光调控植物种子萌发分子机制研究进展

萌发是种子植物进入农业生态系统的重要发育阶段。对于需光类种子,光是调控其萌发最重要的环境信号因子之一,红光促进而远红光抑制种子萌发。光敏色素是调控种子萌发的主要光受体。活化的光敏色素诱导萌发主效抑制因子PIF1发生蛋白降解,调节赤霉素和脱落酸代谢和信号途径相关基因的表达,从而促进种子的萌发。同时,一系列的表观遗传因子通过改变染色质结构,动态调节萌发相关基因的表达从而影响种子的萌发进程。该论文重点论述了光调控种子萌发的转录及表观遗传机制研究进展,并对其在农业生产中的应用进行了展望。     

隐花色素

光形态建成是光辐射影响下的植物生长发育过程,首先被人们观察到的是光对植物形态结构方面的影响。光照使黄化植物转绿而茁壮生长、影响种子萌发、并且调节植物的开花进程。这三类重要的现象推动这个领域的研究深入发展,通过红光和远红光可逆生理作用的认识导致了植物光敏色素的发现。使光敏色素活     

光照和温度对滇丁香种子萌发的影响

研究了不同光照和温度条件对滇丁香(Luculia pinciana)种子萌发的影响。结果表明,滇丁香种子是需光种子,有明显的光休眠现象;种子在光下萌发的最适温度范围为20~25℃,8~10d开始萌发,2~3周萌发完全,萌发率可达97%,温度的升高或降低均会降低种子萌发率。在15~30℃,用250mg/L GA₃处理24h能代替光照解除光休眠。     

光质、光强对入侵植物紫茎泽兰种子萌发及幼苗状态的影响

为了探讨紫茎泽兰种子需光萌发特性,对光质（不同颜色、红光/远红光）和光强对种子萌发率及幼苗状态进行了研究。研究发现：不同颜色光对发芽率影响显著,其中黄光、橙光和红光等波长在（591~750 nm）之间的光更有利于提高种子的萌发率（74.3%~83.3%）,而较短波长的光（<570 nm,紫光、蓝光和绿光）促进效果显著降低（62.3%~66.7%）（p<0.05）。光强对紫茎泽兰种子发芽和幼苗影响较光的颜色更具有规律性。黑暗条件下发芽率22%,随着光照提高,发芽率指数增加(r²=0.96),芽长指数下降(r²=0.99)、根长指数增加(r²=0.98),而鲜重呈现线性增加(r²=0.70)。适量红光（630 nm）和远红光（730 nm）照射能够打破和引起休眠,红光照射量与发芽率提高量成线性正相关（r²=0.98）,而远红光照射率与发芽率降低量呈线性正相关（r²=0.92）,说明紫茎泽兰需光萌发是一个光敏色素引起的过程。紫茎泽兰通常在裸地和人为干扰土壤上泛滥,这可能与其种子需光萌发有关。而对其生态学控制可以考虑从光强、特别是光质角度（如人工造林）控制种子萌发来实现。     

拟南芥突变体种子休眠与萌发的研究进展

种子的休眠和萌发是一个复杂的过程,至今尚未能清楚阐明其调控机制。目前已从拟南芥突变体中鉴定了一些与种子萌发和休眠相关的基因,有助于阐明种子休眠和萌发的分子机制。本文综述了拟南芥突变体种子休眠与萌发方面的研究进展。赤霉素是促进种子萌发的主要因素之一,RGL、SPY、GCR、SLY和GAR等基因的表达参与赤霉素对种子萌发的调控。脱落酸与种子休眠有关,ABI1、ABI2、ABI3、ABI4、ABI5、FUS3、LEC、MARD和CIPK等基因参与了脱落酸的调控过程。对3类乙烯反应的突变体（ein、etr和ctr）以及油菜素内酯突变体（det和bri）的研究表明乙烯和油菜素内酯是通过拮抗脱落酸而促进种子萌发的。光对种子萌发的调节,是通过具有Ser／Thr蛋白激酶活性的光敏色素PhyA、PhyB、PhyC、PhyD和PhyE,以磷酸化／去磷酸化方式调节其它与萌发相关基因的表达。含氮化合物对种子萌发的促进,可能是以一种依赖一氧化氮的方式解除种子休眠。     

光照和温度对望天树种子萌发的影响

在实验室内人工气候箱控制的条件下, 研究了我国重要珍稀濒危植物望天树 (Shorea wantianshuea,龙脑香科 (Dipterocarpaceae)) 种子萌发对持续光照、14小时光照 / 10小时黑暗周期性光照的反应; 同时研究了望天树种子的萌发对不同温度的反应和低温贮藏对种子活力的影响。结果表明, 不论是持续光照还是周期性光照都不能提高望天树种子的萌发率, 相反, 持续光照和周期性光照都不同程度降低了种子萌发率。光照能通过加速或延迟种子萌发的进程、或改变幼苗活力指数和萌发指数而影响种子萌发的质量; 持续光照延迟种子萌发的进程, 而周期性光照加快种子萌发的进程。30℃是种子的最适萌发温度, 虽然15℃和5℃的相对低温对幼苗活力指数影响不大, 但大大延迟了种子萌发进程, 并提高萌发率。望天树种子不能耐受5℃低温贮藏, 但具有在15℃下短期贮藏的潜力和一定程度的生理性休眠。     

光照和温度对望天树种子萌发的影响

在实验室内人工气候箱控制的条件下,研究了我国重要珍稀濒危植物望天树（Shorea wantianshuea,龙脑香科（Dipterocarpaceae））种子萌发对持续光照、14小时光照／10小时黑暗周期性光照的反应;同时研究了望天树种子的萌发对不同温度的反应和低温贮藏对种子活力的影响。结果表明,不论是持续光照还是周期性光照都不能提高望天树种子的萌发率,相反,持续光照和周期性光照都不同程度降低了种子萌发率。光照能通过加速或延迟种子萌发的进程、或改变幼苗活力指数和萌发指数而影响种子萌发的质量;持续光照延迟种子萌发的进程,而周期性光照加快种子萌发的进程。30℃是种子的最适萌发温度,虽然15℃和5℃的相对低温对幼苗活力指数影响不大,但大大延迟了种子萌发进程,并提高萌发率。望天树种子不能耐受5℃低温贮藏,但具有在15℃下短期贮藏的潜力和一定程度的生理性休眠。     

拟南芥突变体种子休眠与萌发的研究进展

种子的休眠和萌发是一个复杂的过程, 至今尚未能清楚阐明其调控机制。目前已从拟南芥突变体中鉴定了一些与种子萌发和休眠相关的基因, 有助于阐明种子休眠和萌发的分子机制。本文综述了拟南芥突变体种子休眠与萌发方面的研究进展。赤霉素是促进种子萌发的主要因素之一, RGL、SPY、GCR、SLY和GAR等基因的表达参与赤霉素对种子萌发的调控。脱落酸与种子休眠有关, ABI1、ABI2、ABI3、ABI4、ABI5、FUS3、LEC、MARD和CIPK等基因参与了脱落酸的调控过程。对3类乙烯反应的突变体 (ein、etr和ctr) 以及油菜素内酯突变体 (det和bri) 的研究表明乙烯和油菜素内酯是通过拮抗脱落酸而促进种子萌发的。光对种子萌发的调节, 是通过具有Ser/Thr蛋白激酶活性的光敏色素PhyA、PhyB、 PhyC、PhyD和PhyE, 以磷酸化/去磷酸化方式调节其它与萌发相关基因的表达。含氮化合物对种子萌发的促进, 可能是以一种依赖一氧化氮的方式解除种子休眠。     

光信号与激素调控种子休眠和萌发研究进展

休眠是种子植物在长期进化过程中产生的适应性性状, 通过抑制种子在不适宜的环境中萌发进而保证植物能够在逆境中生存。此外, 休眠有助于种子的长距离运输和扩散, 因此休眠对种子延续和物种保存具有重要意义。种子由休眠向萌发的发育转变不仅关系到物种的繁衍, 而且对保证农业生产中作物的产量和品质也具有重要作用。种子的休眠和萌发受到内源激素和外源光信号的共同调控。其中, 外源光信号主要通过调控内源ABA和GA的生物合成及信号转导进而调控种子休眠和萌发。该文系统综述了外源光信号和内源激素调控种子休眠和萌发的作用通路以及两类信号通路之间的交互作用, 旨在为农业生产中利用光和激素调控种子休眠与萌发提供参考。     

光信号与激素调控种子休眠和萌发研究进展

休眠是种子植物在长期进化过程中产生的适应性性状, 通过抑制种子在不适宜的环境中萌发进而保证植物能够在逆境中生存。此外, 休眠有助于种子的长距离运输和扩散, 因此休眠对种子延续和物种保存具有重要意义。种子由休眠向萌发的发育转变不仅关系到物种的繁衍, 而且对保证农业生产中作物的产量和品质也具有重要作用。种子的休眠和萌发受到内源激素和外源光信号的共同调控。其中, 外源光信号主要通过调控内源ABA和GA的生物合成及信号转导进而调控种子休眠和萌发。该文系统综述了外源光信号和内源激素调控种子休眠和萌发的作用通路以及两类信号通路之间的交互作用, 旨在为农业生产中利用光和激素调控种子休眠与萌发提供参考。     

Light activates the degradation of PIL5 protein to promote seed germination through gibberellin in Arabidopsis

Angiosperm seeds integrate various environmental signals, such as water availability and light conditions, to make a proper decision to germinate. Once the optimal conditions are sensed, gibberellin (GA) is synthesized, triggering germination. Among environmental signals, light conditions are perceived by phytochromes. However, it is not well understood how phytochromes regulate GA biosynthesis. Here we investigated whether phytochromes regulate GA biosynthesis through PIL5, a phytochrome-interacting bHLH protein, in Arabidopsis. We found that pil5 seed germination was inhibited by paclobutrazol, the ga1 mutation was epistatic to the pil5 mutation, and the inhibitory effect of PIL5 overexpression on seed germination could be rescued by exogenous GA, collectively indicating that PIL5 regulates seed germination negatively through GA. Expression analysis revealed that PIL5 repressed the expression of GA biosynthetic genes (GA3ox1 and GA3ox2), and activated the expression of a GA catabolic gene (GA2ox) in both PHYA- and PHYB-dependent germination assays. Consistent with these gene-expression patterns, the amount of bioactive GA was higher in the pil5 mutant and lower in the PIL5 overexpression line. Lastly, we showed that red and far-red light signals trigger PIL5 protein degradation through the 26S proteasome, thus releasing the inhibition of bioactive GA biosynthesis by PIL5. Taken together, our data indicate that phytochromes promote seed germination by degrading PIL5, which leads to increased GA biosynthesis and decreased GA degradation.     

Accumulation of Storage Materials,Precocious Germination and Development of Desiccation Tolerance During Seed Maturation in Mustard (Sinapis alba L.)

Under defined environmental conditions (20°C, continuous light of 15 klx) development of mustard seeds from artificial pollination to maturity takes about 60 d. After surpassing the period of embryo cell division and histodifferentiation (12–14d after pollination = dap), the seed enters into a maturation period. The time courses of various physiological, biochemical, and structural changes of embryo and testa during seed maturation were analyzed in detail (dry and fresh mass changes, osmotic and water potential changes, respiration, DNA amplification by endomitosis, total ribosome and polysome formation, storage protein synthesis and accumulation, storage lipid accumulation). In addition to the final storage products protein and lipid, embryo and testa accumulate transiently large amounts of starch within the chloroplasts during early maturation. Concomitantly with the subsequent total breakdown of the starch, the plastids lose most of their internal structure and chlorophyll and shrink into proplastids, typical for the mature seed. At about 30 dap the seeds shift from a desiccation-sensitive to a desiccation-tolerant state and are able then to germinate rapidly upon drying and reimbibition. If isolated from the immature fruit and sown directly on water, the seeds demonstrate precocious germination from about 13 dap onwards. Young seeds (isolated ≦ 38 dap) germinate only after surpassing a lag-phase of several days (after-ripening) during which the embryo continues to accumulate storage protein and lipid at the expense of the surrounding seed tissues. We conclude from these results that the maturing seed represents a rather closed developmental system which is able to continue its development up to successful germination without any specific regulatory influence from the mother plant. Immature seeds are able to germinate without a preceding dehydration treatment, which means that partial or full desiccation does not serve as an environmental signal for reprogramming seed development from maturation to germination. Instead, it is argued that the water relations of the seed are a critical element in the control of maturation and germination: during maturation on the mother plant the embryo is subject to a considerable turgor pressure (of the order of 12 bar) accompanied by a low water potential (of the order of ?12 bar). This turgor permits maturation growth but is subcritical for germination growth. However, upon imbibition in water, the low water potential provides a driving force for a burst of water uptake overcoming the critical turgor threshold and thereby inducing germination.     

盐生植物角果碱蓬种子二型性对环境的适应策略

角果碱蓬(Suaeda corniculata)是藜科一年生盐生植物, 在我国分布于北方盐碱滩涂和盐碱荒漠地区。角果碱蓬具有棕色和黑色两种异型体种子(简称棕色和黑色种子)。对采自内蒙古鄂托克前旗盐渍化生境的角果碱蓬二型种子的形态、休眠和萌发特性开展对比研究, 测定了二型种子休眠和萌发行为对温度、光照和盐分(NaCl)的响应, 以揭示盐生植物异型种子对温带盐漠生境的适应对策。结果表明: (1)二型性种子在大小、种皮特性和结实比例方面有显著差异。与黑色种子相比, 棕色种子个体较大, 种皮透水性强。黑色种子与棕色种子的结实比例约为5.6 : 1。(2)新成熟的棕色种子的萌发对各温度梯度和光照条件不敏感, 萌发率较高(84%-100%); 而新成熟的黑色种子萌发率较低(8%-78%), 萌发对光照敏感。(3)黑色种子具有浅度生理休眠, 种皮划破、赤霉素处理和低温层积均可有效地提高种子的萌发率。(4)二型种子萌发对土壤盐分的胁迫具有不同的响应。与黑色种子相比, 棕色种子对盐分胁迫不敏感, 在较高的盐分浓度下仍有较高的萌发率, 低温层积处理能够降低黑色种子对盐胁迫的敏感性, 有效地提高种子的初始萌发率、萌发恢复率和最终萌发率。角果碱蓬二型种子不同的形态、休眠和萌发特性, 提高了该物种在高度异质性生境中的适合度, 对种群成功地适应温带盐漠环境具有重要的意义。     

Photoinhibition of radish (Raphanus sativus L) seed germination: control of growth potential by cell-wall yielding in the embryo

The biophysical mechanism underlying photoinhibition of radish (Raphanus sativus L.) seed germination was investigated using three cultivars differing in sensitivity to continuous irradiation with far-red light (high-irradiance reaction of phytochrome). Sensitivity of germination to the inhibitory action of light was assessed by probing germination under osmotic stress (incubation in media of low water potentials adjusted with polyethylene glycol 6000) and expressed in terms of ‘germination potential’ (positive value of the water potential at which germination is inhibited by 50%). Far-red light decreases the germination potential to various degrees in the different cultivars, reflecting the light-sensitivity of germination in water. Removal of the seed coat increases the germination potential by a constant amount in darkness and light. It is concluded that germination depends on the expansive force of the embryo which can be drastically diminished by far-red light. Seed-coat constraint and expansive force of the embryo interact additively on the level of the germination potential. Photoinhibition of germination was accompanied by an inhibition of water uptake into the seed. Analysis of seed water relations showed that osmotic pressure and turgor assumed higher levels in photoinhibited seeds, compared to seeds germinating in darkness, while the water potential was close to zero under both conditions. Far-red light produced a shift (to less negative values) in the curve relating water-uptake rate to external water potential, i.e. a reduction in the driving force for water uptake. It is concluded that photoinhibition of germination results from the maintenance of a high threshold of cell-wall extensibility in the embryo.     

Germination and water dispersal of seeds from a threatened plant species Penthorum chinense

To conserve a threatened plant species (Penthorum chinense Pursh) in Japan, seed germination responses to pretreatment (imbibition and/or chilled), temperature and light, and seed dispersal by water were examined. The seeds collected from abandoned paddy fields in a warm temperate region, central Japan, germinated in light (14 h photoperiod; light 22°C, dark 21°C) after a moist-chilled treatment. After this pretreatment, the seeds germinated well at 10–25°C (optimum temperature 15°C), but did not germinate in darkness even at the optimum temperature. Most of the seeds floated on distilled water, but 20–60% of the seeds that were collected from several populations sank in distilled water, indicating dimorphism in seed dispersal by water. The floating and sunken seeds did not show significant differences in weight and germination rate within a population. The addition of a surface-active agent in distilled water submerged the seeds, indicating that the buoyancy of the seeds is attributable to an oil coating on the seed surface that enhances the interfacial tension on the seeds. Three times the number of seeds sank in river water collected from a rural area than in distilled water. A greater number of seeds also sank in water that had increasing concentrations of linear alkylbenzenesulfonate, which is a major component of synthetic detergents. This suggests that the water dispersal of this species is suppressed by surface-active agents, including detergents, in river water.     

Phytochrome regulation and differential expression of gibberellin 3beta-hydroxylase genes in germinating Arabidopsis seeds.

Despite extensive studies on the roles of phytochrome in photostimulated seed germination, the mechanisms downstream of the photoreceptor that promote germination are largely unknown. Previous studies have indicated that light-induced germination of Arabidopsis seeds is mediated by the hormone gibberellin (GA). Using RNA gel blot analyses, we studied the regulation of two Arabidopsis genes, GA4 and GA4H (for GA4 homolog), both of which encode GA 3beta-hydroxylases that catalyze the final biosynthetic step to produce bioactive GAs. The newly isolated GA4H gene was expressed predominantly during seed germination. We show that expression of both GA4 and GA4H genes in imbibed seeds was induced within 1 hr after a brief red (R) light treatment. In the phytochrome B-deficient phyB-1 mutant, GA4H expression was not induced by R light, but GA4 expression still was, indicating that R light-induced GA4 and GA4H expression is mediated by different phytochromes. In contrast to the GA4 gene, the GA4H gene was not regulated by the feedback inhibition mechanism in germinating seeds. Our data demonstrate that expression of GA 3beta-hydroxylase genes is elevated by R light, which may result in an increase in biosynthesis of active GAs to promote seed germination. Furthermore, our results suggest that each GA 3beta-hydroxylase gene plays a unique physiological role during light-induced seed germination.     

Seed anatomy and water uptake in relation to seed dormancy in Opuntia tomentosa (Cactaceae, Opuntioideae)

BACKGROUND AND AIMS: There is considerable confusion in the literature concerning impermeability of seeds with 'hard' seed coats, because the ability to take up (imbibe) water has not been tested in most of them. Seeds of Opuntia tomentosa were reported recently to have a water-impermeable seed coat sensu lato (i.e. physical dormancy), in combination with physiological dormancy. However, physical dormancy is not known to occur in Cactaceae. Therefore, the aim of this study was to determine if seeds of O. tomentosa are water-permeable or water-impermeable, i.e. if they have physical dormancy. METHODS: The micromorphology of the seed coat and associated structures were characterized by SEM and light microscopy. Permeability of the seed-covering layers was assessed by an increase in mass of seeds on a wet substrate and by dye-tracking and uptake of tritiated water by intact versus scarified seeds. KEY RESULTS: A germination valve and a water channel are formed in the hilum-micropyle region during dehydration and ageing in seeds of O. tomentosa. The funicular envelope undoubtedly plays a role in germination of Opuntia seeds via restriction of water uptake and mechanical resistance to expansion of the embryo. However, seeds do not exhibit any of three features characteristic of those with physical dormancy. Thus, they do not have a water-impermeable layer(s) of palisade cells (macrosclereids) or a water gap sensu stricto and they imbibe water without the seed coat being disrupted. CONCLUSIONS: Although dormancy in seeds of this species can be broken by scarification, they have physiological dormancy only. Further, based on information in the literature, it is concluded that it is unlikely that any species of Opuntia has physical dormancy. This is the first integrative study of the anatomy, dynamics of water uptake and dormancy in seeds of Cactaceae subfamily Opuntioideae.     

喀什霸王的结实和种子萌发特性

喀什霸王(Zygophyllum kaschgaricum)是生长于中国新疆南部荒漠环境的稀有种及二级保护植物。当前, 该物种在自然种群中呈分散式及片段化分布, 且种群密度低, 种群老龄化较严重。因此, 为了了解该物种种子萌发特性及其对荒漠环境的响应, 该文采用室内控制实验方法, 对该物种的自然坐果率、结籽率、种子吸水特性、种子休眠和萌发特性及对干旱胁迫的响应进行了比较研究。结果表明: 喀什霸王在自然种群中的坐果率及结籽率较低, 且种子的败育率较高。不同干藏时间种子的吸水速率间存在显著差异; 随着干藏时间的延长, 种子的吸水率逐步增强。刚成熟的种子在不同温度及光周期下均可萌发; 其中高温(10/20 ℃, 20/30 ℃)及黑暗条件下的萌发率比低温(10/5 ℃, 5/2 ℃)及光照条件下的萌发率高。不同干藏时间的种子在不同浓度赤霉素(GA₃)下的萌发率均较高; 但低温储藏时间对该物种种子的打破休眠及萌发率没有促进作用。以上结果说明该物种存在非深度生理休眠; 而干藏时间、高温且黑暗及高浓度(50 mmoloL ^-1) GA₃是打破休眠及促进种子萌发的最合适条件。高温条件下的干旱胁迫对喀什霸王种子萌发具有抑制作用; 春季和秋季降水量决定种子的萌发率。总之, 喀什霸王种子在物候上表现出的春秋季萌动及非深度生理休眠以提高幼苗存活力及保障种群稳定性, 是一种对新疆南部干旱及高温胁迫荒漠环境的适应策略。