异子蓬二型种子的萌发与休眠特性及其生态适应

异子蓬为中亚荒漠的特有种,其果实为胞果,花被片在结果时宿存包被果实形成浆果状的散布单位.对异子蓬的散布单位、果实形态进行比较观察,并在5/15℃、5/25℃、15/25 ℃(暗/光=12h/12h)3个温变周期下进行种子萌发实验,结果表明:(1)异子蓬的散布单位绿色、浆果状,有大小两种形态;大小两种散布单位内包被的果实在形状、大小、颜色上存在着明显差异.(2)大的散布单位内包被的果实圆形,扁平,褐色,无光泽,直径(2.64±0.03) mm,重量(3.68±0.04) mg;小的散布单位内包被的果实双凸镜形,黑色,有光泽,直径(2.40±0.04) mm,重量(2.86±0.06) mg.(3)褐色种子在3个温变周期下的萌发率都在81%以上,萌发速度快,而黑色种子萌发率低,萌发慢,具有休眠现象.(4)划破种皮及低温层积处理可显著提高黑色种子在3个温变周期下的萌发率,延长储存时间也可有效地促进黑色种子的萌发,表明黑色种子处于非深度生理休眠状态.异子蓬产生的两种不同类型的种子及其在时间和空间上的萌发差异对荒漠异质环境具有重要的适应意义.     

盐分、变温和激素处理对盐生植物异子蓬异型性种子萌发及成苗的影响

利用培养皿种子萌发及土壤盆栽方法对新疆特有藜科1年生盐生植物异子蓬(Suaeda aralocaspica)在不同盐分(NaCl和Na2SO4)、变温(4～30℃)和激素(GA3)处理下的种子萌发及成苗行为进行了研究.结果表明:异子蓬2种种子在NaCl和Na2SO42种盐胁迫下的萌发和休眠特性表现出明显差异,其中黑色种子的萌发率比褐色种子低;黑色种子属于非深度生理休眠,可分别通过15/30℃变温处理、4℃低温层积30 d及用800mg·L-1赤霉素处理促进其萌发;盐处理及温度变化对褐色种子的出苗率和成苗率影响显著,对黑色种子则不明显,说明黑色种子萌发存在着持续性的缓萌机制,有利于其形成持久种子库,以上特性均显示出其对初春盐生环境的适应性.     

外源激素ABA影响新疆荒漠盐生植物异子蓬异型种子萌发机制

以新疆荒漠盐生植物异子蓬(Suaeda aralocaspica)异型种子为材料,通过外源ABA直播(长时间)与浸种(短时间)处理分别研究不同浓度(0、0.5、1.0、5.0、10.0μmol/L)ABA对异型种子萌发、幼苗生长的影响,并通过定量PCR技术分析与萌发相关的ABC转运蛋白基因、丝裂原活化蛋白激酶激酶基因和DNA修复和转录因子基因在5.0μmol/L ABA处理下的表达规律,探讨外源ABA影响异子蓬异型种子萌发差异的机制。结果显示:(1)外源ABA直播处理下,褐色种子萌发及幼苗生长均受到明显抑制,黑色种子的萌发和幼苗生长在低浓度被促进,较高浓度被显著抑制;短时间浸种处理,能够同时促进异型种子的萌发及幼苗生长,且随ABA浓度增加促进效应增强。(2)直播萌发后,褐色种子中3个基因的表达量与对照相比均不变或降低表达(除了DNA修复和转录因子基因在8 h显著升高),黑色种子与对照相比均上调表达(除了ABC转运蛋白基因在8h显著下调);短时间浸种萌发后,褐色种子中3个基因的表达量均比对照显著升高,黑色种子中升高不明显。基因表达规律与种子萌发结果趋势一致,暗示ABA浸种可能触发了异子蓬种子萌发内在机制并对随后的萌发过程产生促进作用。而黑色种子对ABA处理表现出较好的萌发响应,可能是其幼苗能抵御荒漠地区逆境环境的重要原因。     

种子异型植物异子蓬的生殖分配和结实格局

选择能产生异型果实和种子的一年生盐生植物异子蓬(Suaeda aralocaspica)为研究对象, 研究其生殖分配和结实格局。结果表明, 异子蓬具有较高的生殖分配和规律性的“谨慎型”结实格局。异子蓬的生殖分配高达56%。在资源充裕时, 该植物对扁圆形棕色种子(采取机会主义的萌发策略)的投资多于双凸镜形黑色种子(采取谨慎的萌发策略)的投资。异子蓬的花序类型为二歧聚伞花序, 单个果序含果实(种子)的数目为1-15个, 最多可分为4级。果序内果实的分布具有一定的规律: 第一级1个果实, 第二级2个, 第三级4个, 第四级8个。异子蓬优先将资源供给黑色种子的果实, 利用不同类型种子的发育顺序, 首先保证黑色种子的产出。具有较高的生殖分配和采取“谨慎”的生殖策略是异子蓬与其所处盐漠环境长期适应的结果。     

异子蓬二型性种子的种皮结构及离子含量差异

异子蓬(Borszczowia aralocaspica)是中亚荒漠区系特有的一年生盐生植物,其种子具有二型性。通过检测异子蓬二型性种子中的生理生化物质以及观察种皮的结构,比较异子蓬二型性种子中营养物质、养分全量以及种皮结构的差异,结果表明:(1)异子蓬的棕色种子只有一层薄种皮,而黑色种子不但具有一层薄的内种皮还有一层坚硬致密的外种皮;(2)棕色种子的质量大、质量频数分布较为集中,并且吸水迅速,在吸水率较低时就能够大量萌发;(3)棕色种子的可溶性糖、可溶性蛋白质、全氮、全磷以及K+、Na+、K+/Na+和Mg2+/Na+都显著高于黑色种子。说明在繁殖子代、产生二型性种子时,异子蓬对棕色种子和黑色种子之间的繁殖投入存在着巨大的差异,这可能是导致异子蓬二型性种子产生差异性萌发机制的重要原因。     

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。     

新疆干旱区植物藜的种子异型性及其萌发机理

新疆干旱区分布的植物藜(Chenopodium album)的种子有黑色和褐色两种类型。对藜的异型性种子从形态结构、不同环境因素及激素或化学物质对萌发的影响以及同工酶谱等方面进行了研究,并对其萌发及适应异质环境的机理进行了讨论。结果表明:(1)藜的异型性种子在形态结构、萌发休眠特性等方面都存在明显差异:黑色种子种皮厚且硬,休眠,萌发慢,萌发率低;褐色种子种皮薄而软,不休眠,萌发快且萌发率高;(2)黑色种子的休眠可通过切除胚根外缘种皮得以完全解除;(3)赤霉素、乙烯利对黑色种子的萌发无明显促进作用;KNO3可较显著促进黑色种子的萌发;协同使用乙烯利和KNO3时,可显著提高黑种子萌发率,完全打破休眠;(4)黑色种子和褐色种子的酯酶、过氧化物酶及过氧化氢酶同工酶谱带存在差异;(5)黑色种子的萌发需要光照,而褐色种子则对光不敏感;低温贮藏对二者的萌发均无显著影响,尽管黑色种子的萌发率有波动。研究结果初步显示黑色种子的休眠是内源(胚)和外源(种皮)因素共同所致。藜的种子异型性及其萌发机理的形成是其对新疆干旱区异质化环境的高度适应。     

准噶尔荒漠异苞滨藜(Atriplex micrantha)的种子二型性及其萌发行为

异苞滨藜的果实(种子)存在二型性,这两种类型的果实在形状、大小、颜色及包被其苞片的大小上均有差异。黑色果实果皮光滑,有光泽,直径1.481mm±0.186mm,包被果实的苞片较小;褐色果实扁平,直径2.642mm±0.254mm,包被果实的苞片较大。以异苞滨藜的两种果实(种子)为材料,在3个变温条件(5/25℃、5/15℃、15/25℃,暗12h/光12h)下进行萌发实验,褐色种子成熟后即具有一定的萌发能力,最终萌发率都在80%以上。黑色种子仅在低的夜间温度(5℃)和高的昼间温度(25℃)条件下有较高的萌发率(>70%),而在5/15℃、15/25℃条件下,种子的最终萌发率较低(<20%),2星期的低温层积处理能够有效地加速和提高黑色种子在3个变温条件下的萌发,划破果皮和种皮也能不同程度的加速和促进黑色种子的萌发,表明黑色种子处于非深度生理休眠状态。苞片中盐含量低,苞片对褐色种子萌发无抑制作用。异苞滨藜的果实(种子)二型性及其萌发行为是对荒漠异质环境的适应。     

盐地碱蓬二型性种子的萌发和休眠及生态适应特性

二型性种子可能增强植物(尤其是盐生植物)应对难于预测环境变化的能力。该研究以运城盐湖盐地碱蓬(Suaeda salsa)二型性种子(黑色和棕色)为材料,采用室内观察和萌发方法对两种种子的形态、休眠和萌发特性进行比较,以揭示盐生植物二型性种子对盐渍化环境的适应对策。结果表明:(1)黑色种子体积小,种皮角质有光泽,而棕色种子体积大,种皮膜质无光泽。(2)新成熟的棕色种子在实验温度范围内萌发率均超过90%,而黑色种子仅在15/25℃下萌发最高(但低于85%),并且棕色种子萌发速度较黑色更快。(3)低盐(0.39mol/L)条件下,光照或黑暗对棕色种子萌发没有显著影响;高盐(0.39mol/L)条件下,棕色种子在黑暗中的萌发显著高于光照环境,而黑色种子在光照条件下的萌发率显著高于黑暗;所有温度和光照处理中,2种种子均随盐分浓度的升高而下降,但棕色种子在盐分浓度为0.78mol/L时萌发率仍能够达到30%以上,而黑色种子在盐分浓度大于0.59mol/L时几乎不萌发。(4)棕色种子为非休眠种子,而黑色种子具有非深度生理休眠特性;划破种皮、干储、低温层积和GA3处理均能够提高黑色种子的萌发率。研究发现,盐地碱蓬通过二型性种子休眠和萌发行为上的差异,采用"两头下注"对策提高其对盐渍化异质生境的适应性。     

埋藏对65种半干旱草地植物种子萌发特性的影响

比较了半干旱草原科尔沁沙地65种植物新采种子和埋藏1年种子的萌发特性.结果表明：埋藏1年后,有38种植物的种子仍具有活力.新采种子萌发率>80%的植物种子埋藏后萌发率下降；而萌发率<80%的植物种子埋藏后萌发率均有较大幅度的提高.埋藏对种子萌发特性的影响因植物生态类群(杂草、草原植物、草甸植物、沙生植物)而异.种子埋藏1年后,多年生植物萌发率没有明显提高,而1年生植物萌发率显著提高(P<0.05).对于新采种子和埋藏1年种子萌发率间相差10%以下的物种,可根据萌发格局将植物种分为3类：埋藏促进型(如差巴嘎蒿)、埋藏抑制型(如大籽蒿)和埋藏不变型(如鹤虱).     

Effects of germination time on seed morph ratio in a seed-dimorphic species and possible ecological significance

Background and Aims Diaspores of heteromorphic species may germinate at different times due to distinct dormancy-breaking and germination requirements, and this difference can influence life history traits. The primary aim of this study was to determine the effect of germination time of the two seed morphs of Suaeda corniculata subsp. mongolica on life history traits of the offspring.Methods Germinated brown and black seeds were sown on the 20th of each month from April to September in a simulated but near-natural habitat of the species. Phenological and vegetative traits of the maternal plants, and number, size and germination percentage of the offspring were determined.Key Results Germinated seeds sown late in the year produced smaller plants that had a higher proportion of non-dormant brown than dormant black seeds, and these brown seeds were larger than those produced by germinated seeds sown early in the year. The length of the seedling stage for brown seeds was shorter than that for black seeds, and the root/shoot ratio and reproductive allocation of plants from brown seeds were more variable than they were for plants from black seeds. Late-germinating brown seeds produced larger plants than late-germinating black seeds.Conclusions Altering the proportion of the two seed types in response to germination timing can help alleviate the adverse effects of delayed germination. The flexible strategy of a species, such as S. corniculata, that produces different proportions of dimorphic seeds in response to variation in germination timing may favour the maintenance and regeneration of the population in its unpredictable environment.     

Effects of salinity and nitrate on production and germination of dimorphic seeds applied both through the mother plant and exogenously during germination in Suaeda salsa

Salinity and nitrogen are two important environmental factors that affect the distribution of halophytes in their natural saline habitats. Seeds of the euhalophyte Suaeda salsa L. were harvested from plants that had been treated with 1 or 500 mm NaCl combined with 0.5 or 5 mm NO₃^?‐N (nitrate) for 115 days in a glasshouse. Germination was evaluated under different concentrations of NaCl and nitrate. Plants exposed to high salinity (500 mm ) and low nitrate (0.5 mm ) tended to produce heavy seeds. Either high salinity (500 mm ) or high nitrate (5 mm ) increased the brown/black seed ratio. The concentrations of Na⁺, K⁺, and Cl^? were higher in brown than in black seeds, and NO₃^? concentrations were higher in black than in brown seeds, regardless of NaCl and nitrate treatments during plant culture. Regardless of NaCl and nitrate concentrations during germination, seeds from plants grown with 0.5 mm nitrate generally germinated more rapidly than seeds from plants grown with 5 mm nitrate, and the difference was greater for black than for brown seeds. Exogenous nitrate during germination enhanced the germination of brown seeds less than that of black seeds. Producing more brown seeds and heavy black or brown seeds appears to be an adaptation of S. suaeda to saline environments. Producing more black seeds, which tend to remain dormant, should reduce competition for nitrogen and appears to be an adaptation to nitrogen‐limited environments. In conclusion, nitrate provided exogenously or by mother plants to black seeds may act as a signal molecule that enhances the germination of black S. suaeda seeds.     

The role of the seed coat in adaptation of dimorphic seeds of the euhalophyte Suaeda salsa to salinity

The role of the seed coat in adaptation of dimorphic seeds of the euhalophyte Suaeda salsa to salinity was investigated during germination and early seedling growth. Black and brown seeds were treated with chloroform for 1 min before the extract was used to analyze waxes and the seeds to investigate the protective role of the seed coat under saline conditions. Waxes in black seed coats were more abundant than those in brown seed coats. Salinity (500 mM NaCl) increased the concentration of Na⁺ and decreased the concentration of K⁺ in both black and brown seeds regardless of chloroform treatment. Chloroform treatment alone (in the absence of NaCl) had no effect on the concentration of Na⁺ or K⁺ in black or brown seeds and in the presence of 500 mM NaCl had no effect on the concentration of Na⁺ or K⁺ in brown seeds. However, chloroform treatment increased Na⁺ and decreased K⁺ in black seeds with 500 mM NaCl. A change of MDA (malondialdehyde) concentration in black and brown seeds treated with or without chloroform was similar to the change of Na⁺ concentration. High salinity (1500 mM NaCl) pretreatment for 40 days had a less adverse effect on germination of black seeds compared with brown seeds after they were transferred to fresh water regardless of chloroform treatment. Similar results were found for seedling emergence. In conclusion, a black seed coat may be more protective than a brown seed coat, probably by shielding the embryo from ion toxicity, because of its higher content of waxes. Thus black seeds can better maintain seed viability than brown seeds for extended periods under hypersaline conditions.     

散枝猪毛菜的种子多型性及其萌发行为

散枝猪毛菜(Salsola bracchita)主要分布于新疆准噶尔荒漠,具有很强的抗干旱和抗盐碱能力。该文对散枝猪毛菜的果实进行了观察,表明散枝猪毛菜具有4种类型的散布单位和果实,这4种果实在形状、大小、颜色和着生方式上均有显著差异。A型果实绿色、球形、大,着生方式为横生,宿存花被革质,背部有紫红色翅状附属物,直径为(2.161±0.138) mm,单粒重为(3.810±0.113) mg;B型果实绿色、大、扁平、长圆形,着生方式为直立,宿存花被革质,背部有翅状附属物,长为(2.062±0.188) mm,宽为(1.720±0.148) mm,单粒重为(2.665±0.113) mg;C型果实绿色、大、扁平、长圆形,着生方式为直立,宿存花被膜质,背部无翅状附属物,长为(2.239±0.277) mm,宽为(1.844±0.150) mm,单粒重为(2.723±0.559) mg;D型果实黄色、小、扁平、长卵形,着生方式为直立,宿存花被膜质,背部无翅状附属物,长为(1.678±0.163) mm,宽为(1.390±0.110) mm,单粒重为(0.928±0.025) mg。A型、B型和C型种子(果实)在5 ℃/15 ℃、5 ℃/25 ℃、15 ℃/25 ℃(暗12 h/光12 h)变温条件下萌发率>68%,且B型和C型种子比A型种子有较高的萌发率和萌发速率。D型种子在3种变温条件下萌发缓慢,最终萌发率始终维持在较低水平(<20%),划破果皮和种皮能够显著提高D型种子的萌发率和萌发指数,表明D型种子处于生理休眠状态。散枝猪毛菜的种子多型性是对荒漠异质环境的一种适应。     

Carry-over of differential salt tolerance in plants grown from dimorphic seeds of Suaeda splendens

Background and Aims

Halophytic species often show seed dimorphism, where seed morphs produced by a single individual may differ in germination characteristics. Particular morphs are adapted to different windows of opportunity for germination in the seasonally fluctuating and heterogeneous salt-marsh environment. The possibility that plants derived from the two morphs may also differ physiologically has not been investigated previously.

Methods

Experiments were designed to investigate the germination characteristics of black and brown seed morphs of Suaeda splendens, an annual, C₄ shrub of non-tidal, saline steppes. The resulting seedlings were transferred to hydroponic culture to investigate their growth and photosynthetic (PSII photochemistry and gas exchange) responses to salinity.

Key Results

Black seeds germinated at low salinity but were particularly sensitive to increasing salt concentrations, and strongly inhibited by light. Brown seeds were unaffected by light, able to germinate at higher salinities and generally germinated more rapidly. Ungerminated black seeds maintained viability for longer than brown ones, particularly at high salinity. Seedlings derived from both seed morphs grew well at high salinity (400 mol m⁻³ NaCl). However, seedlings derived from brown seeds performed poorly at low salinity, as reflected in relative growth rate, numbers of branches produced, F_v/F_m and net rate of CO₂ assimilation.

Conclusions

The seeds most likely to germinate at high salinity in the Mediterranean summer (brown ones) retain a requirement for higher salinity as seedlings that might be of adaptive value. On the other hand, black seeds, which are likely to delay germination until lower salinity prevails, produce seedlings that are less sensitive to salinity. It is not clear why performance at low salinity, later in the life cycle, might have been sacrificed by the brown seeds, to achieve higher fitness at the germination stage under high salinity. Analyses of adaptive syndromes associated with seed dimorphism may need to take account of differences over the entire life cycle, rather than just at the germination stage.Key words: Chlorophyll fluorescence, germination, growth rate, halophyte, photosynthesis, photosystem II, salt tolerance, seed dimorphism, seed viability, Suaeda splendens