Factors influencing seed dormancy and germination in sand,and seedling survival under desiccation,of Psammochloa villosa (Poaceae), inhabiting the moving sand dunes of Ordos,China

Psammochloa villosa (Trin.) Bor. (Poaceae), is distributed primarily in moving sand dunes of the Ordos Plateau, China. Freshly harvested caryopses (seeds) are in non-deep physiological dormancy (non-deep PD). Germination is slow and low and only over a narrow temperature range. A treatment of four weeks cold stratification at 3 to 5 °C in darkness was required to break non-deep PD, allowing germination to reach high percentages at higher temperatures and without light requirement. Rate and percentages of germination were increased by scarifying the caryopsis coat and by artificial removal of different proportions of the endosperm. However, seedling dry weight and increases in root and shoot lengths, were significantly influenced by the proportion of the endosperm that remained on the caryopses. Higher percentages of seedling emergence were obtained from the shallowly buried caryopses, ranging from 0.5–2.0 cm, and the depth of the caryopses in the sand affected the time of germination. The deeper the caryopses were buried, the more that remained ungerminated and in enforced dormancy. The caryopses germinated when the upper sand layer was removed and buried caryopses were at a suitable sand depth for germination, or when the sand was aerated. In natural habitats, germinated seedlings may be wholly exposed to the air by sand erosion and thus be exposed to drought stress. However, young seedlings have the ability to tolerate desiccation and to recover after rehydration. Root length at the `point of no return' is 4 mm.     

Morphology of caryopses,seedlings and seedling emergence of the grass Calamovilfa longifolia from various depths in sand

Summary The spikelets of the grass Calamovilfa longifolia (Hook) Scribn. are one flowered and the dispersal unit is composed of a caryopsis (3.5 mm long and 0.9 mm wide) enclosed in lemma and palea. The highest germination of caryopses and emergence of seedlings occurred from 1–2 cm depths and seedling emergence decreased with increasing depth of burial. The maximum depth of sand from which a seedling can emerge is about 8 cm. Seedlings emerging from deep locations had first internodes which were more elongated than those of seedlings from more shallow plantings. Coleoptile lengths of seedlings from shallow or deeply buried caryopses were similar.     

沙埋和种子大小对固沙禾草沙鞭的种子萌发与幼苗出土的影响

该文研究了野外条件下不同深度的沙埋对沙鞭(Psammochloa villosa)种子萌发和幼苗出土的影响,以及温室条件下种子大小对不同深度沙埋后的种子萌发和幼苗出土的影响。结果表明,沙埋深度显著影响沙鞭的种子萌发率、幼苗出土率和种子休眠率。沙子表面的种子不能萌发。2 cm的浅层沙埋时的种子萌发率和幼苗出土率最高,1 cm 沙埋的种子萌发率和幼苗出土率次之。沙埋深度超过2 cm之后,沙鞭的种子萌发率和幼苗出土率与沙埋深度呈负相关。2 cm的种子休眠率最低。从2 ~12 cm,种子休眠率随着沙埋深度的增加而增加。在幼苗能够出土的深度(1~6 cm),幼苗首次出土所需的时间随着沙埋深度的增加而延长。种子大小对沙鞭的种子萌发率没有显著影响。但是在深层沙埋(6 cm)时,与小种子相比,大种子产生的幼苗的出土率较高。从2~6 cm,大种子形成的幼苗的茎长度都较长。     

Seed dormancy release and germination characteristics of <i>Corispermum lehmannianum</i> Bunge,an endemic species in the Gurbantunggut desert of China

Seed dormancy release and germination of Corispermum lehmannianum Bunge were tested using various treatments: temperature, cold stratification, gibberelins (GA3), dry storage and sand burial. Results showed that temperature and light did not affect the germination of fresh seeds, cold stratification and GA3 could improve seed germination, whereas dry storage and sand burial did not. The germination percentage was highest at 35/20 °C after the cold stratification and GA3 treatments. Corispermum lehmannianum seeds were classified as non-deep, Type-2, physiological dormancy (PD), whose seed dormancy could be released by cold stratification and GA3.     

Seed Dormancy,Seedling Establishment and Dynamics of the Soil Seed Bank of Stipa bungeana (Poaceae) on the Loess Plateau of Northwestern China

Studying seed dormancy and its consequent effect can provide important information for vegetation restoration and management. The present study investigated seed dormancy, seedling emergence and seed survival in the soil seed bank of Stipa bungeana, a grass species used in restoration of degraded land on the Loess Plateau in northwest China. Dormancy of fresh seeds was determined by incubation of seeds over a range of temperatures in both light and dark. Seed germination was evaluated after mechanical removal of palea and lemma (hulls), chemical scarification and dry storage. Fresh and one-year-stored seeds were sown in the field, and seedling emergence was monitored weekly for 8 weeks. Furthermore, seeds were buried at different soil depths, and then retrieved every 1 or 2 months to determine seed dormancy and seed viability in the laboratory. Fresh seeds (caryopses enclosed by palea and lemma) had non-deep physiological dormancy. Removal of palea and lemma, chemical scarification, dry storage (afterripening), gibberellin (GA₃) and potassium nitrate (KNO₃) significantly improved germination. Dormancy was completely released by removal of the hulls, but seeds on which hulls were put back to their original position germinated to only 46%. Pretreatment of seeds with a 30% NaOH solution for 60 min increased germination from 25% to 82%. Speed of seedling emergence from fresh seeds was significantly lower than that of seeds stored for 1 year. However, final percentage of seedling emergence did not differ significantly for seeds sown at depths of 0 and 1 cm. Most fresh seeds of S. bungeana buried in the field in early July either had germinated or lost viability by September. All seeds buried at a depth of 5 cm had lost viability after 5 months, whereas 12% and 4% seeds of those sown on the soil surface were viable after 5 and 12 months, respectively.     

Responses of Caryopsis Germination, Seedling Emergence, and Development to Sand Water Content of Agropyron cristatum (L.) Gaertn. and Bromus inermis Leyss.

Responses of caryopsis germination, seedling emergence, and development of Agropyron cristatum (L.) Gaertn. (Gramineae) and Bromus inermis Leyss. (Gramineae), two dominant perennial grasses in the Otindag Sandland of China, to different sand water content (SWC; 1%, 2%, 3%, 4%, 6%, 8%, 12%,16%, and 20%) were studied comparatively. The results showed that the germination responses of the two grasses to SWC were similar (i.e. caryopses could not germinate when the SWC was below 3%; at SWC ranging from 3% to 12%, the higher the SWC, the higher the germination percentage; and at a SWC of 12 %, germination reached similarly high percentages). At a sand burial depth of 0.5 cm, the threshold of SWC for seedling emergence was 6% forA. cristatum and 8% forB. inermis; at 12%-20% SWC, the seedling emergence of both species reached similarly high percentages. The seedling growth responses of these two species to SWC gradients were different. For A. cristatum, the biomass of seedlings increased with SWC from 6% to 12%, and decreased with SWC from 12% to 20%. For B. inermis, the biomass of seedlings always increased with SWC from 8% to 20%. The results also showed that the seedlings of both species allocated more biomass to the roots with decreases in SWC. The SWC changes from April to October in natural microhabitats of both species suggested that the SWC may play an important role in caryopsis germination,seedling emergence, and the growth characteristics of the two grasses. The responses of caryopsis germination, seedling emergence, and the growth characteristics of these two species to SWC may determine their distribution patterns in the Otindag Sandland.     

Responses of Caryopsis Germination, Seedling Emergence, and Development to Sand Water Content of Agropyron cristatum （L.） Gaertn. and Bromus inermis Leyss.

Responses of caryopsis germination, seedling emergence, and development of Agropyron cristatum （L.） Gaertn. （Gramineae） and Bromus inermis Leyss. （Gramineae）, two dominant perennial grasses in the Otindag Sandland of China, to different sand water content （SWC; 1%, 2%, 3%, 4%, 6%, 8%, 12%, 16%, and 20%） were studied comparatively. The results showed that the germination responses of the two grasses to SWC were similar （i.e. caryopses could not germinate when the SWC was below 3%; at SWC ranging from 3% to 12%, the higher the SWC, the higher the germination percentage; and at a SWC of 12%-20%, germination reached similarly high percentages）. At a sand burial depth of 0.5 cm, the threshold of SWC for seedling emergence was 6% forA. cristatum and 8% forB. inermis; at 12%-20% SWC, the seedling emergence of both species reached similarly high percentages. The seedling growth responses of these two species to SWC gradients were different. For A. cristatum, the biomass of seedlings increased with SWC from 6% to 12%, and decreased with SWC from 12% to 20%. For B. inermis, the biomass of seedlings always increased with SWC from 8% to 20%. The results also showed that the seedlings of both species allocated more biomass to the roots with decreases in SWC. The SWC changes from April to October in natural microhabitats of both species suggested that the SWC may play an important role in caryopsis germination, seedling emergence, and the growth characteristics of the two grasses. The responses of caryopsis germination, seedling emergence, and the growth characteristics of these two species to SWC may determine their distribution pattems in the Otindag Sandland.     

沙埋对无芒雀麦种子萌发和幼苗生长的影响

研究了沙埋对浑善达克沙地植物群落中多年生禾草无芒雀麦种子萌发、出苗和幼苗生长的影响.结果表明:无芒雀麦种子能在深度≤12cm的沙埋中萌发,≤8cm的沙埋中出苗.在此范围内,随着沙埋深度的增加,无芒雀麦种子的萌发率和出苗率逐渐降低.沙埋深度在植株高度的33%时,无芒雀麦1周龄和2周龄幼苗均全部存活,且总生物量≥对照(未沙埋幼苗);当沙埋深度增至植株高度的66%时,1周龄和2周龄幼苗的存活率分别降至70%和25%,生长也受到抑制;而遭受全部沙埋时,1周龄和2周龄幼苗均不能存活.遭受沙埋后,无芒雀麦幼苗分配较多的生物量用于地上部分的生长,其生物量分配模式改变可能是无芒雀麦幼苗对沙埋环境的重要适应对策.     

Achene germination of the spring ephemeroid species Carex physodes in the Gurbantunggut Desert

Much of the seed germination research on Carex has focused on wetland species, and little is known about the species of arid habitats. Here, we investigated seed dormancy and germination of Carex physodes, which is an important component of the plant community of the Gurbantunggut Desert of the Junggar Basin in Xinjiang, China. Our studies included the effects of mechanical and chemical scarification, dry storage, treatment with GA₃, wet‐cold stratification, and burial in the field. No freshly matured achenes germinated over a range of temperature regimes after treatment with GA₃, 6 months of dry storage or removal of part of the endosperm. The mechanical scarification resulted in < 5% achene germination, however, higher percentage of achene germination occurred after removal of the pericarp (60%), H₂SO₄ scarification (30%) or scarification in 10% NaOH (85%). Six and nine months of wet‐cold stratification promoted < 40% achene germination. The optimal germination temperatures ranged from 25/10°C to 35/20°C. Maximum germination after 9 months of burial at a depth of 3 cm in the field was 36%. These results indicate that the seeds have non‐deep physiological dormancy (PD) and that the pericarp contains germination inhibitors and has strong mechanical resistance to germination.     

沙埋对西鄂尔多斯珍稀植物种子萌发和幼苗出土的影响

采用不同沙埋深度(0、1、2、3、5、7、10 cm埋深)处理,对内蒙古西鄂尔多斯4种珍稀植物霸王(Zygophyllum xanthoxylum)、沙冬青(Ammopiptanthus mongolica)、蒙古扁桃(Prunus mongolica)和长叶红砂(Reaumuria trigyna)种子萌发及幼苗出土情况进行研究,以揭示其种子萌发特性.结果表明:表层沙土(0 cm埋深)的种子几乎不萌发;除霸王外,其他3种植物在不同沙埋深度处理下的种子萌发率差异显著(P＜0.05);沙埋对4种珍稀植物的幼苗出土均有显著影响,出苗率随着沙埋深度的增加而降低,且遵循指数方程.霸王在2 cm沙埋深度的出苗率最高,其他物种在1 cm沙埋深度的出苗率最高.沙埋深度对幼苗生物量的影响不显著.     

Germination,seedling occurrence and seedling survival of Spinifex sericeus R. Br. (Poaceae)

Abstract Spinifex sericeus is a clonal, perennial, dioecious, coastal foredune grass that is often used in beach rehabilitation programmes. The objectives of this study were to: (i) investigate the germination behaviour of S. sericeus; and (ii) examine the occurrence and survivorship of 5. sericeus seedlings in the dune systems on the mid-north coast of NSW. Diaspores and naked caryopses of S. sericeus require darkness for germination. Diaspore germination is adversely affected by leaching, probably due to the creation of anaerobic conditions around the caryopsis. Seedling emergence occurs mainly on the front of the foredune, as this is an area of active sand accretion and burial is necessary for germination. The presence of a hypocotyl allows germination and emergence from depths as great as 12.5 cm. In the two populations studied, seedling survival was low and probably made little contribution to population recruitment. The factors that appear to be responsible for the high mortality rate of seedlings are sand erosion, sand deposition and desiccation.     

Soil nitrification and foliar δ15N declined with stand age in trembling aspen and jack pine forests in northern Alberta,Canada

Aims

Seed germination and seedling emergence are vulnerable to water stress in arid environments. When precipitation is low and unpredictable during the early growing season, seeds near the sand surface often suffer from hydration/dehydration during germination. We investigated the responses of seedling emergence and survival of a sand dune grass with high sand stabilization value to amount and frequency of precipitation and depth of burial in sand.

Methods

Effects of amount and frequency of precipitation, burial and hydration/dehydration on seedling emergence of Leymus secalinus, were examined using standard procedures.

Results

Seedling emergence was affected by amount and frequency of monthly precipitation and depth of burial, and it decreased as precipitation frequency decreased with same amount of precipitation. Highest emergence percentage was obtained with 100 or 150 mm precipitation at 1–4 cm depth. Hydration/dehydration treatments decreased germination and increased dormancy percentage. Young seedlings with root lengths of 0–1 mm desiccated up to 30 days revived after rehydration.

Conclusions

Seedling emergence of L. secalinus is adapted to 150 mm monthly precipitation with frequency of 10–30 times per month, 1–4 cm burial depth and dehydration interval of 1–2 days. Alteration of amount and/or frequency of precipitation caused by climate change could markedly affect seedling emergence and population regeneration of this species.     

Effects of sand burial depth on seed germination and seedling emergence of Cirsium pitcheri

A greenhouse study was conducted to determine the effects of sand burial on seed germination and seedling emergence of Cirsium pitcheri, a threatened species along Lake Huron sand dunes. In October 1996, seeds of C. pitcheri were sorted into three groups (small, medium and large) and artificially buried at 2, 4, 6, 8, 10 and 12 cm depths in plastic pots filled with unsterilized sand from Lake Huron sand dunes. These pots were placed outside in sand boxes for natural stratification. In early February, the pots were brought into the greenhouse for germination of seeds. Data showed that percent seed germination and emergence of seedlings were not related to seed size. However, both variables were negatively correlated with depth. Seedling emergence occurred from a maximum depth of 6 cm with most seedlings emerging from 2 cm depth. Only one seedling emerged from 8 cm depth. Seedlings from large seeds produced longer roots than those of small seeds. Larger root system would probably enhance seedling establishment.     

Variation in dormancy and germination in three co-occurring perennial forest herbs

Mesic deciduous forest herbs often disperse seed with morphophysiological dormancy (MPD) that prevents germination during unfavorable periods for seedling survival. However, for seeds of some species with MPD, seasonal separation of root and shoot emergence and variation in dormancy levels can complicate interpretation of seedling emergence timing in the field. We tested whether dormancy-break and germination requirements differed among co-occurring perennial forest herbs, Actaea racemosa, Hydrastis canadensis, and Sanguinaria canadensis, which are wild-harvested for their medicinal properties and known to have MPD. Seeds of all species exhibited a summer → autumn → winter requirement for seedling emergence in spring. However, species differed in seed-bank persistence due to variation in primary dormancy levels and stratification requirement of seeds. A. racemosa and H. canadensis can form short-term persistent seed bank, whereas S. canadensis can form a long-term persistent seed-bank, regardless of whether elaiosomes were removed from seeds prior to burial. A. racemosa seeds are dispersed in autumn with weak physiological dormancy, as seeds germinated to high rates at 15/6°C after 8 weeks. In contrast, most seeds of the summer dispersed species, H. canadensis and S. canadensis, require summer temperatures to overcome physiological dormancy. Consequently, seedling emergence is reduced and delayed by 1 year if seeds are not sown immediately following the period of natural dispersal. Seedling emergence was much lower in the field than in controlled conditions for all species, especially in the small-seeded A. racemosa. Interspecific variation in dormancy levels and germination traits must be considered when establishing populations for conservation purposes and in understanding recruitment limitation in perennial forest herbs.     

Sequential temperature control of multi-phasic dormancy release and germination of Paeonia corsica seeds

Aims The physiological responses during dormancy removal and multi-phasic germination were investigated in seeds of Paeonia corsica (Paeoniaceae).Methods Seeds of P. corsica were incubated in the light at a range of temperatures (10–25 and 25/10°C), without any pre-treatment, after W (3 months at 25°C), C (3 months at 5°C) and W + C (3 months at 25°C followed by 3 months at 5°C) stratification, and a GA 3 treatment (250 mg·l^-1 in the germination substrate). Embryo growth, time from testa to endosperm rupture and radicle emergence were assessed as separate phases. Epicotyl–plumule emergence was evaluated incubating the germinated seeds at 15°C for 2 weeks, at 5 and 25°C for 2 months on agar water before transplanting to the soil substrate at 10, 15 and 20°C and at 15°C for 2 months on the surface agar water with GA 3 .Important findings Embryo growth, testa rupture, endosperm rupture (radicle emergence) and growth of the epicotyl were identified as four sequential steps in seeds of P. corsica. Gibberellic acid alone and warm stratification followed by 15°C promoted embryo growth and subsequent seed germination. Cold stratification induced secondary dormancy, even when applied after warm stratification. After radicle emergence, epicotyl–plumule emergence was delayed for ca. 3 months. Mean time of epicotyl–plumule emergence was positively affected by cold stratification (2 months at 5°C) and GA 3. P. corsica seeds exhibited differential temperature sensitivity for the four sequential steps in the removal of dormancy and germination processes that resulted in the precise and optimal timing of seedling emergence.     

Sand burial effects on seed germination, seedling emergence and establishment of Panicum virgatum

Studies in the field and in a greenhouse were conducted to examine the effects of sand burial on seed germination, seedling emergence and establishment of Panicum virgatum L. on the foredunes of Lake Erie. Under natural conditions, the seedlings emerged from sand burial depths ranging from 0 to 11 cm, with a mean ± SD of 4.73 ± 1.82 cm. The frequency distribution of the depth of emergence of seedlings in the field was significantly skewed to the right and platykurtic. In the greenhouse, some seedlings emerged from a burial depth of as much as 16 cm. Although percent germination of seeds was not affected by sand burial, the percent emergence and the rate of emergence of seedlings were significantly reduced by excessive sand burial. Seedling mortality was found only among seedlings that emerged from sand burial depths of 10 cm or more. In the field, all the seedlings established in one growing season had originally emerged from sand burial depths of less than 12 cm. Within this burial range, seedlings from shallower burial depths had lower chances of establishment than expected, whereas those from deeper burial depths had higher probabilities of establishment than expected.     

Effects of partial endosperm removal on embryo dormancy breaking and salt tolerance of <Emphasis Type="Italic">Hordeum spontaneum</Emphasis> seeds

Seed damage is a common phenomenon in nature and in agricultural production. In this experiment, partial endosperm removal from wild barley (Hordeum spontaneum) caryopses, sampled from three ecotypes originated from xeric environments in Israel, was conducted. The aim was to examine seed dormancy and germination states in damaged caryopses and salt tolerance of young seedlings derived from them. Six treatments were made: (1) control seeds with intact caryopses; (2–4) removal of 0.25, 0.5, and 0.75 of the length of intact caryopses; (5) transection at the points, at which the endosperm and embryo meet; and (6) slitting of endosperm opposite the embryo. A significant negative correlation was found between germination percentage (dormancy release) and the relative distance from the dissection point to embryo. Partial removal of the endosperm could accelerate dormancy release. Seedling salt tolerance was assessed by the ratio of root or coleoptile length in a seedling grown in 100 or 200 mM NaCl solution to that of a seedling grown in water. The seedling salt tolerance was positively correlated with the removed portion of the seed endosperm. For each level of endosperm removal, the salt tolerance to 200 mM NaCl of the seedlings derived from the Dead Sea ecotype was higher than those from both the Sede Boker and the Mehola ecotypes. The results suggest that partial damage to seed endosperms in natural conditions may play a role in increasing the phenotypic plasticity of germination and salt tolerance.     

Germination and emergence of annual species and burial depth: Implications for restoration ecology

Due to the high content of viable seeds, topsoil is usually spread on ground left bare during railway and motorway construction to facilitate the regeneration of vegetation cover. However, during handling of the topsoil, seeds are often buried deeply and they cannot germinate or the seedlings cannot emerge from depth. This study experimentally explores the predictive value of seed mass for seed germination, mortality and seedling emergence at different burial depths for 13 common annual species in semiarid Mediterranean environments. We separate the effect of burial depth on germination and emergence by means of two experiments. In the germination experiment, five replicates of 20 seeds for each species were buried at depths ranging from 0 to 4 cm under greenhouse conditions. Germinated and empty or rotten seeds were counted after 8 weeks. In the emergence experiment, five replicates of four newly-germinated seeds per species were buried at the same depths under controlled conditions and emergence was recorded after 3 weeks. The effect of burial depth on percentage of germination and seedling emergence was dependent on seed size. Although all species showed a decrease in germination with burial depth, this decrease was greater for small-than large-seeded species. Percentage of emergence was positively related to seed mass but negatively related to burial depth. Seed mortality was higher for small-than large-seeded species, but there was no general effect of burial depth on this variable. Thus, the current practice of spreading 30 cm deep layers of topsoil in post-construction restoration projects is unadvisable. In this restoration scenario, thinner layers of topsoil should be used to achieve the maximum potential of the topsoil for germination and seedling establishment.     

Seed germination studies with Heteropogon contortus

Germination characteristics of Heteropogon contortus were investigated in germination cabinets and in the glasshouse using soil trays. Under the former a dual inhibitory system appeared to operate. On the one hand naked caryopses from seeds which were less than 4 months of age were inhibited from germination by a process apparently analogous to epicotyl dormancy. This could readily be broken by applying 1% gibberellic acid and partly broken by repeated wetting and drying. On the other hand between 4 and 12 months, naked caryopses germinated more readily than intact seeds (caryopsis plus glumes, lemmas, etc.). However, the dormancy factor apparently associated with these structures could not be detected when seed was germinated in soil in the glasshouse and it appeared that they are a necessary protection for the caryopsis under field conditions. The optimal temperature for germination was between 30 and 35°C constant. Alternating diurnal temperatures of less than this appeared to cause reduced germination, and no high-temperature stimulation was observed. Germination was not directly promoted by the passage of a normal grass fire.     

Germination in Zostera japonica is determined by cold stratification, tidal elevation and sediment type

The effects of water temperature and bottom sediment type were studied on seed dormancy and germination of Zostera japonica Ascherson & Graebner in mesocosm. To test whether the germination rate is affected by cold stratification, seeds were divided into two groups: those exposed to cold (7 °C) and those left untreated (23–15 °C). Additionally, to mimic tidal variation, we used five tidal depth treatments for germination experiments in mesocosm. In mesocosm tanks, there was a wide range of daily fluctuating temperature at datum line +40 cm (17–25 °C), D.L. +20 cm (15 °C), and D.L. +0 cm (4–7 °C). In contrast, the maximum temperature range at D.L. −20 cm and −40 cm was narrow (5–6 °C). In the no cold stratification group, the maximum germination rates on sandy, muddy sand, and muddy bottom sediment were 3%, 11%, and 3%, respectively. In the cold stratification group (7 °C), the maximum germination rates were 40%, 53%, and 54%, respectively. First germination was observed 36 ± 0 days and 43 ± 6 days after the start of the germination experiment in the cold stratification group and the no cold stratification group, respectively. Bottom sediment type and tidal level did not affect seed germination in the both stratification group. Cold stratification strongly increases germination in all sediment types tested and under varying temperature regimes and at different tidal levels. We also tested whether seed germination is affected by daily fluctuations in temperature (10 °C constant, 15 °C/10 °C, and 20 °C/10 °C were compared) in an indoor incubator. Forty-two days after being sown, the maximum seedling emergence rates in the three groups were 3 ± 5%, 21 ± 7%, and 42 ± 26%, respectively. At 20 °C/10 °C, first germination was observed 11 days after the start of incubation, the germination rate rose sharply after 18 day of incubation, and then it leveled off after 32–42 days of incubation. In the no cold stratification group, seed germination was not observed in any of the three treatments. This finding suggests that the breaking of seed dormancy and germination of Z. japonica seeds are determined strongly by cold temperature and daily fluctuations of temperature, respectively.