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。     

不同生境盐地碱蓬出苗及幼苗抗盐性比较

研究了盐胁迫对不同生境盐地碱蓬出苗、幼苗生长、离子积累和荧光参数等的影响.盐地碱蓬种子具有二型性,即外种皮柔软而半透明的棕色种子和外种皮坚硬的黑色种子.两种生境盐地碱蓬棕色种子的出苗率均明显高于黑色种子.与黑色种子相比,潮间带生境盐地碱蓬棕色种子在高盐环境下的出苗速率和出苗率高于盐碱地生境盐地碱蓬.各处理盐分浓度下,潮间带生境盐地碱蓬叶片的最大光化学效率(Fv/Fm)、PSⅡ实际光化学效率(ΦPSⅡ)和地上部分生物量均低于盐碱地生境盐地碱蓬.高盐浓度下,潮间带生境盐地碱蓬叶片Na+和Cl-含量低于盐碱地生境盐地碱蓬.这些特征可能是盐地碱蓬长期适应不同生境的结果.     

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.     

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。     

盐地碱蓬二型性种子及其幼苗对盐渍环境的适应性

研究了盐地碱蓬二型性种子中离子含量与刚萌发幼苗耐盐性之间的关系,以及盐分对砂培盐地碱蓬二型性种子的幼苗生长、离子含量及光合特性的影响.棕色种子中离子含量显著高于黑色种子.与对照相比,100和400 mmol/L NaCl对棕色种子幼苗伸长没有抑制作用,却显著抑制黑色种子幼苗的伸长.NaCl处理下棕色种子的幼苗地上部分干重和主茎一级分枝数比黑色种子幼苗高,但二型性种子的幼苗叶片中离子含量、叶绿素含量及光合放氧速率却没有明显差异.上述结果说明盐地碱蓬棕色种子较高的离子含量可能是棕色种子刚萌发幼苗耐盐性较强的重要原因.棕色种子幼苗较高的生物量可能与其较多的分枝数有关.二型种子的这些特征可能决定了其在群落建成中所起到的不同作用.     

盐旱互作对不同生境盐地碱蓬种子萌发和幼苗生长的影响

研究了盐旱互作对潮间带和盐碱地生境盐地碱蓬棕色种子萌发、地上部生长和离子积累的影响。不同盐浓度预处理后,未萌发的种子风干后复水,其萌发率与对照相比没有降低,说明两种生境盐地碱蓬种子萌发期间都耐干湿交替。400mmol/LNaCl溶液浇灌的潮间带生境盐地碱蓬幼苗在第3次干旱处理后萎蔫幼苗的百分比高于盐碱地生境的,而复水后正常幼苗的百分比却相反。400mmol/LNaCl溶液处理下,第3次干旱处理复水后,盐碱地生境盐地碱蓬幼苗地上部离子含量(主要是Na+和Cl-)高于潮间带生境的。表明在干旱情况下,盐碱地生境盐地碱蓬幼苗能积累更多的无机离子,降低渗透势,提高根系吸收水分的能力。上述结果说明,盐碱地生境盐地碱蓬幼苗较潮间带生境盐地碱蓬幼苗更耐盐与干旱的交互作用。     

新疆短命植物小拟南芥(Arabidopsis pumila)种子萌发特性及其生态适应性

通过显微结构及不同处理条件下种子萌发率的观察,对早春短命植物小拟南芥(Arabidopsis pumila)种子萌发特性及影响因素进行了研究,并对其生态适应性进行了讨论。结果表明:(1)温度和光照变化对自然生境和温室收获种子的萌发率影响均不显著,说明此种群在前期萌发阶段对光、温不敏感;(2)自然生境中采收的小拟南芥种子萌发率显著低于温室收获种子,说明环境条件的变化对短命植物种子的发育具有重要作用,可显著改变种子的萌发行为;(3)赤霉素使自然生境收获种子胚活性增强从而对萌发有较大促进作用,可使萌发率增加50%以上;(4)对种皮进行各种机械损伤处理使得种皮松弛或透气,可以显著提高自然生境种子的萌发率(超过70%);(5)盐和干旱胁迫对种子萌发均具有明显的抑制作用,但复水后部分被抑制种子可重新萌发,显示盐和干旱胁迫可导致种子产生浅度休眠。结合小拟南芥自然生存环境及本研究的结果,显示其种子萌发特性与生境具有高度适应性。     

NaCl胁迫对2种表型盐地碱蓬种子萌发的渗透效应和离子效应研究

采用高低2个浓度的NaCl、LiCl及等渗甘露醇溶液处理紫红色表型(紫色型)和绿色表型(绿色型)盐地碱蓬种子,通过测定它们的种子萌发率、吸胀速率和胚内离子含量,研究NaCl胁迫对2种表型种子萌发的离子效应和渗透效应.结果表明:(1)2种表型盐地碱蓬种子萌发率在高浓度(300 mmol/L)和低浓度(100 mmol/L)NaCl处理下均显著降低,紫色型种子萌发率在低浓度下显著低于绿色型,而在高浓度下却显著高于绿色型;绿色型种子萌发率在高浓度(30 mmol/L)和低浓度(10 mmol/L)LiCl处理下均未受到显著影响,但紫色型种子萌发率却均极显著降低;2种表型盐地碱蓬种子萌发率在低浓度等渗甘露醇处理下均极显著低于低浓度NaCl处理,而高浓度等渗甘露醇处理却均与高浓度NaCl处理无显著差异.(2)2表型种盐地碱蓬种子的吸胀速率在低浓度NaCl处理下没有受到显著影响,但高浓度NaCl处理及与之等渗的高浓度甘露醇处理下都显著降低,而且紫色型种子的吸胀速率在等渗甘露醇处理时显著高于绿色型.(3)2种表型盐地碱蓬种子胚中的Na 含量和Na /K 在对照和低浓度NaCl处理下无显著差异,但紫色型种子胚中的Na 、K 含量在高浓度NaCl处理时都显著高于对照,且K 含量增加的幅度远大于Na 含量,导致紫色型种子胚中的Na /K 显著低于绿色型.研究发现,盐地碱蓬种子萌发在低浓度NaCl胁迫下主要受离子效应抑制,而高浓度NaCl胁迫下则主要受渗透效应抑制,紫色型种子萌发率在高浓度NaCl胁迫下高于绿色型的原因之一是前者能维持更低的Na /K 比.     

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。     

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。     

Germination of dimorphic seeds of the desert annual halophyte Suaeda aralocaspica (Chenopodiaceae), a C4 plant without Kranz anatomy

Background and Aims

Suaeda aralocaspica is a C₄ summer annual halophyte without Kranz anatomy that is restricted to the deserts of central Asia. It produces two distinct types of seeds that differ in colour, shape and size. The primary aims of the present study were to compare the dormancy and germination characteristics of dimorphic seeds of S. aralocaspica and to develop a conceptual model of their dynamics.

Methods

Temperatures simulating those in the natural habitat of S. aralocaspica were used to test for primary dormancy and germination behaviour of fresh brown and black seeds. The effects of cold stratification, gibberellic acid, seed coat scarification, seed coat removal and dry storage on dormancy breaking were tested in black seeds. Germination percentage and recovery responses of brown seeds, non-treated black seeds and 8-week cold-stratified black seeds to salt stress were tested.

Key Results

Brown seeds were non-dormant, whereas black seeds had non-deep Type 2 physiological dormancy (PD). Germination percentage and rate of germination of brown seeds and of variously pretreated black seeds were significantly higher than those of non-pretreated black seeds. Exposure of seeds to various salinities had significant effects on germination, germination recovery and induction into secondary dormancy. A conceptual model is presented that ties these results together and puts them into an ecological context.

Conclusions

The two seed morphs of S. aralocaspica exhibit distinct differences in dormancy and germination characteristics. Suaeda aralocaspica is the first cold desert halophyte for which non-deep Type 2 PD has been documented.Key words: Borszczowia, cold desert halophyte, physiological seed dormancy, seed germination, Suaeda     

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.     

Germination dimorphism in Suaeda acuminata: A new combination of dormancy types for heteromorphic seeds

Desert annual Suaeda acuminata produces two morphologically distinct types of seeds on the same plant. The main aims of our study were to compare germination characteristics of the dimorphic seeds, ascertain their dormancy types and give the hormonal explanation. The two seed types of S. acuminata absorbed water at different rates with brown seeds imbibing water faster. Germination percentages of brown seeds were significantly higher than those of black seeds in all temperature and light regimes tested. Eight weeks of cold stratification did not break dormancy of black seeds, whereas exogenous GA₃ promoted germination. Excised embryos of untreated black seeds produced normal seedlings. Contents of ZR, GA₃ and ABA of brown seeds were significantly higher than that of black seeds; while contents of IAA of black seeds were significantly higher than that of brown seeds. Brown seeds of S. acuminata are non-dormant, whereas black seeds have intermediate physiological dormancy (PD). Interaction among ZR, ABA and GA₃ may play an important role in dormancy status of both seed types. This is the first report of non-dormancy and intermediate PD in a heteromorphic species.     

Comparison of germination and seed bank dynamics of dimorphic seeds of the cold desert halophyte Suaeda corniculata subsp. mongolica

Background and Aims

Differences in dormancy and germination requirements have been documented in heteromorphic seeds of many species, but it is unknown how this difference contributes to maintenance and regeneration of populations. The primary aim of this study was to compare the seed bank dynamics, including dormancy cycling, of the two seed morphs (black and brown) of the cold desert halophyte Suaeda corniculata and, if differences were found, to determine their influence on regeneration of the species.

Method

Seeds of the two seed morphs were buried, exhumed and tested monthly for 24 months over a range of temperatures and salinities, and germination recovery and viability were determined after exposure to salinity and water stress. Seedling emergence and dynamics of the soil seed bank were also investigated for the two morphs.

Key Results

Black seeds had an annual dormancy/non-dormancy cycle, while brown seeds, which were non-dormant at maturity, remained non-dormant. Black seeds also exhibited an annual cycle in sensitivity of germination to salinity. Seedlings derived from black seeds emerged in July and August and those from brown seeds in May. Seedlings were recruited from 2·6 % of the black seeds and from 2·8 % of the brown seeds in the soil, and only 0·5 % and 0·4 % of the total number of black and brown seeds in the soil, respectively, gave rise to seedlings that survived to produce seeds. Salinity and water stress induced dormancy in black seeds and decreased viability of brown seeds. Brown seeds formed only a transient soil seed bank and black seeds a persistent seed bank.

Conclusions

The presence of a dormancy cycle in black but not in brown seeds of S. corniculata and differences in germination requirements of the two morphs cause them to differ in their germination dynamics. The study contributes to our limited knowledge of dormancy cycling and seed bank formation in species producing heteromorphic seeds.     

The effect of plant growth regulators, nitric oxide, nitrate, nitrite and light on the germination of dimorphic seeds of Suaeda salsa under saline conditions

Suaeda salsa, a leaf succulent shrub in the family Chenopodiaceae, is one of the most important halophytes in China. Suaeda salsa produces dimorphic seeds (soft brown seeds and hard black seeds). Seeds of S. salsa were collected from the coastal salt flats near Huanghua City, China. Experiments were conducted to determine the salinity-alleviating effect of plant growth regulators, nitric oxide, nitrate, nitrite and light on the germination of dimorphic seeds of S. salsa. Brown seeds had a higher germination rate than black seeds in all experiments. Black seeds were more sensitive to salt in the absence of light in comparison to brown seeds. Brown seeds absorbed water more quickly in comparison to black seeds and were found to be more tolerant of salt stress. Our results showed that 1-aminocyclopropane-1-carboxylate (ACC, the immediate precursor of ethylene), nitrite, GA₄ and BA improved seed germination in the presence of salt. However, nitrate, GA₁, GA₃ failed to alleviate salt stress. ABA inhibited seed germination and seedling growth. Possible mechanisms involved in the alleviation of salt stress in S. salsa seeds and the ecological adaptation of the seeds to the environment are discussed.     

Dormancy and germination responses of halophyte seeds to the application of ethylene

Ethylene is invariably produced during seed germination but its role in regulating seed dormancy and germination is poorly understood. Seeds of 22 halophytic species having different life forms – salt secreting dicots, salt secreting monocots, stem succulents and leaf succulents were germinated in Petri dishes kept in a growth chamber set at 20/30 °C (night/day) temperature and a 12 hr light period. Sodium chloride and ethephon were added to the medium from the beginning of the experiment. Seed germination was recorded every other day for twenty days. Application of ethylene did not have any significant effect on releasing seeds from innate dormancy. However, it appeared to have a role in alleviation of salinity effects which varied from negative in certain species to almost complete alleviation of high salinity effects in others.Our data indicates that ethylene appears to have little role in breaking innate seed dormancy however, in most halophyte seeds studied, application of ethylene alleviate the salinity effect to various degrees. Halophyte seeds which could germinate under saline conditions approaching twice the salinity of seawater may offer clues to understand management of seed germination under highly saline conditions. To cite this article: M.A. Khan et al., C. R. Biologies 332 (2009).     

Variation of seed heteromorphism in Chenopodium album and the effect of salinity stress on the descendants

Background and Aims

Chenopodium album is well-known as a serious weed and is a salt-tolerant species inhabiting semi-arid and light-saline environments in Xinjiang, China. It produces large amounts of heteromorphic (black and brown) seeds. The primary aims of the present study were to compare the germination characteristics of heteromorphic seeds, the diversity of plant growth and seed proliferation pattern of the resulting plants, and the correlation between NaCl stress and variation of seed heteromorphism.

Methods

The phenotypic characters of heteromorphic seeds, e.g. seed morphology, seed mass and total seed protein were determined. The effects of dry storage at room temperature on dormancy behaviour, the germination response of seeds to salinity stress, and the effect of salinity on growth and seed proliferation with plants derived from different seed types were investigated.

Key Results

Black and brown seeds differed in seed morphology, mass, total seed protein, dormancy behaviour and salinity tolerance. Brown seeds were large, non-dormant and more salt tolerant, and could germinate rapidly to a high percentage in a wider range of environments; black seeds were salt-sensitive, and a large proportion of seeds were dormant. These characteristics varied between two populations. There was little difference in growth characteristics and seed output of plants produced from the two seed morphs except when plants were subjected to high salinity stress. Plants that suffered higher salinity stress produced more brown (salt-tolerant) seeds.

Conclusions

The two seed morphs of C. album exhibited distinct diversity in germination characteristics. There was a significant difference in plant development and seed proliferation pattern from the two types of seeds only when the parent plants were treated with high salinity. In addition, seed heteromorphism of C. album varied between the two populations, and such variation may be attributed, at least in part, to the salinity.     

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。