. Flooding fields during winter and/or sowing rice relatively early in the growing season may help in establishing rice before seeds of H. limosa germinate.     

Biology and management of Phalaris minor in wheat under a rice/wheat system

Summary Germination of Phalaris minor declined with the increase in duration of imbibition in water from 30 min to 72 h at temperatures above 22 °C . Germination was reduced down to 10 cm and 2 cm soil depth by wheat straw burning in puddled and non-puddled soil, respectively, with maximum reduction near the soil surface. The dormancy of P . minor seed was not more than 60 days under field conditions. In puddled soil, 38–60% of the viable seeds of P. minor remained concentrated in the upper 5-cm layer. Germination decreased with an increase in soil depth. In total, 15% of seeds stored in the laboratory emerged from 10-cm depth, whereas seeds did not germinate below 4.2-cm depth under field conditions. Depth of emergence of P . minor was shallower in zero tillage compared with the conventional method of wheat sowing. The seeds retrieved from rice soils kept under continuous submergence for 60 days exhibited 26% and 57% loss of germination over semi-submergence and semi-wet conditions respectively. There was 100% loss of germination in 10-month-old seeds retrieved from the soil under rice-growing conditions. Plant density of P. minor was lower in zero tillage than with the conventional method of wheat sowing. Cross-ploughing in the upper 2–5 cm of soil (shallow tillage) and drill-sowing of wheat 1 week after shallow tillage reduced germination of P. minor by 44% and 37% and increased grain yield by 21% and 47% over zero-tillage and conventional methods respectively.     

Invasiveness evaluation of fireweed (Crassocephalum crepidioides) based on its seed germination features

In China, fireweed ( Crassocephalum crepioides ) is listed as an invasive plant that is also cultivated as a vegetable. To gain a clearer understanding of its invasiveness and rapid spread, we evaluated its seed dispersal ability, and the influences of light, temperature, pH, NaCl stress, moisture content, and storage periods on its seed germination. Its seed dispersal ability is limited. The seed germination of fireweed is inhibited by darkness, temperatures <10°C or >35°C, and a NaCl solution with a concentration >0.15 mol L⁻¹. The optimal conditions under which nearly all the seeds could germinate are light, with temperatures from 20 to 30°C, and a neutral soil with 40% moisture content. The seeds of fireweed have no apparent dormancy and retain a high viability after room storage for 10 months. Fireweed only has a moderate invasive capacity and its wide distribution in China possibly correlates with its cultivation.     

Germination characteristics of Amaranthus quitensis as affected by seed production date and duration of burial

Thermal requirements for the germination of Amaranthus quitensis, a common annual weed in Argentina, were studied. In addition, temporal changes in dormancy from seeds produced at different times during the growing season were examined. For this second objective, thermal and light requirements for germination were tested in seeds buried at different depths, with or without crop residues. Base and optimum temperatures for germination rates were 12.8°C and 37°C respectively. At dispersal time, maximum percentage germination was 60–70% and this was generally recorded at 35°C/25°C in a 14-h photoperiod. Seed germination tended to increase in later seed collection dates. Seeds of A. quitensis showed seasonal changes in germinability in the soil. In winter, germination of retrieved seeds increased to over 90% until summer, after which there was a decrease until the following winter when germination was close to 40%. There were no differences in germinability between burial depths and crop residue levels. Germination requirements for alternating temperatures and light tended to disappear after burial. Initial viability was 99% and declined slightly during burial. Soil temperature seems to play a crucial role not only by regulating seasonal changes in dormancy, but also by defining the percentage and the germination rate in non-dormant seeds.     

不同季节强碱土土壤呼吸影响因子分析与模型预测

利用LI-8100土壤碳通量测量仪测定了春夏秋三季晴朗天气下强碱土土壤呼吸速率、温度(气温和地温)、湿度(空气相对湿度和土壤湿度)数据,分析了它们之间的相关关系,获得不同季节对土壤呼吸影响较大的因子,并建立不同类别的多种回归模型;在精度检验及简单易行原则基础上,得到各季节土壤呼吸预测的最优模型。结果表明:(1)虽然温湿度均是影响不同季节强碱土土壤呼吸的主要因素,但均以温度的影响较大,其中气温是春秋两季土壤呼吸的最大直接影响因子,地温是夏季的最大直接影响因子,而土壤湿度为各季节最大的间接影响因子。(2)春秋季土壤呼吸的最佳预测模型均为10 cm处气温和土壤湿度所建的双因子方程,该方程具有较小的均方根误差(RMSE)(分别为0.159和0.259),且相对分析误差(RPD)2(分别为2.9、2.094),具有非常好的预测能力。夏季土壤呼吸最佳预测模型则为包含10 cm处气温、地温、空气相对湿度和土壤湿度所建的4因子方程,RMSE为0.248,RPD2(为2.406),可用于精确预测。(3)各季节土壤呼吸变化趋势与其影响因子的变化,因春季的完全同步,夏季基本一致,而秋季一致性较差,故春季土壤呼吸最佳预测模型的预测精度最高(92.67%),夏季次之(84.99%),秋季较差(77.23%)。     

干旱半干旱地区双垄地布覆盖对土壤水分的影响

为探明一种可替代覆盖材料的可行性,在甘肃省定西市安定区开展了为期1 a的监测试验,试验设防草地布加地膜覆垄(MB)、防草地布覆垄(DB)和裸地起垄(CK)3种处理,利用土壤水分、水势传感器分别对沟内地表下5、15 cm的土壤含水率、水势进行监测,结果表明:(1)表层5 cm土壤水分日变化呈复杂波形,受不同覆垄处理和季节性天气变化的影响显著;(2)0～20 cm土层的水分年变化主要受降雨、露水和蒸发强度的影响,表现为春冬干、夏秋湿的特点,在11月至翌年2月期间MB和DB覆垄处理土壤水分净损失量要高于CK裸地垄,而在作物生育期内(5—9月)覆垄处理土壤储水净增量为DB最大(36.35 mm)、MB次之(30.73 mm)、CK最小(16.30 mm);(3)MB和DB覆垄能明显加快雨露叠加,增加土壤水储存,而CK处理下叠加效应弱,且深层土壤对降雨不敏感,具有滞后性,但随着连续降雨的发生,表层土壤储水量加大,这种滞后性明显减弱;(4) 0～20 cm土层土壤储水量日变化幅度为夏季最大(平均1.20 mm·d~(-1)),春季次之(1.03 mm·d~(-1)),秋季最小(0.79 mm·d~(-1)),全年水分净收获总量为DB最大(24.9 mm)、MB略低(21.5 mm)、CK最小(11.4 mm)。整体而言,只用地布覆垄具有最好的集水效果,这种防草地布可多年使用,无地膜碎片化污染问题,但在无降雨时期,垄沟内因土壤水分高,其蒸发强度略高于裸地处理。     

Purple nutsedge tuber sprouting

Purple nutsedge ( Cyperus rotundus L.) tubers remain viable for several years and serve as its principal means of survival. The maintenance of high moisture content is essential to tuber survival. Apical dominance influences bud dormancy within a tuber and in a chain of tubers, and dormancy increases with tuber age. Several growth inhibitors were identified in tubers, but their role in tuber dormancy has not been established. Moisture levels in soil must increase to a critical level before sprouting occurs, but excess soil moisture deters sprouting. Oxygen may be a limiting factor for tuber sprouting in waterlogged soils. Although light is not a requirement for sprouting, it has promoted sprouting. Temperature regulates sprouting; no sprouting occured below 10°C and above 45°C. Optimum sprouting occurred between 25 and 35°C when provided with constant temperatures. However, daily alternating temperatures greatly stimulated sprouting. A daily short duration (0.5 h) of high temperature increased sprouting to nearly 100%, whereas less than 50% sprouting occurred without the daily high temperature pulse. Bud break occurred readily for most tubers at 20°C and in nearly 100% of the tubers with a single 0.5 h exposure to a high temperature (35°C) pulse. However, most buds did not elongate if the tuber remained at 20°C. Bud elongation occurred at higher temperatures, and daily alternating temperatures stimulated shoot elongation up to eightfold greater than at the respective mean constant temperatures. Daily soil temperature fluctuation may be a major signal for purple nutsedge emergence, such as when the plant canopy is removed, or when soils are solarized. Future research is needed to determine tuber sprouting for different ecotypes, and on the role of the rhizome chain. Systems to manipulate sprouting may provide new strategies for purple nutsedge management.     

Variation in the development of secondary dormancy in oilseed rape genotypes under conditions of stress

Summary A substantial amount of seed is left in the fields before and during harvest of oilseed rape. Although this crop exhibits little or no primary dormancy, the absence of certain environmental cues that promote germination of imbibed seeds induces secondary dormancy. The work reported investigated the extent to which environmental stress conditions, including osmotic stress, low oxygen stress and anaerobiosis, induce secondary dormancy in oilseed rape, and examined the variation in development of secondary dormancy between and within genotypes. Osmotic stress was most effective in inducing dormancy. Anaerobic treatment produced very few dormant seeds, as did an atmosphere low in oxygen and high in nitrogen. The development of secondary dormancy under osmotic stress varied considerably between and within genotypes. Dormancy ranged from almost zero to about 60% for winter genotypes and about 85% for spring types. Within genotypes, variations occurred between seed lots and years of harvest. Temperature variations affected the percentage of dormant seeds. More dormant seeds were likely to be produced with incubation under water stress at 20 °C than at 12 °C. In winter genotypes, fewer dormant seeds were produced when incubation temperature and germination test temperatures differed. Thus, incubating at 20 °C and 12 °C, followed by germination tests at 20 °C and 12 °C, respectively, produced most dormant seeds. Also, in the winter genotypes, the potential development of secondary dormancy was positively correlated with the pattern and speed of germination of untreated seeds.