安徽大龙山国家森林公园东亚小金发藓配子体与孢子体个体性状及其相互关系

植物性状是植物对环境变化响应和适应的综合反映。目前对苔藓植物个体水平功能性状的研究较为匮乏。以安徽大龙山国家森林公园分布的东亚小金发藓（Pogonatum inflexum）雌株为例,在其孢子体成熟期测定地上部分孢子体和配子体的形态特征及生物量,系统分析了植株性状变异特征、异速生长关系及协变（整合）格局。结果表明,东亚小金发藓配子体形态性状变异性高于孢子体,且二者生物量变异系数最大。孢子体高度是配子体的2倍,但其生物量仅占地上部分的23%。孢子体和配子体功能性状之间有一定相关性;形态性状（Y）与生物量（X）之间多为指数<1.0异速生长关系,而孢子体和配子体生物量之间为等速生长关系（指数为1.135±0.158）。孢子体生物量分配比例随个体增大而显著减小,体现出显著的负的个体大小依赖。主成分分析表明,东亚小金发藓孢子体和配子体主要个体性状具有不同的协变方向（即两个不同的性状群）,其中孢子体生物量是两类性状群的关键结点。综合来看,东亚小金发藓个体性状间具有与维管植物相似的协变关系。     

The composition of lipids and fatty acids determined at various stages of haploid and diploid generations in the fern Adiantum capillus-veneris

The composition of lipids and fatty acids was determined at various developmental stages of gametophytes and sporophytes in the fern Adiantum capillus-veneris L. When the lipid content was expressed per g fresh weight, triacylglycerol was a major reserve lipid in spores and remained the dominant lipid in the gametophytes up to the 15th day after germination, but it was a minor one in the pinnae of sporophytes. In contrast, triterpenoids were abundant in sporophytes but not detected in gametophytes. When the content of fatty acids was expressed in percentage of the total fatty acids, the content of arachidonic acid in phosphatidylcholine and linolenic acid in diacyglyceryltrimethylhomoserine and phosphatidylcholine was significantly higher in both pinnae and petioles of sporophtes than in gametophytes. Significant differences in fatty acid composition of phosphatidylethanolamine, phosphatidylserine, phosphatidylinositol and triacylglycerol were also noted between gametophytes and pinnae of sporophytes. All these differences were attributed to the difference of generation rather than tissue differences or age.
The content of monogalactosyldiacylglycerol, hexadecatrienoic acid in monogalactosyldiacylglycerol, and 3- trans hexadecenoic acid in phosphatidylglycerol was higher in the pinnae of sporophytes than in gametophytes or in young leaves of sporophytes. Electron microscopy of the ultrastructure of chloroplasts indicated that the high content of these components was consistent with the development of chloroplasts in pinnae.     

The evolution of root hairs and rhizoids

Background

Almost all land plants develop tip-growing filamentous cells at the interface between the plant and substrate (the soil). Root hairs form on the surface of roots of sporophytes (the multicellular diploid phase of the life cycle) in vascular plants. Rhizoids develop on the free-living gametophytes of vascular and non-vascular plants and on both gametophytes and sporophytes of the extinct rhyniophytes. Extant lycophytes (clubmosses and quillworts) and monilophytes (ferns and horsetails) develop both free-living gametophytes and free-living sporophytes. These gametophytes and sporophytes grow in close contact with the soil and develop rhizoids and root hairs, respectively.

Scope

Here we review the development and function of rhizoids and root hairs in extant groups of land plants. Root hairs are important for the uptake of nutrients with limited mobility in the soil such as phosphate. Rhizoids have a variety of functions including water transport and adhesion to surfaces in some mosses and liverworts.

Conclusions

A similar gene regulatory network controls the development of rhizoids in moss gametophytes and root hairs on the roots of vascular plant sporophytes. It is likely that this gene regulatory network first operated in the gametophyte of the earliest land plants. We propose that later it functioned in sporophytes as the diploid phase evolved a free-living habit and developed an interface with the soil. This transference of gene function from gametophyte to sporophyte could provide a mechanism that, at least in part, explains the increase in morphological diversity of sporophytes that occurred during the radiation of land plants in the Devonian Period.     

The evolutionary ecology (evo-eco) of plant asexual reproduction

The different modes of asexual plant reproduction are reviewed in the context of the evolutionary ecology (evo-eco) of retaining old (and colonizing new) habitats and exploring new ones. The modes of asexual reproduction are contrasted between the green algae and the nonvascular and vascular land plants (bryophytes and tracheophytes, respectively). The chronology of their appearance is also traced in the fossil record. Among bryophytes, the indeterminate growth of protonema and the subsequent fragmentation of gametophytes provide a successful asexual reproductive stratagem, one that is paralleled by indeterminate rhizomaty and subsequent fragmentation of tracheophyte sporophytes. Self-compatible (autogamous) bisexual (monoicous) gametophytes capable of producing viable homozygous sporophytes offer a method for colonizing new sites by means of a single spore, a stratagem that is paralleled in tracheophyte sporophytes by the co-option of sexual reproductive structures (e.g., flowers) to produce diploid embryos asexually (e.g., variants of apomixis such as nucellar embryony). A review of the fossil record and the biology of extant taxa show that some of these modes of asexual reproduction are ancient and present in the green ancestors of the land plants (i.e., the charophycean algae). An overview of the data indicates that asexual reproduction provides a bet-hedging stratagem that can obviate some of the many costs of sexual reproduction, and preserve well-adapted genomes within previously occupied sites.     

Vegetative and reproductive innovations of early land plants: implications for a unified phylogeny

As the oldest extant lineages of land plants, bryophytes provide a living laboratory in which to evaluate morphological adaptations associated with early land existence. In this paper we examine reproductive and structural innovations in the gametophyte and sporophyte generations of hornworts, liverworts, mosses and basal pteridophytes. Reproductive features relating to spermatogenesis and the architecture of motile male gametes are overviewed and evaluated from an evolutionary perspective. Phylogenetic analyses of a data set derived from spermatogenesis and one derived from comprehensive morphogenetic data are compared with a molecular analysis of nuclear and mitochondrial small subunit rDNA sequences. Although relatively small because of a reliance on water for sexual reproduction, gametophytes of bryophytes are the most elaborate of those produced by any land plant. Phenotypic variability in gametophytic habit ranges from leafy to thalloid forms with the greatest diversity exhibited by hepatics. Appendages, including leaves, slime papillae and hairs, predominate in liverworts and mosses, while hornwort gametophytes are strictly thalloid with no organized external structures. Internalization of reproductive and vegetative structures within mucilage-filled spaces is an adaptive strategy exhibited by hornworts. The formative stages of gametangial development are similar in the three bryophyte groups, with the exception that in mosses apical growth is intercalated into early organogenesis, a feature echoed in moss sporophyte ontogeny. A monosporangiate, unbranched sporophyte typifies bryophytes, but developmental and structural innovations suggest the three bryophyte groups diverged prior to elaboration of this generation. Sporophyte morphogenesis in hornworts involves non-synchronized sporogenesis and the continued elongation of the single sporangium, features unique among archegoniates. In hepatics, elongation of the sporophyte seta and archegoniophore is rapid and requires instantaneous wall expandability and hydrostatic support. Unicellular, spiralled elaters and capsule dehiscence through the formation of four regular valves are autapomorphies of liverworts. Sporophytic sophistications in the moss clade include conducting tissue, stomata, an assimilative layer and an elaborate peristome for extended spore dispersal. Characters such as stomata and conducting cells that are shared among sporophvtes of mosses, hornworts and pteridophytes are interpreted as parallelisms and not homologies. Our phylogenetic analysis of three different data sets is the most comprehensive to date and points to a single phylogenetic solution for the evolution of basal embryophytes. Hornworts are supported as the earliest divergent embryophyte clade with a moss/liverwort clade sister to tracheophytes. Among pteridophytes, lycophytes are monophyletic and an assemblage containing ferns, Equisetum and psilophytes is sister to seed plants. Congruence between morphological and molecular hypotheses indicates that these data sets are tracking the same phylogenetic signal and reinforces our phylogenetic conclusions. It appears that total evidence approaches are valuable in resolving ancient radiations such as those characterizing the evolution of early embryophytes. More information on land plant phylogeny can be found at: http: //www.science.siu.edu/ landplants/index.html.     

How was apical growth regulated in the ancestral land plant? Insights from the development of non-seed plants

Land plant life cycles are separated into distinct haploid gametophyte and diploid sporophyte stages. Indeterminate apical growth evolved independently in bryophyte (moss, liverwort, and hornwort) and fern gametophytes, and tracheophyte (vascular plant) sporophytes. The extent to which apical growth in tracheophytes co-opted conserved gametophytic gene networks, or exploited ancestral sporophytic networks, is a long-standing question in plant evolution. The recent phylogenetic confirmation of bryophytes and tracheophytes as sister groups has led to a reassessment of the nature of the ancestral land plant. Here, we review developmental genetic studies of apical regulators and speculate on their likely evolutionary history.

The combined results of recent developmental genetics and phylogenetics studies suggest that the ancestral sporophyte was more complex than previously thought.     

PHOTOINHIBITION AND RECOVERY AFTER HIGH LIGHT STRESS IN DIFFERENT DEVELOPMENTAL AND LIFE-HISTORY STAGES OF LAMINARIA SACCHARINA (PHAEOPHYTA)1,2

The capacity to cope with high light stress was investigated in different life-history and developmental stages of Laminaria saccharina Lamour. sporophytes and gametophytes. Changes in photosynthetic efficiency and in the level of photoinhibition were measured by in vivo fluorescence changes of photosystem II. Pigment content was studied using high performance liquid chromatography. Additionally, the morphology of the various developmental stages during the life cycle was studied by light microscopy in relation to the photosynthetic parameters. High light stress (2 h, 500 μmol.m-².s^?1) induced photoinhibition of photosynthesis with fast kinetics in older sporophytes and gametophytes. In contrast, the absolute degree of photoinhibition after light stress was higher in younger than in older sporophytes. Photosynthesis recovered faster in older sporophytes and gametophytes compared to young sporophytes. In very young sporophytes, photosynthesis did not recover fully even after 12 h exposure to low light, indicating severe photodamage. Kinetics of recovery in old sporophytes and in gametophytes showed a fast and a slow phase, whereas younger sporophytes recovered only with a slow phase, The fast phase is indicative of a decline of the photoprotective process, whereas the slow phase indicates a recovery from photodamage. The capacity to cope with high light stress in Laminaria sporophytes increased with increasing age of the thalli. The gametophytes are less sensitive to high light stress and may be selected to endure unfavorable white light conditions. Investigation of the xanthophylls showed that the higher resistance to high light is not caused solely by a higher content of xanthophyll cycle pigments. Additionally, changes in the thallus structure during the development of the sporophytes seemed to cause a higher resistance to high light. The observed changes in the ability to cope with high light in the different life-history and developmental stages of Laminaria saccharina may influence the distribution of the species on the shore.     

Reproductive biology of the fernPhegopteris decursive-pinnata

Four diploid plants and four tetraploid plants ofPhegopteris decursive-pinnata were investigated for determination of the reproductive characteristics of their gametophytes and two major features were recognized. First, gametophytes of the diploids showed an ontogenetic sequence of gametangium formation which is unfavorable for intragametophytic selfing, whereas those of the tetraploids showed that favorable for intragametophytic selfing. Second, 41 to 72% of the isolated gametophytes of the diploids produced sporophytes in the intragmetophytic selfing tests, whereas all of the isolated gametrophytes of the tetraploids produced sporophytes in the tests. Based on these developmental and genetic features of gametophytes, the dissimilar mating systems of the diploids and the tetraploids of this species are discussed.     

SEASONAL LIGHT AND TEMPERATURE INTERACTION EFFECTS ON DEVELOPMENT OF LAMINARIA SACCHARINA (PHAEOPHYTA) GAMETOPHYTES AND JUVENILE SPOROPHYTES1

In previous studies, Laminaria saccharina L. (Lamour.) sporophytes were found to exhibit two major peaks of sporogenesis and an annual life cycle in Long Island Sound, New York. Young sporophytes were observed shortly after the sporogenesis peaks in early autumn and spring, but most of the mature sporophytes decayed during summer. A new study was conducted to determine if the spring sporogenesis activity contributed to the recruitment observed in autumn through oversummering of gametophytic and juvenile sporophytic stages, as previously suggested. Reproduction and growth in gametophytes and growth in juvenile sporophytes were studied under crossed gradients of light and temperature. Periodic outplantings of substrata seeded with gametophytic and sporophytic stages to the field were conducted to assess actual survival. The optimum temperature and light conditions for gametophyte development, growth and reproduction varied with the time of year meiospores were obtained. Most of this variation was attributable to temperature. A seasonal adaptation to temperature in most developmental stages was observed. Higher temperatures resulted in greater numbers of male gametophytes. Gametophytic stages could develop at all times, suggesting that oversummering in this stage was possible. Juvenile sporophytes had a narrower optimum temperature range and again photon fluence rate contributed little to observed variances. Out planting of sporophytic stages at various times during the year indicated only sporophytes prepared from autumn and winter could survive summer conditions. The thalli of these plants grew rapidly in spring and eroded back to the meristematic region in summer. Most of these plants then quickly became reproductive, resulting in another autumn sporogenesis peak. Gametophytic and sporophytic outplantings prepared from spring meiospores did not survive the summer. Thus, the recruitment observed in autumn can only be the result of the previous autumn's sporogenesis activity. The sporogenous activities of spring and early summer appear to be unimportant, despite the fact that all reproductive indices are superior at those times.     

A methodology for large-scale Athyrium sheareri gametophyte proliferation and sporophyte production using tissue culture

Tissue culture methods using gametophytes are considered the easiest ways to mass-produce fern sporophytes. The aim of this study was to develop a practical propagation method for the ornamental fern, Athyrium sheareri. The gametophytes obtained from in vitro spore germination were used as experimental materials. We used the chopping method to investigate the culturing conditions for proliferating gametophytes and the blending method for evaluating the mass production of sporophytes in mixed soil. Gametophyte proliferation was determined via Knop medium, various concentrations of Murashige and Skoog (MS) basal medium (1, 1/2, 1/4), and media components (sucrose, nitrogen source, and activated charcoal). The fresh weight of the gametophytes increased by more than 24-fold in 1/2 MS medium. In addition, 1 g of gametophyte could produce a maximum of 255.3 sporophytes in a mixed soil of 7.5 cm² area. Treating gametophytes with exogenous plant growth regulators promoted the formation and growth of sporophytes. The cultivated young sporophytes were acclimated and successfully grown in greenhouses. We developed a mass production protocol for A. sheareri sporophytes suitable for field application, which is expected to have commercial value.

     

银粉背蕨配子体及幼孢子体发育的研究

银粉背蕨是一种小型观赏蕨类植物,但目前我国对该蕨的研究还不够成熟。本文利用改良Knop's培养基和腐殖土培养银粉背蕨的孢子,观察其配子体及幼孢子体形态发育特征,并研究了其配子体发育的最适培养基pH值。研究结果显示：（1）银粉背蕨孢子黄褐色,具三裂缝,极面观三角圆形,赤道面观为近半圆形,孢子具网状纹饰;孢子萌发为书带蕨型;原叶体发育为水蕨型;颈卵器和精子器为薄囊蕨型;成熟原叶体为对称的心脏形,不具毛状体;上述特征为银粉背蕨孢子和配子体发育的稳定特征。（2）培养基pH值在7.0~9.0时随着碱性的增强,银粉背蕨孢子萌发和配子体生长发育速度逐渐增加。（3）利用腐殖土培养银粉背蕨孢子,7~8周可发育成幼叶,成苗率达90%,成苗健壮,根系发达,是扩繁银粉背蕨的适宜方式。本文为资源植物银粉背蕨人工繁殖和演化研究提供科学依据。     

Contribution of fern gametophytes to the growth of produced sporophytes on the basis of carbon gain

Sporophytes appeared on most gametophytes of Thelypteris palustris (Salisb.) Schott that reached a certain size, which is interpreted to be a critical size of gametophytes for the production of sporophytes. After sporophytes were produced, attached gametophytes ceased dry weight growth, but the gametophytes which did not produce sporophytes grew successively. It was hypothesized that matter produced by gametophytes was being supplied to young sporophytes. Photosynthesis and respiration of gametophytes with attached sporophytes were not significantly different from that of gametophytes without sporophytes. Photosynthetic activity of gametophytes dropped from 0.18 to 0.03 mol CO₂ g^–1 s^–1 during the growth period. The higher photosynthetic rates of gametophytes in the early growth stage were important for reaching the critical size for sporophyte production in a short time. Sporophytes in the one leaf stage averaged 0.14 mol CO₂ g^–1 s^–1 of photosynthetic activity. The results show that sporophytes that had expanded the first leaf grow by their own photosynthetic production. Gametophytes allocated the photosynthate for sporophytes and it was an important aid before the one-leaf stage. The supportive role of gametophytes ended at that stage.     

Aglaophyton major, a non-vascular land-plant from the Devonian Rhynie Chert

EDWARDS, D. S., 1986. Aglaophyton major , a non-vascular land-plant from the Devonian Rhynie Chert. The genus Aglaophyton is described to accommodate the plant formerly known as Rhynia major , re-examined using new material and techniques. The plant, which is a sporophyte, is suggested to have been shorter than originally supposed and to have formed extensive stands of decumbent axes. These features, with information on the angle of dichotomous branching and the frequency of sporangia, are incorporated into a new reconstruction of the plant. The conducting strand within the axes consists of three zones, a central region of thin-walled cells, a middle region of thick-walled cells and an outer region of thin-walled cells. There is no indication of differential wall thickenings in the cells of any of these zones and the strand is interpreted in terms of a bryophytic conducting strand, the inner two regions being comparable to hydroids and the outer region comparable to the leptoids found in polytrichaceous moss gametophytes and sporophyles. The plant thus exhibits characters intermediate between tracheophytes and some mosses and cannot be considered a member of either group; at the present time it is not assigned to a higher category. A rediagnosis of Rhynia gwynne-vaughanii , based on published results and new data, is also included.     

Gametophyte contribution to sporophyte growth on the basis of carbon gain in the fern <Emphasis Type="Italic">Thelypteris palustris</Emphasis>: effect of gametophyte organic-matter production on sporophytes

At an early stage of growth gametophytes support the sporophytes of ferns. Young sporophytes become independent of gametophytes when the first leaves develop. Although large fern gametophytes produce multiple archegonia simultaneously, only one sporophyte is typically established on one gametophyte. The number of sporophytes is believed to be controlled in two possible directions, from gametophyte to sporophyte or from preceding sporophyte to another sporophyte. To investigate the effects of gametophytes on their sporophytes, we studied the relationship between organic matter production by gametophytes and the growth of young sporophytes of Thelypteris palustris. We cut gametophytes in half (CGs) to reduce the gametophytes’ production of matter. There was no significant difference between the growth of sporophytes on intact gametophytes (IGs) and that on CGs. According to our estimates, based on the rate of organic matter production, the large gametophyte was able to produce two or more sporophytes. The resources required for CGs to make similar-sized sporophytes was twice that for IGs. In polyembryony each of the multiple sporophytes was similar in size to the single sporophytes. Resource limitation does not seem to explain why fern gametophytes establish single sporophytes.     

Isolation and culture of protoplasts from the marine green algaMonostroma angicava

Protoplasts were isolated enzymatically from gametophytes of the marine green algaMonostroma angicava. The protoplasts regenerated in PES medium after gradual reduction of the osmoticum. Three types of developmental process were recognized in the protoplast regeneration: an original type, in which the protoplasts regenerated into leafy gametophytes; an apogamic type, in which they regenerated into sporophytes; a callus type, in which they regenerated into callus-like tissues. The resulting gametophytes and apogamic sporophytes became fertile in successive cultures.     

HAPLOID PARTHENOGENETIC SPOROPHYTES OF LAMINARIA JAPONICA (PHAEOPHYCEAE)1

Parthenogenetic sporophytes were obtained from three strains of Laminaria japonica Areschoug. These sporophytes grew to maturity in the sea, producine spores that all grew into female gametophytes. These female gametophytes gave rise to another generation of parthenogenetic sporophytes during the next year, so that by the year 1990 parthenogenetic sporophytes had been cultivated for 12, 9, and 7 generations, respectively, for the three strains. When female gametophytes from parthenogenetic sporophytes were combined with normal male gametophytes, normal sporophytes that reproduced and gave rise to both female and male gametophytes were obtained. The parthenogenetic sporophytes were shorter and narrower than the normal sporophytes of the same strain. Chromosome counts on mature sporophytes showed that normal sporophytes (from fertilized eggs) were diploid (2n = approximately 40) and that the spores they produced were haploid (n = approximately 20), while nuclei from both somatic and sporangial cells in parthenogenetic sporophytes were haploid. All gametophytes were haploid. Young sporophytes derived from cultures with both female and male gametophytes were diploid, while young, sporophytes obtained from female gametophytes from parthenogenetic sporophytes had haploid, diploid, or polyploidy chromosome numbers. Polyploidy was associated with abnormal cell shapes. The presence of haploid parthenogenetic sporophytes should be use in breeding kelp strains with useful characteristics, since the sporophyte phenotype is expressed from a haploid genotype which can be more readily selected.     

Gametophyte development and reproduction of the Asian fern Polystichum polyblepharum (Roem. ex Kunze) C. Presl (Dryopteridaceae,Polypodiopsida)

Abstract

The gametophytic generation of Polystichum polyblepharum (Dryopteridaceae), including spore germination, morphological development of the gametophytes, major vegetative features and reproduction strategies, was studied. Spore germination was of the Vittaria model and the developmental pattern was of the Aspidium model. Adult gametophytes were cordate and hairy, with unicellular hairs located at the margins and at the prothallus surface. The marginal ones were secretory. The gametophytes produced archegonia located in the central area between the notch and the rhizoids, but antheridia were never detected. Archegonia were of the normal type described for leptosporangiate ferns. Apogamous sporophytes appeared from a cellular outgrowth developed just under the apical notch. Initially, the outgrowth appeared completely surrounded by glandular hairs and scales, which persisted throughout the subsequent stages of leaf development.     

Tissue-direct PCR, a rapid and extraction-free method for barcoding of ferns

Fern gametophytes and young sporophytes often provide too little material for DNA extraction and are particularly difficult to identify to genus. Here we developed an efficient procedure called 'Tissue-direct PCR', in which a slice of fern tissue is mixed with PCR reagents and primers, allowing certain genomic regions to be amplified directly in the thermal cycler. For these diminutive and featureless stages of ferns, Tissue-direct PCR combined with amplifying plant barcodes promises to make the identification of immature ferns easy and rapid. Tissue-direct PCR would also be very helpful for large-scale ecological studies surveying distribution and population structure.     

Generational differences in response to desiccation stress in the desert moss Tortula inermis

BACKGROUND AND AIMS: Active growth in post-embryonic sporophytes of desert mosses is restricted to the cooler, wetter months. However, most desert mosses have perennial gametophytes. It is hypothesized that these life history patterns are due, in part, to a reduced desiccation tolerance for sporophytes relative to gametophytes. METHODS: Gametophytes with attached post-embryonic sporophytes of Tortula inermis (early seta elongation phenophase) were exposed to two levels of desiccation stress, one rapid-dry cycle and two rapid-dry cycles, then moistened and allowed to recover, resume development, and/or regenerate for 35 d in a growth chamber. KEY RESULTS: Gametophytes tolerated the desiccation treatments well, with 93 % survival through regenerated shoot buds and/or protonemata. At the high stress treatment, a significantly higher frequency of burned leaves and browned shoots occurred. Sporophytes were far more sensitive to desiccation stress, with only 23 % surviving after the low desiccation stress treatment, and 3 % surviving after the high desiccation stress treatment. While the timing of protonemal production and sporophytic phenophases was relatively unaffected by desiccation stress, shoots exposed to one rapid-dry cycle produced shoots more rapidly than shoots exposed to two rapid-dry cycles. CONCLUSIONS: It is concluded that sporophytes of Tortula inermis are more sensitive to rapid drying than are maternal gametophytes, and that sporophyte abortion in response to desiccation results from either reduced desiccation tolerance of sporophytes relative to gametophytes, or from a termination of the sporophyte on the part of the gametophyte in response to stress.