环境因子对蕨类植物孢子萌发的影响

蕨类植物通过孢子萌发形成独立生活的配子体,配子体能够形成精子器和颈卵器,进而通过受精作用形成新的孢子体。孢子萌发是蕨类植物生活史过程中配子体世代向孢子体世代转变的关键步骤。同时,此过程不仅受到多种环境因子的影响,也是研究细胞核极性移动、细胞不对称分裂、假根极性生长等独特的细胞学事件的良好模型。迄今为止,人们已经研究发现多种环境因子对约200余种蕨类植物孢子萌发有影响。总结了环境因子对蕨类植物孢子萌发影响的规律如下:(1)孢子萌发除了受到光照强度影响外,主要受光质的影响,光质的影响主要表现为4种方式:①孢子萌发受红光刺激与远红光抑制像开关一样调控;②孢子萌发不受远红光抑制;③孢子萌发受蓝光抑制;④孢子只能在黑暗条件下萌发。(2)重力作用会影响孢子细胞核移动,进而影响孢子细胞发育的极性。(3)赤霉素(GA)能增加孢子萌发率或帮助孢子打破休眠。成精子囊素与GA作用相似,启动或促进孢子萌发。而脱落酸(ABA)、茉莉酸(JA)和乙烯等其它激素对孢子萌发的影响相对较小。(4)不同植物孢子有着各自最适的萌发培养基条件,如不同种类孢子对MS培养基中无机盐含量、蔗糖含量、pH值的要求不同。孢子外被中的Ca2+、Mn2+和Mg2+,培养基中的Cd2+和La3+,以及孢子接种密度、萌发空间CO2含量也会对孢子萌发造成影响。(5)多数蕨类植物孢子在15-30℃可以萌发,最适萌发温度为25℃。(6)4℃和液氮储藏可以延长孢子寿命并保持较高萌发率。     

蕨类植物孢子与种子植物花粉萌发的比较

蕨类植物孢子与种子植物花粉在有性生殖过程中都具有重要的作用。花粉作为种子植物的雄配子体,通过萌发后极性生长的花粉管将精细胞送到胚囊完成受精作用。蕨类植物孢子作为配子体的原始细胞,通过不对称的有丝分裂产生一大一小两个细胞,小细胞萌发出极性生长的假根,大细胞继续分裂发育为原叶体（配子体）。成熟的花粉和蕨类植物孢子都是代谢高度静止的细胞,两者的萌发过程不仅都受到各种不同环境因子的影响,而且在信号转导、极性建立和能量代谢等方面可能有着相似的调控机制。本文综述了蕨类植物孢子和种子植物花粉萌发过程的差异和保守性特征。     

蕨类植物孢子与种子植物花粉萌发的比较

蕨类植物孢子与种子植物花粉在有性生殖过程中都具有重要的作用。花粉作为种子植物的雄配子体, 通过萌发后极性生长的花粉管将精细胞送到胚囊完成受精作用。蕨类植物孢子作为配子体的原始细胞, 通过不对称的有丝分裂产生一大一小两个细胞, 小细胞萌发出极性生长的假根, 大细胞继续分裂发育为原叶体(配子体)。成熟的花粉和蕨类植物孢子都是代谢高度静止的细胞, 两者的萌发过程不仅都受到各种不同环境因子的影响, 而且在信号转导、极性建立和能量代谢等方面可能有着相似的调控机制。本文综述了蕨类植物孢子和种子植物花粉萌发过程的差异和保守性特征。     

蕨类植物性别分化对环境的响应

蕨类植物是维管植物中唯一的孢子体和配子体都能独立生活的类群.同型孢子蕨类配子体的性别分化受到激素和环境因子的影响.生理学研究表明,成精子囊素与赤霉素能诱导雄配子体发育,抑制雌配子体发育;脱落酸阻止成精子囊素诱导的精子器形成;乙烯合成前体ACC促进赤霉素诱导的精子器形成,而乙烯合成抑制因子AOA通过抑制细胞分化来抑制精子器形成.光照对不同种类蕨类配子体分化的影响存在差异.糖类能够促进雄配子体形成,并可加速成熟雌配子体向两性分化.钙离子、钴离子和甲硫氨酸等分别参与了蓝光和赤霉素对配子体性别分化的调控过程.培养密度影响配子体生长及性别表达,高密度下雄性和无性配子体居多,而低密度下两性和雌性配子体居多.近年来的突变体表型分析与分子生物学研究表明,成精子囊素通过影响ANI1、HER、TRA、FEM和MAN等基因的表达调控配子体性别分化.综述了蕨类植物性别分化对环境响应的研究进展.     

不同光照度下水蕨配子体叶绿素荧光特性的研究

以模式植物水蕨的两性配子体和雄配子体为材料,研究了不同光照度下蕨类植物配子体叶绿素荧光参数的差异,探讨水蕨配子体光系统Ⅱ(PSⅡ)对不同光照的响应规律.结果表明:同一光照度条件下,水蕨两性配子体的实际光化学效率(Ф_(PSⅡ))和潜在光化学效率(F_v/F_m)均高于雄配子体,但无显著差异,此时两者间的光化学猝灭系数(q_p)和非光化学猝灭系数(q_N)值也无显著差异;在4 000 lx光照条件下,两性配子体PSⅡ驱动的电子传递速率(ETR_(PSⅡ))显著高于雄配子体.随着光照度的增加,水蕨两性配子体的Ф_(PSⅡ)、ETR_(PSⅡ)和q_p值逐渐增加,且在4 000 lx光照下显著增加,但F_v/F_m和q_N值无显著变化;而雄配子体的Ф_(PSⅡ)、ETR_(PSⅡ)、F_v/F_m、q_p和q_N值均呈先增后减趋势,但各光照处理间无显著差异.研究发现,水蕨两性配子体PSⅡ的活性相对比雄配子体高;两性配子体和雄配子体的光保护能力相似;光照度对水蕨雄配子体的光合能力没有显著影响,而4 000 lx的光照度对水蕨两性配子体的生长最有利.     

我国的蕨类植物杂交种

蕨类植物是植物界中的一大类群,是靠孢子进行繁殖的孢子植物。它的生活史中具有一个独立生活的配子体世代和一个独立生活的孢子体世代。配子体极小,其上发育着精子器、颈卵器。蕨类植物的精子具鞭毛,受精过程离不开水,通过水精子才能达到颈卵器与卵子结合受精。在自然界潮湿的环境中许多不同种的蕨类植物的配子体生活在一起时,就有可能发生不同种的精子和卵子结合成受精卵而发育出杂交后代的可能性。但杂交后代能否正常发育并存     

蕨类植物配子体发育及其性器官分化的研究进展

作者概述了蕨类植物生活周期中不同发育阶段的特点,并着重介绍了光、植物生长调节物质、水和温度等因素对蕨类植物配子体发育的影响以及成精子囊素、光、钙离子和其它植物生长调节物质在性器官形成和分化过程中的作用。同时作者对蕨类植物性器官分化的不稳定性进行了探讨、并结合目前蕨类植物性器官分化的遗传学和分子生物学研究的最新进展,说明蕨类植物作为研究系统在植物生物学研究领域中的优势。     

天柱山黄山松种内与种间竞争的研究

研究无菌条件下不同浓度砷对蜈蚣草的孢子萌发、假根和原叶体生长发育的影响。结果表明,培养基质中As浓度大于30mg/L时,孢子萌发速度、假根和原叶体的生长发育受到抑制,假根的着生点位置发生改变;而与高浓度As影响不同的是,低浓度As(30mg/L)可促进孢子变绿的速率和初期原叶体(播后8d内)的生长。因此,高浓度As能抑制配子体生长发育并改变假根着生点位置,而低浓度As则可促进配子体早期的生长。     

瓦韦的“胎生现象”

蕨类植物的生活史中有2种植物体,即孢子体和配子体。其中配子体是能自养的、独立生存且有固定形态、发育模式、结构的植物体,它在蕨类植物生活史中占据较长的阶段．但常常不占优势．配子体世代通常是从孢予萌发开始,孢子萌发后发育为原叶体（即配子体）。存大多数蕨类植物中,     

亮毛蕨的配子体发育特征及其系统学意义

采用原生境土培养方法对亮毛蕨(Acystopteris japonica)进行培养,并对其配子体发育过程进行了观察。结果表明:亮毛蕨植物的孢子二面体型,极面观为椭圆形,赤道面观肾形,单裂缝,无周壁,外壁具棒状纹饰,孢子萌发为书带蕨型(Vittaria-type),原叶体发育为铁线蕨型(Adiantum-type)。丝状体4～7个细胞,片状体小楔状,仅3～5个细胞宽,成熟原叶体心形,裸露,初生假根具叶绿体,颈卵器较短;配子体发育较迅速,假根具分枝和膨大,精子器囊由3个壁细胞构成,颈卵器略弯曲。本研究首次观察到亮毛蕨的颈卵器壁细胞内含有球形颗粒,但其功能不详。亮毛蕨的配子体发育过程兼有原始和进化的特点。在适宜的光照和温度下,土壤培养亮毛蕨原叶体的成苗率达95%以上,移栽成活率达89%以上。     

Phytochrome-dependent modulation and re-induction of growth of the first rhizoid in Dryopteris paleacea Sw

Phytochrome-dependent growth in Dryopteris paleacea Sw. was investigated in young, developing gametophytes with respect to formation and differentiation of rhizoids. Under continuous red light (Rc), the first rhizoids grew synchronously by tip elongation at a constant rate of 240 μm · d⁻¹ until formation and outgrowth of the second rhizoid. Cessation of growth of the first rhizoids and outgrowth of the second rhizoids showed a correlation in time assumed to be mediated by intercellular signaling. The first rhizoids showed two modes of response to actinic irradiations: (i) modulation of rhizoid growth, and (ii) re-induction of growth in non-growing rhizoids. In the former, the promotory effect of actinic irradiations on rhizoids pre-cultured under Rc determined both the time for which rhizoids continued to grow after transfer into darkness and the final rhizoid length. In the latter, re-induced growth was studied using non-growing rhizoids which were obtained after irradiation with a far-red light (FR) pulse at the end of the pre-culture in Rc and transfer into darkness for 3 d to stop growth. Re-induction of growth occurred with a lag phase of 36 to 48 h after formation of the FR-absorbing form of phytochrome (Pfr) by a red light (R) pulse. From the incomplete R/FR reversibility it is evident that, here, coupling of Pfr to signal transduction is possible within minutes. Re-induction of growth possesses the advantage that the effect of actinic irradiations can be studied as an all-or-none response at the level of single gametophytes in future experiments. The present results clearly indicate that the developmental stage of the whole gametophyte, i.e. temporal and spatial patterns undergone during development, affects the regulation of rhizoid growth by the external factor light. Received: 8 June 1998 / Accepted: 22 December 1998     

Anomalous gravitropic response of Chara rhizoids during enhanced accelerations

Centrifugal accelerations of 50-250 g were applied to rhizoids of Chara globularis Thuill. at stimulation angles (alpha) of 5-90 degrees between the acceleration vector and the rhizoid axis. After the start of centrifugation, the statoliths were pressed asymmetrically onto the centrifugal flank of the apical cell wall. In contrast to the well-known bending (by bowing) under 1 g, the rhizoids responded in two distinct phases. Following an initial phase of sharp bending (by bulging), which is similar to the negatively gravitropic response of Chara protonemata, rhizoids stopped bending and, in the second phase, grew straight in directions clearly deviating from the direction of acceleration. These response angles (beta) between the axis of the bent part of the rhizoid and the acceleration vector were strictly correlated with the g-level of acceleration. The higher the acceleration the greater was beta. Except for the sharp bending, the shape and growth rate of the centrifuged rhizoids were not different from those of gravistimulated control rhizoids at 1 g. These results indicate that gravitropic bending of rhizoids during enhanced accelerations (5 degrees < or = alpha < or = 90 degrees) is caused not only by subapical differential flank growth, as it is the case at 1 g, but also by also by the centripetal displacement of the growth centre as was recently discussed for the negative gravitropism of Chara protonemata. A hypothesis for cytoskeletally mediated polar growth is presented based on data from positive gravitropic bending of Chara rhizoids at 1 g and from the anomalous gravitropic bending of rhizoids compared with the negatively gravitropic bending of Chara protonemata. The data obtained are also relevant to a general understanding of graviperception in higher-plant organs.     

Jasmonic acid promotes division of fern protoplasts, elongation of rhizoids and early development of gametophytes

The effects of jasmonic acid (JA) on spore germination, early gametophyte development and sporophytic protoplast culture of the fern Platycerium bifurcalum (Cav.) C. Chr. were investigated. JA and no influence on spore germination and primary rhizoid initiation, but significantly promoted early gametophyte development, which was evident from the longer primary rhizoids as well as the higher number of rhizoids and cells per gametophyte. Jasmonic acid (1 μ M ) also promoted the transition of gametophytes from a filamentous to a planar growth. Optimal primary rhizoid elongation and highest cell division activities in the gametophytes were observed at 0.01–1 μ M JA, while the highest number of rhizoids on gametophytes was obtained at 0.1–1 μ M JA. Jasmonic acid (0.01 μ M ) also stimulated initial protoplast divisions. Except for the experiment in which the effect of JA on germination was tested. JA concentrations exceeding 1 μ M did not promote cell elongation or cell division but were instead inhibitory. On the basis of these findings, we propose that JA may be involved in early stages of P. bifurcatum development.     

Association of spectrin-like proteins with the actin-organized aggregate of endoplasmic reticulum in the Spitzenkörper of gravitropically tip-growing plant cells

Spectrin-like epitopes were immunochemically detected and immunofluorescently localized in gravitropically tip-growing rhizoids and protonemata of characean algae. Antiserum against spectrin from chicken erythrocytes showed cross-reactivity with rhizoid proteins at molecular masses of about 170 and 195 kD. Confocal microscopy revealed a distinct spherical labeling of spectrin-like proteins in the apices of both cell types tightly associated with an apical actin array and a specific subdomain of endoplasmic reticulum (ER), the ER aggregate. The presence of spectrin-like epitopes, the ER aggregate, and the actin cytoskeleton are strictly correlated with active tip growth. Application of cytochalasin D and A23187 has shown that interfering with actin or with the calcium gradient, which cause the disintegration of the ER aggregate and abolish tip growth, inhibits labeling of spectrin-like proteins. At the beginning of the graviresponse in rhizoids the labeling of spectrin-like proteins remained in its symmetrical position at the cell tip, but was clearly displaced to the upper flank in gravistimulated protonemata. These findings support the hypothesis that a displacement of the Spitzenk?rper is required for the negative gravitropic response in protonemata, but not for the positive gravitropic response in rhizoids. It is evident that the actin/spectrin system plays a role in maintaining the organization of the ER aggregate and represents an essential part in the mechanism of gravitropic tip growth.     

Circular arrangement of cortical microtubules around the subapical part of a tip-growing fern protonema

Summary Cortical microtubule arrays in tip-growing protonemal and rhizoid cells of the fernAdiantum gametophytes were observed by immunofluorescence microscopy. A circular arrangement of cortical microtubules was demonstrated around the subapical part of protonemal cells growing under red light conditions. However, such an arrangement was not found in growing rhizoids either by immunofluorescence microscopy or by electron microscopy. The different patterns of microtubule arrays around the apices of tip-growing protonemal and rhizoid cells suggest the possible existence of different mechanisms in regulating the cell diameter in the two types of cylindrical cell.     

Observations on the mature gametophyte of Phylloglossum (Lycopodiaceae)

Mature, gametangia-bearing photosynthetic gametophytes of Phylloglossum can be grown from the nonphotosynthetic, cylindrical, negatively gravitropic immature stages in illuminated axenic culture on a nutrient medium with or without 0.2% glucose. The gametangial-bearing region of these gametophytes, the photosynthetic crown, grows horizontally from the apex of the immature cylindrical stage. At maturity the photosynthetic crown is thickened and bilaterally symmetrical. It is usually narrow and bean-shaped with dorsal and ventral regions. Occasionally, the ventral region becomes thicker and the crown is deltoid in cross section. The dorsal edge or ridge of the crown is the gametangial region with archegonia and antheridia. The gametangia are often hidden because they are interspersed among numerous uni- or bicellular paraphyses. The lateral surfaces below the gametangial region lack outgrowths, and the ventral region is covered with long rhizoids. The apical meristem at the anterior end of the crown is overarched by the developing dorsal tissue of the gametangial ridge. The lower derivatives of the meristem form the ventral region with rhizoids. Phylloglossum gametophytes are unusual because their mode of organic nutrition changes from mycorrhizal to photosynthetic during ontogeny.     

EFFECT OF 2-CHLOROETHYLTRIMETHYLAMMONIUM CHLORIDE ON FERN GAMETOPHYTES

Kelley , A. G., and S. N. Postlethwait . (Purdue U., Lafayette, Ind.) Effect of 2-chloroethyltrimethylammonium chloride on fern gametophytes. Amer. Jour. Bot. 49(7): 778–786. Illus. 1962.—2-chloroethyltrimethylammonium chloride (CCC) promotes growth of fern gametophytes by stimulating the rate of cell division. The transition from filamentous to biplanar to triplanar cell division is hastened, resulting in earlier and more numerous gametangia on treated prothalli. Growth is stimulated in the range of 10^-4 m –10^-2 m CCC, with the optimal effect at 10^-3 m CCC. The time sequence of rhizoid production is altered by 10^-2 m CCC; the resultant production of rhizoids by prothalli treated with 10^-2 m CCC does not exceed that of controls.     

Involvement of microtubules in rhizoid differentiation of Spirogyra species

Summary.  Some species of Spirogyra form rosette-shaped or rod-shaped rhizoids in the terminal cell of the filaments. In the present study, we analyzed an involvement of microtubules (MTs) in rhizoid differentiation. Before rhizoid differentiation, cortical MTs were arranged transversely to the long axis of cylindrical cells, reflecting the diffuse growth. At the beginning of rhizoid differentiation, MTs were absent from the extreme tip of the terminal cell. In the other area of the cell, however, MTs were arranged transversely to the long axis of the cell. In the fully differentiated rosette-shaped rhizoid, MTs were randomly organized. However, at a younger stage of rosette-shaped rhizoids, MTs were sometimes arranged almost transversely in the lobes of the rosette. In the rod-shaped rhizoid, MTs were arranged almost transversely. MT-destabilizing drugs (oryzalin and propyzamide) induced swelling of rhizoids, and neither rosette-shaped nor rod-shaped rhizoids were formed. The role of MTs in rhizoid differentiation was discussed. Received June 17, 2002; accepted November 11, 2002; published online April 8, 2003 RID="*" ID="*" Correspondence and reprints: Department of Life Science, Graduate School of Science, Himeji Institute of Technology, Harima Science Park City, Hyogo 678-1297, Japan.     

Staining and osmotic properties of young gametophytes ofPolypodium vulgare L. and their bearing on rhizoid function

Summary The rhizoids of gametophytes ofPolypodium vulgare L. rapidly absorb vital stains whereas the protonemal cells are impermeable to these stains, which can only enter the cells from the rhizoids. The protonemal cells which bear rhizoids were found to have a slightly higher osmotic equivalent than did the rhizoids or the protonemal cells on either side. From the results of several staining procedures it was demonstrated that the rhizoid walls contain free carboxyl groups and thus possess cation exchange properties. Most of the carboxyl groups are probably present in a yellow-brown wall matrix substance, which shows high resistance to acid and alkali extraction. The precise nature of this substance has not been determined but it could be an acid mucopolysaccharide. Carboxyl groups are detectable in the protonemal cell walls only after saponification and are probably esterified in the untreated wall. Several other chemical and physiological differences were found between the rhizoids and the protonemal cells and it was concluded that the specific properties of the rhizoids are related to their function as organs of uptake.     

Phototropism of Thalli and Rhizoids Developed from the Thallus Segments of Bryopsis hypnoides Lamouroux

Newly regenerated thalli were used to study the phototropism of Bryopsis hypnoides Lamouroux under different qualities of light. Positive phototropism in the thalli and negative phototropiam In the rhizoida of B. hypnoides were investigated and analyzed in terms of bending. Both thaiii and rhlzoids developed from thallus segments exhibited typical tip growth, and their photoreceptive sites for phototroplam were also restricted to the apical hemisphere. The bending curvature of rhizoids and thalli were determined with unilateral lights at various wavelengths and different fluence rates after a fixed duration of Illumination. The trends of bending from the rhizoid and thallus were coincident, which showed that the action spectrum had a large range, from ultraviolet radiation （366.5 nm） to green light （524 nm）. Based on the bending curvatures, blue light had the highest efficiency, while the efficiency of longer wavelengths （〉500 nm） was significantly lower. External Ca^2＋ had no effect on the bending curvature of thalli and rhlzolda. Blue light （440 nm） induced thallus branching from rhizoids, while red light （650 nm） had no such effect. Fast-occurring chloroplast accumulation In the outermost cytoplasmic layer of the blue light （440 nm）-Irradiated region In the rhizoid was observed, from which protrusions （new thalli） arose after 4 h of the onset of illumination, and this action was thought to be driven by the dynamics of actin microfilamenta.