蔷薇种子的休眠及解除方法

分析了蔷薇(Rosa L.)种子休眠原因、解除休眠方法以及环境条件对休眠与萌发的影响.蔷薇种子休眠的主要原因有瘦果果皮和种皮的限制作用,胚生理休眠以及果肉、瘦果果皮、种皮和胚中的抑制物质.解除休眠的方法包括去除瘦果果皮限制、解除胚的生理休眠、去除抑制物质等.种子发育过程中及成熟后,环境因子,如温度、水分和光照,对种子休眠和萌发有影响.此外,微生物、果实采集时间也对种子休眠及萌发有较大影响.蔷薇种子的休眠机制复杂,且种间差异很大.     

濒危植物巴东木莲种子休眠与萌发特性的研究

巴东木莲(Manglietia patungensis)为我国特有种, 属国家重点保护植物。为找出其生殖环节中的致危因素, 作者对巴东木莲种子休眠与后熟过程中的形态和萌发特性进行了研究。结果表明, 巴东木莲种胚发育不完全可能是种子休眠的主要原因, 在其后熟过程中胚不断分化、发育成熟; 种皮具有较好的透性, 与休眠的关系不大; 种子不同部位均存在萌发抑制物, 胚乳中高含量的萌发抑制物是影响胚萌发的重要因素。内源激素ABA和IAA在巴东木莲种子休眠与萌发过程中起着重要作用, ABA是引起休眠的关键因素, IAA有助于种子的萌发, IAA/ABA相对含量的变化对种子的休眠和萌发产生重要影响。巴东木莲种子的休眠是由种子本身的形态和生理特点引起的综合休眠, 在4℃低温保湿条件下才能完成其形态和生理后熟过程, 而自然条件下, 巴东木莲种子成熟时正值秋季少雨, 很容易失水而不能完成其后熟过程而失去生活力, 这可能是导致该物种自然更新困难的重要原因。     

脱落酸调控种子休眠和萌发的分子机制

脱落酸(ABA)是调控种子休眠和萌发过程的主要植物激素。种子内源ABA含量和种胚对ABA敏感性共同调控种子休眠和萌发过程, 确保植物种子以休眠状态在逆境中保持其自身繁衍能力, 并在适宜的环境下启动萌发程序。种子ABA合成代谢和ABA信号转导途径涉及许多重要基因家族, 它们通过复杂的调控网络精确地控制着种胚发生、种子成熟、休眠及萌发进程。该文对ABA调控种子休眠和萌发的分子机制最新研究进展进行综述, 并展望了今后的研究方向。     

野生玫瑰果实发育过程中种子的细胞组织结构和内源激素变化对其休眠性的影响

以吉林珲春自然群落的野生玫瑰(Rosa rugosa Thunb.)为试验材料,探究野生玫瑰在果实形成过程中种子的发育及休眠性的形成和变化。选取青果期(1~35 d)、转色期(35~60 d)、红果期(60~75 d)的果实及种子,结合形态学、组织细胞学观察法及高效液相色谱技术,对果实各发育时期的种子形态、种胚及内果皮的发育进行研究,并分析种子内源激素含量变化与果实发育、种子休眠之间的关系。结果表明:果实和种子在青果期(1~35 d)时发育速度最快,种胚在花后24 d发育完全,种子不存在形态休眠。花后24 d内果皮开始沉积木质素并逐渐木质化,种子开始产生机械休眠。种子激素含量的变化与果实的发育、转色及内果皮的木质化密切相关,种子内源GA3和ABA含量在青果期(1~35 d)达到峰值,内源IAA含量在果实转色期(35~60 d)达到最大值,高浓度的ABA含量是种子尚未脱离果实时便已进入生理休眠的主要原因。     

GA3和变温层积对天女木兰种子萌发及内源激素的影响

用不同质量浓度GA3浸泡天女木兰种子并结合变温层积处理,应用高效液相色谱法对不同时期种子中4种激素GA3、IAA、ABA、ZR含量进行测定,并测量种胚长和萌发率,以探讨天女木兰种胚发育,内源激素含量变化与种子休眠萌发之间的调控关系,为进一步研究种子休眠机理提供理论基础。结果表明:(1)天女木兰成熟种子胚发育不完全,胚乳内高浓度ABA和低浓度GA3是其休眠的主要原因。(2)GA3处理能促使天女木兰种子提前30d完成形态后熟,并以1 500mg·L-1 GA3处理效果最佳。(3)在变温层积过程,天女木兰种胚发育可分三个阶段:阶段Ⅰ(0~70d)完成种胚进一步分化;阶段Ⅱ(70~120d)种胚快速生长时期;阶段Ⅲ(120~150d)休眠完全解除,种子具备发芽能力。种子能否打破休眠主要取决于阶段Ⅰ和Ⅱ的状况。(4)GA3/ABA、IAA/ABA和ZR/ABA在种子后熟期间的变化同胚生长发育存在一致性,认为内源激素的相对水平对种子休眠具有重要的调控作用。     

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。     

葡萄属种子发育的物候、萌发行为及其对冷层积的反应

以3个起源和分布中心的不同葡萄种及其杂交种作为实验材料,观察了开花、果实和种子成熟的物候,研究了成熟种子的基本特性和最初萌发率以及冷层积对成熟种子萌发的影响。结果表明,葡萄种子具有休眠特性,生理成熟时都具有分化完全的胚,在休眠解除过程中胚的形态不发生变化;美洲种休眠最深,欧亚种休眠最浅,其它种类的葡萄种子的休眠程度介于美洲种和欧亚种之间;不同种葡萄花期、果实和种子成熟过程的物候存在差异,果实成熟与种子成熟不同步,其间隔时间越短,种子休眠程度越深。冷层积60天能有效地解除东亚种、欧亚种、欧山杂种和蘡欧杂种的种子休眠,但对解除美洲种、美洲种种间杂种和欧美杂种的种子休眠效果较差。     

葡萄属种子发育的物候、萌发行为及其对冷层积的反应

以3个起源和分布中心的不同葡萄种及其杂交种作为实验材料, 观察了开花、果实和种子成熟的物候, 研究了成熟种子的基本特性和最初萌发率以及冷层积对成熟种子萌发的影响。结果表明, 葡萄种子具有休眠特性, 生理成熟时都具有分化完全的胚, 在休眠解除过程中胚的形态不发生变化; 美洲种休眠最深, 欧亚种休眠最浅, 其它种类的葡萄种子的休眠程度介于美洲种和欧亚种之间; 不同种葡萄花期、果实和种子成熟过程的物候存在差异, 果实成熟与种子成熟不同步, 其间隔时间越短, 种子休眠程度越深。冷层积60天能有效地解除东亚种、欧亚种、欧山杂种和蘡欧杂种的种子休眠, 但对解除美洲种、美洲种种间杂种和欧美杂种的种子休眠效果较差。     

珍稀濒危植物金丝李种子的休眠机理

珍稀濒危植物金丝李(Garcinia paucinervis)种子的萌发十分缓慢,探讨其休眠机理,可为该物种的种质资源保育与可持续利用提供理论依据。本文对金丝李种子种皮结构及其透水性,剔除部分种皮和胚乳后种子的萌发情况,胚乳和胚等粗提物的活性,储藏、层积和不同温度下种子萌发情况,萌发过程中内源激素含量等进行了研究。结果表明:金丝李种皮无栅栏细胞层,下表面的角质层较薄;种皮对种子的吸胀阻碍小;随着种孔端剔除种皮和胚乳程度的加深,金丝李种子的萌发进程逐渐延长,甚至降低其萌发率,种脐端削除处理对种子萌发影响不大;内果皮、种皮、胚乳和胚中可能存在抑制金丝李种子萌发和生长的内源抑制物;新鲜种子胚率达86.12%,低温层积后胚率无显著变化;低温层积处理延缓其萌发进程,对萌发率无显著影响,4℃低温层积是储藏金丝李种子的较好方法;种子萌发对温度敏感,在32℃培养下可打破种子休眠,萌发速度显著加快。种子萌发过程中ABA含量降低,GA与ABA、IAA与ABA的比值随种子萌发显著升高,萌发促进与抑制物比例逐渐趋于提高。因此,金丝李种子存在内源抑制物,同时缺乏萌发促进物质,导致生理休眠。该种子休眠特性使其幼苗生长能应对生境的季节变化,种群在风险环境中得以延续,避免大量幼苗竞争。植被破坏导致种子萌发阶段受阻是造成金丝李濒危的原因之一。     

小花草玉梅种子萌发过程中常见内源激素含量的变化及萌发机制

植物内源激素在调节种子休眠和萌发过程中具有极其重要的作用。本研究运用反式高效液相色谱(RP-HPLC)与紫外检测器联用的方法对小花草玉梅干种子、吸胀种子和露白种子中的赤霉素GAs、脱落酸ABA、玉米素ZT和生长素IAA含量进行了检测,旨在研究植物内源激素水平在种子萌发过程中的变化是否直接关系着小花草玉梅种子在光照或黑暗条件下的萌发能力。结果显示,种子吸胀过程中,光照促进了ZT同时抑制了ABA的积累,并且ZT对ABA萌发抑制作用的解除也受光的促进,露白种子中的ABA/ZT,ABA/GAs和(ABA+IAA)/(GAs+ZT)水平在黑暗条件下高于光照条件,上述均是导致光照条件下种子萌发率较高的重要原因;相对于干种子,IAA含量在种子吸胀初期急速下降,(ABA+IAA)/(GAs+ZT)在种子萌发过程中有所降低,而ABA/GAs却表现出明显的上升趋势;各激素水平所受光照的影响均在种子开始露白时显著减弱,另外,吸胀第9天是小花草玉梅种子萌发过程中激素变化的一个关键的时间节点。总之,种子萌发并非直接关系着GA含量的升高和ABA含量的降低。因光照直接促进了小花草玉梅种子的萌发,本研究认为高寒草甸充足的光照和较强的太阳辐射通过调节种子内源激素水平在一定程度上对种子萌发过程中的休眠解除具有重要作用。     

Dormancy and Germination of Abscisic Acid-Deficient Tomato Seeds : Studies with the sitiens Mutant

The role of abscisic acid (ABA) in the dormancy induction of tomato (Lycopersicon esculentum) seeds was studied by comparison of the germination behavior of the ABA-deficient sitiens mutant with that of the isogenic wild-type genotype. Freshly harvested mutant seeds, in contrast to wild-type seeds, always readily germinate and even exhibit viviparous germination in overripe fruits. Crosses between mutant and wild-type and self-pollination of heterozygous plants show that in particular the ABA fraction of embryo and endosperm is decisive for the induction of dormancy. After-ripened wild-type seeds fully germinate in water but are more sensitive toward osmotic inhibition than mutant seeds. Germination of both wild-type and mutant seeds is equally sensitive toward inhibition by exogenous ABA. ABA content of mature wild-type seeds is about 10-fold the level found in mutant seeds. Nevertheless, it is argued that the differences in dormancy between the seeds of both genotypes are not a result of actual ABA levels in the mature seeds or fruits but a result of differences in ABA levels during seed development. It is hypothesized that the high levels of ABA that occur during seed development in wild-type seeds induce an inhibition of cell elongation of the radicle that can still be observed after long periods of dry storage.     

Diversities in morphological characteristics and seed germination behavior in fruits ofSalsola komarovii Iljin

Two different types of dispersal units (called fruits in this study) were observed inSalsola komarovii Iljin. One is a fruit which has dark brown lignified tepals with long wings and a green seed and falls easily from the mother plant (long-winged type). Another has light brown lignified tepals with short wings and a yellow seed and attaches tightly to the mother plant (short-winged type). This difference of fruit type appeared independently from maturity of fruit. Seeds in the short-winged fruits were in dormancy for a longer period of time than those in the long-winged fruits. The germination rate was significantly higher in the seeds of long-winged fruits. The dormancy in seeds of the long-winged fruits was effectively terminated by reducing the temperature but the effect of chilling was very weak in seeds of the short-winged fruits. It was concluded from these observations that there exists a dimorphism in the fruits of, or a heterocarpy in,S. komarovii. The plants grown under water stress produced mostly short-winged fruits and those grown under well-watered conditions bore fruits of both types. Exogenously applied abscisic acid (ABA) tended to produce the short-winged fruits, suggesting that the heterocarpy was, at least partly, regulated by ABA.     

Seed Dormancy in Acer: The Role of Testa-imposed and Embryo Dormancy in Acer velutinum

The embryo dormancy shown in freshly harvested samples of Acervelutinum seeds is weakly established and very short-lived.Loss of this embryo dormancy occurred during post-harvest fruitstorage at either 5 or 17 C. In contrast, the dormancy of intactfruits and seeds was overcome only during storage at the lowertemperature. Removal of the cotyledons from embryos of freshlyharvested fruits allowed more rapid germination of the embryonicaxes, indicating that the cotyledons exert an inhibitory effect,although the axes still retained a measure of innate dormancy.The inhibitory effect of the cotyledons became less marked withincreasing duration of fruit storage, this loss of inhibitoryeffect occurring at both storage temperatures. Applied ABA stronglysuppressed germinative capacity in intact embryos and isolatedembryonic axes from freshly harvested fruits, but when ABA wasapplied to embryos of fruits that had been stored for variousperiods at 5 or 17 C, the inhibitory effect was first weakenedand then lost with increased storage. Although dormancy in the seeds of A. velutinum may be describedas intermediate between testa-imposed dormancy and true dormancy,it is perhaps more properly included in the former category. Acer velutinum Boiss. var. vanvolxemii, abscisic acid, embryo dormancy, germination, seed storage, testa-imposed dormancy, tissue sensitivity     

Induction of dormancy during seed development by endogenous abscisic acid: studies on abscisic acid deficient genotypes of Arabidopsis thaliana (L.) Heynh.

Mutant lines of Arabidopsis thaliana (L.) Heynh., which are characterized by symptoms of withering and the absence of seed dormancy, showed much lower levels of endogenous abscisic acid (ABA) in developing seeds and fruits (siliquae) than the wild type. Reciprocal crosses of wild type and ABA-deficient mutants showed a dual origin of ABA in developing seeds. The genotype of the mother plant regulated a sharp rise in ABA content half-way seed development (maternal ABA). The genotype of the embryo and endosperm was responsible for a second ABA fraction (embryonic ABA), which reached much lower levels, but persisted for some time after the maximum in maternal ABA. The onset of dormancy correlated well with the presence of the embryonic ABA fraction and not with the maternal ABA. Dormancy developed in both the absence and presence of maternal ABA in the seeds. In this respect maternal ABA resembled exogenously applied ABA which did not induce dormancy in ABA-deficient seeds. However, both maternal and applied ABA stimulated the formation of a mucilage layer around the testa, which could be observed during imbibition of the mature seeds. In the mature state, ABA-deficient seeds germinated in the siliquae on the plant, but only when the atmosphere surrounding the plant was kept at high relative humidity. In younger stages germination in siliquae occurred after isolation from the plants and incubation on wet filter paper. Therefore, it seems that limited access to water is the primary trigger for the developmental arrest in these seeds.     

Seed dormancy in Acer: Endogenous germination inhibitors and dormancy in Acer pseudoplatanus L.

Summary Dormant seeds of Acer pseudoplatanus L. contain two zones of inhibition on paper chromatograms in 10:1:1 as detected by the lettuce and cress seed germination, and the wheat coleoptile bioassays. One zone at R_f 0.6–0.8 was partitioned into ethyl acetate at acid pH and was shown to contain ABA by its behaviour on GLC and isomerization under ultra-violet light. The other zone at R_f 0.9 was detected only in the germination bioassays and was partitioned into ethyl acetate over a range of pH indicating the presence of one or more neutral compounds.The inhibitors present in the embryo of dormant sycamore seeds inhibited the germination of non-dormant sycamore seeds at relatively low concentrations. A comparison with the effects of application of exogenous ABA indicated that endogenous ABA could not solely account for the inhibitory activity of seed extracts, which appeared to be due partly to the presence of ABA and partly to that of neutral compounds present in the embryo. Leaching treatments that removed dormancy led to a decrease in the level of inhibitors present mainly in the basic fraction. The exogenous application of kinetin to dormant sycamore seeds increased germination whereas gibberellic acid had no effect. Similar responses were obtained with lettuce seeds inhibited by the basic fraction of dormant sycamore seeds.It is suggested that an inhibitor-cytokinin interaction may be involved in the dormancy of sycamore seeds.     

珍稀濒危植物珙桐种子休眠萌发过程中内源激素的变化

珙桐是我国特有珍稀濒危植物,休眠期长且具二次休眠现象。将处于休眠萌发过程中的珙桐种子依据胚根长度划分为4个阶段,利用高效液相色谱(HPLC)测定各阶段种子及其内果皮中ABA(脱落酸)、GA(赤霉素)、KT(细胞分裂素)、IAA(3-吲哚乙酸)4种内源激素含量,分析其比值动态变化,并与幼苗阶段进行比较。结果显示:未破壳种子的内果皮中内源激素含量以ABA最高,其次是GA、IAA、KT,随着种子破壳后四种激素含量显著降低。除ABA外,种子中GA、IAA和KT含量随着胚根的伸长逐渐升高,但仍低于幼苗阶段。此外,随着胚根伸长,种子中GA/ABA、IAA/ABA、KT/ABA比值逐渐增大,其中以GA/ABA的变化最显著。因此,珙桐种子的休眠和萌发可能主要受ABA和GA的平衡和拮抗来调控。     

The Role of Water Uptake in the Transition of Recalcitrant Seeds from Dormancy to Germination

In recalcitrant seeds of horse chestnut (Aesculus hippocastanum L.) maintaining a high water content during winter, dormancy is determined by the presence and influence of the seed coat, while the axial organs of the embryos excised from these seeds are not dormant. Such axial organs were capable for active water uptake and rapid fresh weight increase, so that their fresh weights exceeded those in intact seeds at the time of radicle protrusion. Fructose plays an essential role in the water uptake as a major osmotically active compound. ABA interferes with the water uptake by the axial organs and thus delays the commencement of their growth. The manifestation of seed response to ABA during the entire dormancy period indicates the presence of active ABA receptors and the pathways of its signal transduction. The content of endogenous ABA in the embryo axes doubled in the middle of dormancy period, which coincided with a partial suppression of water uptake by the axes. During seed dormancy release and imbibition before radicle protrusion, the level of endogenous ABA in axes declined gradually. Application of exogenous ABA can imitate dormancy by limiting water absorption by axial organs. Fusicoccin A (FC A) treatment neutralized completely this ABA effect. Endogenous FC-like ligands were detected in the seed axial organs during dormancy release and germination. Apparently, endogenous FC stimulates water uptake via the activation of plasmalemmal H⁺-ATPase, acidification of cell walls, their loosening, and turgor pressure reduction. FC can evidently counteract the ABA-induced suppression of water uptake by controlling the activity of H⁺-ATPase. It is likely that, in dormant intact recalcitrant seeds, axial organs, maintaining a high water content, are competent to elevate their water content and to start their preparation for germination under the influence of FC when coat-imposed dormancy becomes weaker.