Sex ratio and sex expression in an urban population of the silver moss,Bryum argenteum Hedw.

ABSTRACT

Introduction. A well-supported pattern among dioicous bryophytes is male rarity. However, few assessments of bryophyte sex ratios have been made across environmental gradients to assess the role of environment in shaping population sex ratios.

Methods. We systematically surveyed 200 shoots from a 20?m² urban population of Bryum argenteum, and regenerated each shoot apex until sex expression occurred (up to 315 days).

Key results. Female shoots outnumbered male shoots 132 to 68, giving a sex ratio of 1.94♀: 1♂. The female bias was found in two transects in higher light environments but not in the third transect, which had a lower light level and an equal sex ratio. Female shoots took longer than male shoots to reach gametangial induction (122 vs. 60 days) and longer to produce 5 inflorescences (120 vs. 80 days). Male shoots produced an average of 10× the total number of inflorescences compared to female shoots (34 vs. 3.5 inflorescences). Despite producing more inflorescences, male plants also produced more regenerant shoots, thus contradicting the prediction that a higher prefertilisation reproductive effort in males trades off with vegetative proliferation. Female plants harboured significantly more associated microbes than male plants.

Conclusions. Our results support the role of light in influencing sex ratios in this species, suggest that trade-offs between reproduction and vegetative growth may not be strong for males, and indicate a potential role of a sex-specific microbiome in influencing sex ratios.     

Hormonal Regulation of Gametangial Formation in the Moss Bryum argenteum Hedw

Phytohormones such as auxins, cytokinins, gibberellins, andabscisic acid differentially affect gametangial induction inmale and female clones of Bryum argenteum. Both IAA and GA³increased the percentage of fertile gametophores in the maleclone, and inhibited the response in the female clone. GA₃ wasmore effective than IAA in eliciting the response in the maleclone. Cytokinins, on the other hand, increased the productionof fertile gametophores in the female clone, and inhibited itslightly in the male clone. The two cytokinins tested (kinetinand DMAAP) were almost equally effective for the female clone. An Interaction of IAA and kinetin nullified their individualinhibitory effects on the female and male clones, respectively.Cyclic AMP prevented the inhibitory effect of kinetin in themale clone; whereas, in the female clone, it stimulated theresponse elicited by kinetin. Abscisic acid (ABA) acted as ageneral inhibitor of vegetative growth and gametangial inductionin this moss. However, the inhibition of gametangial inductionwas greater in the female clone which is also more sensitiveto ABA than the male clone.     

Thermal tolerance of dried shoots of the moss Bryum argenteum

We conducted laboratory experiments to determine the lethal temperatures of the shoots of dried Bryum argenteum and to determine how this restoration species responds to extreme environments. We specifically assessed changes in gene expression levels in the shoots of dried B. argenteum plants that were subjected to sudden heat shock (control (20 ± 2°C), 80°C, 100°C, 110°C or 120°C) followed by exposure to heat for an additional 10, 20, 30 or 60 min. After they were exposed to heat, the samples were placed in wet sand medium, and their survival and regeneration abilities were evaluated daily for 56 days. The results showed that lethal temperatures significantly reduced the shoot regeneration potential, delayed both shoot and protonemal emergence times and reduced the protonemal emergence area. In addition, the expression of nine genes (HSF3, HSP70, ERF, LEA, ELIP, LHCA, LHCB, Tr288 and DHN) was induced by temperature stress, as assessed after 30 min of exposure. Additionally, a new thermal tolerance level for dried B. argenteum – 120°C for 20 min – was determined, which was the highest temperature recorded for this moss; this tolerance exceeded the previous record of 110°C for 10 min. These findings help elucidate the survival mechanism of this species under heat shock stress and facilitate the recovery and restoration of destroyed ecosystems.     

Subcellular Localization of Adenylate Cyclase in the Shoot Apices of Bryum argenteum Hedw

Adenylate cyclase activity has been detected in the shoot apicesof Bryum argenteum at the ultrastructural level, using adenylylimidodiphosphate as substrate. Heavy deposits of the reactionproduct were observed along the plasma membrane, tonoplast andthe membranous system of developing plastids. No clear stainingwas associated with other subcellular membranes. Mosses, adenylate cyclase, Bryum, cyclic AMP     

Effect of Chelating Agents and Metal Ions on Gametangial Formation in the Moss Bryum argenteum Hedw

Chelating agents such as EDTA and EDDHA markedly stimulate theformation of gametangia in the dioecious moss Bryum argenteum,and the effect is more pronounced on male than on female clones.EDTA-stimulated gametangial formation is associated with significantincreases in endogenous iron and copper. Ferric citrate alsoenhances gametangial formation, but copper sulphate is inhibitory.The present findings are discussed in the context of earlierinvestigations on other plants in an attempt to explain thepossible involvement of chelating agents and metal ions in stimulatingthe onset of the reproductive phase in this moss. Bryum argenteum, gametangial formation, chelating agents, metal ions     

How to define nativeness in vagile organisms: lessons from the cosmopolitan moss Bryum argenteum on the island of Tenerife (Canary Islands)

The distinction between native and introduced biotas presents unique challenges that culminate in organisms with high long‐distance dispersal capacities in a rapidly changing world. Bryophytes, in particular, exhibit large distribution ranges, and some species can truly be qualified as cosmopolitan. Cosmopolitan species, however, typically occur in disturbed environments, raising the question of their nativeness throughout their range. Here, we employ genetic data to address the question of the origin of the cosmopolitan, weedy moss Bryum argenteum on the island of Tenerife. The genetic diversity of B. argenteum on Tenerife was comparable to that found in continental areas due to recurrent colonisation events, erasing any signature of a bottleneck that would be expected in the case of a recent colonisation event. The molecular dating analyses indicated that the first colonisation of the island took place more than 100,000 years ago, i.e. well before the first human settlements. Furthermore, the significant signal for isolation‐by‐distance found in B. argenteum within Tenerife points to the substantial role of genetic drift in establishing the observed patterns of genetic variation. Together, the results support the hypothesis that B. argenteum is native on Tenerife; although the existence of haplotypes shared between Tenerife and continental areas suggests that more recent, potentially man‐mediated introduction also took place. While defining nativeness in organisms that are not deliberately introduced, and wherein the fossil record is extremely scarce, is an exceedingly challenging task, our results suggest that population genetic analyses can represent a useful tool to help distinguish native from alien populations.     

RAPD analysis of genetic variation and dispersal of the moss Bryum argenteum in Ross Island and Victoria Land, Antarctica

The Random Amplified Polymorphic DNA technique was used to assess the level of genetic diversity in Bryum argenteum from Ross Island and southern Victoria Land, Antarctica. Samples were collected from three separate transects, and from other geographically distinct populations within 150 km of Ross Island. Moss growth in two transects, sampled down small exposed meltstream channels at Cape Royds and Cape Chocolate, was very sparse with no other moss colonies found within 0.4 or 4 km, respectively. However, samples from these channels showed similar levels of genetic variation to those from a transect at Granite Harbour, where moss colonies were large, luxuriant and turf-like between boulders. In all transects, high levels of genetic diversity were apparent both within and between colonies, and some spatial relationships were observed down the length of the channels, with more extensive variation at the top than the bottom of two transects. Samples from other sites in the region showed varying but high levels of genetic diversity; overall, the majority showed some clustering according to site of collection, with short-distance dispersal of propagules by water and transmission between sites presumably by wind. The extensive genetic diversity observed appears mainly due to somatic mutation within colonies, with some contribution by immigration of propagules from elsewhere into established colonies. Accepted: 10 January 1999     

Historical ozone concentrations and flavonoid levels in herbarium specimens of the Antarctic moss Bryum argenteum

Depletion of stratospheric ozone since the mid 1970s has led to significant increases in ultraviolet B (UVB) irradiation over Antarctica. The detrimental effects of UVB on plants are many, but plants produce photoprotective flavonoids that reduce cellular damage. We used herbarium samples of the moss Bryum argenteum collected in Antarctica to compare the levels of flavone aglycones in plants collected before and after the formation of the ozone hole. The interpretation of historical data is difficult, because environmental conditions immediately before sample collection are unknown. Factors such as cloud cover can have a significant influence on UVB dose at ground level, modifying the flavonoid content of the specimen and adding considerable variability to the results. Nevertheless, our results revealed significant relationships between total flavone concentration and pooled year classes ( P = 0.001). Furthermore, regression analysis showed a significant negative relationship of total flavone concentration and the level of ozone immediately before the time of collection ( P = 0.016). In addition, the ratio of luteolin (an ortho -dihydroxylated flavone) to apigenin (a monohydroxylated flavone) increased significantly with several environmental parameters. These included (a) increasing modelled midday UVB radiation ( P = 0.002), (b) increasing modelled midday UVB/PAR ratio ( P < 0.001), and (c) decreasing ozone concentration ( P < 0.001). We emphasise the utility of this ratio in interpreting the historical ozone trends rather than relying on changes in total flavone concentrations alone. These results illustrate that herbarium specimens may reveal historical levels of UVB radiation.     

Selective sweeps and intercontinental migration in the cosmopolitan moss Ceratodon purpureus (Hedw.) Brid

The moss Ceratodon purpureus has long been used as a model system in plant development and physiology. However, the molecular population genetics of the species remains virtually unexplored. In this study, we used population genetic analyses of DNA sequence data from three unlinked loci (atpB-rbcL spacer, adk, and phy2) to examine biogeographical patterns in a global sample of this species. The three loci differed significantly in mutation frequency spectra and implied population structure. Pairs of haplotypes from single populations were frequently more divergent than haplotypes sampled from widely disjunct populations. In the atpB-rbcL spacer and adk samples, Australasian haplotypes were more closely related to Northern Hemisphere haplotypes than to haplotypes found in the equatorial regions. In contrast, the phy2 sample showed that the north and south temperate regions were genetically divergent, with the equatorial regions intermediate. Maximum-likelihood estimates (MLE) of the rates of migration between the two hemispheres were significantly different for the two nuclear genes. The frequency spectra of mutations indicated that differences in implied population structure among the three loci resulted from directional selection on the chloroplast genome and on the chromosomal segment containing adk. Collectively, these data suggest that long-distance migration within the Northern Hemisphere and Australasian regions is common (relative to the mutation rate) and that migration between these two regions, potentially via equatorial populations, is more frequent than migration among equatorial populations.     

A new microfungal isolate, Embellisia sp., associated with the Antarctic moss Bryum argenteum

A microfungal isolate of Embellisia sp. (Simmons), assigned Embellisia sp2, not previously described from the Antarctic, has been identified by morphological means and internal transcribed spacer (ITS) region sequencing. Embellisia sp2 was shown to be associated with the bryophyte Bryum argenteum, collected from Marble Point in Southern Victoria Land, Antarctica, and grew only in samples plated from crushed moss tissue and surface-sterilized leafy stems. Two other types of microfungi, Penicillium sp. and Trichoderma sp., were cultivated from a surface rinse of the moss. Accepted: 6 May 2000     

沿海岛屿与内陆藓类植物遗传多样性和分化的比较研究——以真藓(Bryum argenteum Hedw.)为例

基于11个内陆居群和32个岛屿居群的252份标本,采用ISSR分子标记技术对真藓( Bryum argenteum Hedw.)的遗传多样性进行了研究。结果显示:岛屿与内陆居群间的遗传多样性差异显著,岛屿居群间的分化程度( G st = 0. 453)大于内陆居群( G st = 0. 387),岛屿居群的遗传分化与地理来源间存在极显著相关性( r = 0. 478, n = 175, P < 0. 001)。地理隔离效应是导致岛屿居群间遗传分化的重要因素。岛内居群间的遗传分化水平较低,仅有29. 4%~ 29. 7%的遗传多样性存在于居群间。聚类分析表明,43个居群可划分为10大类群,真藓遗传关系受地理因素和生境异质性的影响,水域隔离影响了真藓繁殖体在岛屿间的传播。     

沿海岛屿与内陆藓类植物遗传多样性和分化的比较研究——以真藓(Bryum argenteum Hedw.)为例

基于11个内陆居群和32个岛屿居群的252份标本,采用ISSR分子标记技术对真藓（Bryum argenteum Hedw.）的遗传多样性进行了研究。结果显示：岛屿与内陆居群间的遗传多样性差异显著,岛屿居群间的分化程度（G_st=0.453）大于内陆居群（G_st=0.387）,岛屿居群的遗传分化与地理来源间存在极显著相关性（r=0.478,n=175,P<0.001）。地理隔离效应是导致岛屿居群间遗传分化的重要因素。岛内居群间的遗传分化水平较低,仅有29.4%~29.7%的遗传多样性存在于居群间。聚类分析表明,43个居群可划分为10大类群,真藓遗传关系受地理因素和生境异质性的影响,水域隔离影响了真藓繁殖体在岛屿间的传播。     

A Study on the Spore Morphology of Bryum Hedw. in China

Spore morphology of thirteen species of the genus Bryum Hedw. were observed by LM and SEM. The results show that the ornamentation of spore exine could be divided into three types: Type I , blunt at the top of baculate processes, to which four species belong: Bryum argenteum, B. lonchocaulon , B. uliginosum and B. arcticum. Type Ⅱ, sharp or with small processes at the top of baculate processes, represented by seven species: B. pallescens, B. caespiticium, B. pallens, B. pseudotriquetrum, B. paradoxum, B. alpinum and B. thomsoii. Type Ⅲ, expanded into hemispherical-shaped at the top of baculate processes, represented by two species: B.coronatum and B.sauteri. The Bryum species may also be divided into three groups according to the variation of spore diameter. Group I , with spore diameter under 10 μm, including one species, B. uliginosum. Group Ⅱ, with spore diameter 11～20 μm, including seven species: B. argenteum, B. alpinum, B. coronatum, B. pallens, B.paradoxum, B.sauteri, B.thomsonii. Group Ⅲ, spore diameter 21～30 μm, with five species: B. pallescens, B. caespiticium, B. pseudotriquetrum, B. lonchocaulon, B. arcticum. There are resemblances of spore morphology and exine ornamentation among the thirteen species. In the view of palynology, the genus Bryum is a natural taxon which is more advanced than the genus Pohlia Hedw. But spores of thirteen species are different at some characters such as diameter, shape ofproximal leptoma, etc., which indicates the genetic differentiation in the genus Bryum.     

Multiple colonization and dispersal events hide the early origin and induce a lack of genetic structure of the moss Bryum argenteum in Antarctica

The dispersal routes of taxa with transoceanic disjunctions remain poorly understood, with the potential roles of Antarctica not yet demonstrated. Mosses are suitable organisms to test direct intra‐Antarctic dispersal, as major component of the extant Antarctic flora, with the cosmopolitan moss Bryum argenteum as ideal target species. We analyzed the genetic structure of B. argenteum to provide an evolutionary time frame for its radiation and shed light into its historical biogeography in the Antarctic region. We tested two alternative scenarios: (a) intra‐Antarctic panmixia and (b) intra‐Antarctic genetic differentiation. Furthermore, we tested for evidence of the existence of specific intra‐Antarctic dispersal routes. Sixty‐seven new samples (40 collected in Antarctica) were sequenced for ITS nrDNA and rps4 cpDNA regions, and phylogenetic trees of B. argenteum were constructed, with a focus on its Southern Hemisphere. Combining our new nrDNA dataset with previously published datasets, we estimated time‐calibrated phylogenies based on two different substitution rates (derived from angiosperms and bryophytes) along with ancestral area estimations. Minimum spanning network and pairwise genetic distances were also calculated. B. argenteum was potentially distributed across Africa and Antarctica soon after its origin. Its earliest intra‐Antarctic dispersal and diversification occurred during a warming period in the Pliocene. On the same timescale, a radiation took place involving a dispersal event from Antarctica to the sub‐Antarctic islands. A more recent event of dispersal and diversification within Antarctica occurred during a warm period in the Pleistocene, creating favorable conditions also for its colonization outside the Antarctic continent worldwide. We provide evidence supporting the hypothesis that contemporary populations of B. argenteum in Antarctica integrate a history of both multiple long‐range dispersal events and local persistence combined with in situ diversification. Our data support the hypothesis that B. argenteum has been characterized by strong connectivity within Antarctica, suggesting the existence of intra‐Antarctic dispersal routes.