Acquisition of physical dormancy and ontogeny of the micropyle--water-gap complex in developing seeds of Geranium carolinianum (Geraniaceae)

Background and Aims

The ‘hinged valve gap’ has been previously identified as the initial site of water entry (i.e. water gap) in physically dormant (PY) seeds of Geranium carolinianum (Geraniaceae). However, neither the ontogeny of the hinged valve gap nor acquisition of PY by seeds of Geraniaceae has been studied previously. The aims of the present study were to investigate the physiological events related to acquisition of PY and the ontogeny of the hinged valve gap and seed coat of G. carolinianum.

Methods

Seeds of G. carolinianum were studied from the ovule stage until dispersal. The developmental stages of acquisition of germinability, physiological maturity and PY were determined by seed measurement, germination and imbibition experiments using intact seeds and isolated embryos of both fresh and slow-dried seeds. Ontogeny of the seed coat and water gap was studied using light microscopy.

Key Results

Developing seeds achieved germinability, physiological maturity and PY on days 9, 14 and 20 after pollination (DAP), respectively. The critical moisture content of seeds on acquisition of PY was 11 %. Slow-drying caused the stage of acquisition of PY to shift from 20 to 13 DAP. Greater extent of cell division and differentiation at the micropyle, water gap and chalaza than at the rest of the seed coat resulted in particular anatomical features. Palisade and subpalisade cells of varying forms developed in these sites. A clear demarcation between the water gap and micropyle is not evident due to their close proximity.

Conclusions

Acquisition of PY in seeds of G. carolinianum occurs after physiological maturity and is triggered by maturation drying. The micropyle and water gap cannot be considered as two separate entities, and thus it is more appropriate to consider them together as a ‘micropyle–water-gap complex’.     

Morphology and anatomy of physical dormancy in Ipomoea lacunosa: identification of the water gap in seeds of Convolvulaceae (Solanales)

BACKGROUND AND AIMS: Convolvulaceae is the most advanced plant family (asterid clade) that produces seeds with physical dormancy (water-impermeable seed coat). There are several different opinions about the nature of the specialized structure ('water gap') in the seed coat through which water initially enters seeds of Convolvulaceae, but none of them has been documented clearly. The primary aim of the study was to identify the water gap in seeds of Ipomoea lacunosa (Convolvulaceae) and to describe its morphology, anatomy and function. METHODS: Light microscopy, scanning electron microscopy, tissue-sectioning, dye-tracking and blocking experiments were used to describe the morphology, anatomy and function of the water gap in seeds of I. lacunosa. KEY RESULTS: Dormancy-breaking treatments caused slits to form around the two bulges on the seed coat adjacent to the hilum, and dye entered the seed only via the disrupted bulges. Bulge anatomy differs from that of the rest of the seed coat. Sclereid cells of the bulges are more compacted and elongated than those in the hilum pad and in the rest of the seed coat away from the bulges. CONCLUSIONS: The transition area between elongated and square-shaped sclereid cells is the place where the water gap opens. Morphology/anatomy of the water gap in Convolvulaceae differs from that of taxa in the other 11 angiosperm plant families that produce seeds with physical dormancy for which it has been described.     

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.     

Identification and characterization of the water gap in the physically dormant seeds of Dodonaea petiolaris: a first report for Sapindaceae

Background and Aims

The Sapindaceae is one of 17 plant families in which seed dormancy is caused by a water-impermeable seed or fruit coat (physical dormancy, PY). However, until now the water gap in Sapindaceae had not been identified. The primary aim of this study was to identify the water gap in Dodonaea petiolaris (Sapindaceae) seeds and to describe its basic morphology and anatomy.

Methods

Seed fill, viability, water-uptake (imbibition) and other characteristics were assessed for D. petiolaris seeds. The location and structure of the water gap were investigated using a blocking experiment, time series photography, scanning electron microscopy and light microscopy. Dodonaea petiolaris seeds with PY also were assessed for loss of PY at four ecologically significant temperatures under moist and dry conditions. Seeds of three other species of Sapindaceae were examined for presence of a water gap.

Key Results

The water gap in D. petiolaris seeds was identified as a small plug in the seed coat adjacent to the hilum and opposite the area where the radicle emerges. The plug was dislodged (i.e. water gap opened = dormancy break) by dipping seeds in boiling water for 2·5 min or by incubating seeds on a moist substrate at 20/35 °C for 24 weeks. Layers of cells in the plug, including palisade and subpalisade, are similar to those in the rest of the seed coat. The same kind of water gap was found in three other species of Sapindaceae, Diplopeltis huegelii, Distichostemon hispidulus and Dodonaea aptera.

Conclusions

Following dormancy break (opening of water gap), initial uptake of water by the seed occurs only through the water gap. Thus, the plug must be dislodged before the otherwise intact seed can germinate. The anatomy of the plug is similar to water gaps in some of the other plant families with PY.     

9种形态生理休眠的种子脱水对萌发和胚胎生长的影响

9种形态生理休眠的种子脱水对萌发和胚胎生长的影响在具有形态生理种子休眠(MPD)的物种中,吸胀种子脱水对胚胎生长和萌发的影响鲜为人知。我们研究了9种不同MPD水平的种子对脱水的反应。对每个物种进行对照实验,使种子永久保持水化并暴露在最佳层积-培养顺序中以促进胚胎生长。同时也开展了室温条件下脱水中断种子层积处理1个月的实验。研究结果显示,具有非深度简单MPD的白藤铁线莲(Clematis vitalba)和高山茶藨子(Ribes alpinum)的胚生长 和种子活力均不受干燥影响,但干燥使高山茶藨子的萌发力下降了16%。具有深度简单上胚轴MPD的黄 水仙(Narcissus pseudonarcissus)种子在不同的胚生长阶段呈现脱水耐受性,但其萌发力略有下降。具有不同 MPD复杂水平的物种对脱水的反应更为多变：具有中度复杂MPD的Delphinium fissum亚种与具有深度复杂MPD的峨参(Anthriscus sylvestris)和熊根芹(Meum athamanticum),具有脱水耐受性。与之相反,具有非深度复杂MPD的鹅莓(Ribes uva-crispa)、中度复杂MPD的Lonicera pyrenaica和深度复杂MPD的Chaerophyllum aureum,脱水后萌发力下降。虽然具有MPD简单水平的种子能够具备脱水耐受性,但一些具有复杂水平MPD的种子也具有很高的耐受性。因此,脱水不诱导胚生长后期的次生休眠。9种植物中大多数的吸胀种子的脱水耐受性可能表征其对地中海地区气候变化的适应性。     

Identification and characterization of the water gap in physically dormant seeds of Geraniaceae, with special reference to Geranium carolinianum

Background and Aims

Physical dormancy in seeds of species of Geraniaceae is caused by a water-impermeable palisade layer in the outer integument of the seed coat and a closed chalaza. The chalazal cleft has been reported to be the water gap (i.e. location of initial water entry) in innately permeable seeds of Geraniaceae. The primary aim of this study was to re-evaluate the location of the water gap and to characterize its morphology and anatomy in physically dormant seeds of Geraniaceae, with particular reference to G. carolinianum.

Methods

Length, width, mass, anatomy and germination of two seed types (light brown and dark brown) of G. carolinianum were compared. Location, anatomy and morphology of the water gap were characterized using free-hand and microtome tissue sectioning, light microscopy, scanning electron microscopy, dye tracking, blocking and seed-burial experiments.

Key Results

Treatment with dry heat caused a colour change in the palisade cells adjacent to the micropyle. When placed in water, the ‘hinged valve’ (blister) erupted at the site of the colour change, exposing the water gap. The morphology and anatomy in the water-gap region differs from those of the rest of the seed coat. The morphology of the seed coat of the water-gap region is similar in G. carolinianum, G. columbinum, G. molle and G. pusillum and differs from that of the closely related species Erodium cicutarium.

Conclusions

Dislodgment of swollen ‘hinged valve’ palisade cells adjacent to the micropyle caused the water gap to open in physically dormant seeds of G. carolinianum, and it was clear that initial water uptake takes place through this gap and not via the chalazal opening as previously reported. This water gap (‘hinged valve gap’) differs from water gaps previously described for other families in morphology, anatomy and location in the seed coat.     

不同居群准噶尔无叶豆果实和种子特性及种子萌发差异

研究准噶尔无叶豆(Eremosparton songoricum) 6个居群间果实和种子特性及种子萌发差异, 以揭示异质生境下准噶尔无叶豆果实和种子的生态适应机制。结果显示: 居群间准噶尔无叶豆的植株距离(F = 2.34, p = 0.03)和植株冠幅(F = 8.49, p < 0.01)存在显著差异, 沙漠北缘E、F居群和沙漠腹地C居群(受人类干扰剧烈)的植株距离和植株冠幅高于沙漠腹地A、B、D居群; 居群间准噶尔无叶豆果实和种子的长度、宽度、厚度、重量存在显著差异, 居群E、F和C的大部分参数显著高于其他沙漠腹地居群; 居群间果实多子性(F = 6.96, p < 0.01)也存在显著差异, 居群C的果实多子性最高((32.50 ± 4.79)%); 萌发结果表明, 居群间新鲜的成熟种子萌发不存在显著差异, 且萌发率都低于15%; 所有居群的大部分种子都存在物理性休眠现象, 人为划破种皮能显著提高种子萌发率, 但在低温(15/5 ℃)条件下, 所有居群的种子萌发率都较低, 说明低温抑制了种子萌发; 经人为划破种皮解除物理休眠后, 种子的休眠没有完全释放, 居群C、E和F (大种子居群)的种子萌发率显著高于居群A、B和D (小种子居群) (F = 30.77, p < 0.01), 说明准噶尔无叶豆种子不仅存在物理性休眠现象, 也可能存在生理休眠现象。不同程度的种子复合休眠可能是准噶尔无叶豆不同居群适应古尔班通古特沙漠的重要生存策略。     

Seed development in Ipomoea lacunosa (Convolvulaceae), with particular reference to anatomy of the water gap

BACKGROUND AND AIMS: Disruption of one or both of the bulges (water gap) in the seed coat adjacent to the micropyle is responsible for breaking physical dormancy (PY) in seeds of Ipomoea lacunosa and other taxa of Convolvulaceae. Hitherto, neither ontogeny of these bulges nor onset of PY together with anatomical development and maturation drying of the seed had been studied in this family. The aims of this study were to monitor physiological and anatomical changes that occur during seed development in I. lacunosa, with particular reference to ontogeny of the water gap. METHODS: Developmental anatomy (ontogeny) of seed coat and dry mass, length, moisture content, germinability and onset of seed coat impermeability to water were monitored from pollination to seed maturity. Blocking/drying and dye-tracking experiments were done to identify site of moisture loss during the final stages of seed drying. KEY RESULTS: Physiological maturity of seeds occurred 22 d after pollination (DAP), and 100 % of seeds germinated 24 DAP. Impermeability of the seed coat developed 27-30 DAP, when seed moisture content was 13 %. The hilar fissure was identified as the site of moisture loss during the final stages of seed drying. The entire seed coat developed from the two outermost layers of the integument. A transition zone, i.e. a weak margin where seed coat ruptures during dormancy break, formed between the bulge and hilar ring and seed coat away from the bulge. Sclereid cells in the transition zone were square, whereas they were elongated under the bulge. CONCLUSIONS: Although the bulge and other areas of the seed coat have the same origin, these two cell layers underwent a different series of periclinal and anticlinal divisions during bulge development (beginning a few hours after pollination) than they did during development of the seed coat away from the bulge. Further, the boundary between the square sclereids in the transition zone and the elongated ones of the bulge delineate the edge of the water gap.     

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

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Morphophysiological dormancy in seeds of two North American and one Eurasian species of Sambucus (Caprifoliaceae) with underdeveloped spatulate embryos

In contrast to previous reports, the endocarps ("seed coats") of Sambucus species are not impermeable to water; thus, the seeds do not have physical dormancy. Seeds of the North American species Sambucus canadensis and S. pubens and of the European species S. racemosa have spatulate shaped embryos that are ～60% fully developed (elongated) at seed maturity. The embryo has to extend to the full length of the seed to germinate. Embryos in freshly matured seeds of S. canadensis and in those of S. pubens grew better at 25°/15°C than at 5°C, whereas the rate of embryo growth in S. racemosa was higher at 5°C than at 25°/15°C. Seeds of all three species germinated to significantly higher percentages in light (14-h photoperiod) than in darkness. Fresh seeds of neither species germinated during 2 wk of incubation over a range of thermoperiods. Warm followed by cold stratification broke dormancy in seeds of S. canadensis and in those of S. pubens. Thus, seeds of these two North American species have deep simple morphophysiological dormancy (MPD). In comparison, seeds of the European species S. racemosa required a cold stratification period only for dormancy break, and thus they have intermediate complex MPD. GA(3) was much more effective in breaking dormancy in seeds of S. racemosa than it was in those of S. canadensis or S. pubens.     

TRANSPARENT TESTA1 interacts with R2R3-MYB factors and affects early and late steps of flavonoid biosynthesis in the endothelium of Arabidopsis thaliana seeds

Wild type seed coats of Arabidopsis thaliana are brown due to the accumulation of proanthocyanidin pigments (PAs). The pigmentation requires activation of phenylpropanoid biosynthesis genes and mutations in some of these genes cause a yellow appearance of seeds, termed transparent testa (tt) phenotype. The TT1 gene encodes a WIP‐type zinc finger protein and is expressed in the seed coat endothelium where most of the PAs accumulate in wild type plants. In this study we show that TT1 is not only required for correct expression of PA‐specific genes in the seed coat, but also affects CHS, encoding the first enzyme of flavonoid biosynthesis. Many steps of this pathway are controlled by complexes of MYB and BHLH proteins with the WD40 factor TTG1. We demonstrate that TT1 can interact with the R2R3 MYB protein TT2 and that ectopic expression of TT2 can partially restore the lack in PA production in tt1. Reduced seed coat pigmentation was obtained using a TT1 variant lacking nuclear localisation signals. Based on our results we propose that the TT2/TT8/TTG1 regulon may also comprise early genes like CHS and discuss steps to further unravel the regulatory network controlling flavonoid accumulation in endothelium cells during A. thaliana seed development.