Davallia (Polypodiales: Davalliaceae) macrofossils from Early Miocene Otago (New Zealand) with in situ spores

Fossil fern fronds referable to the extant fern genus Davallia (Polypodiales: Davalliaceae) bearing sporangia with in situ spores are described from the Early Miocene Foulden Maar diatomite deposit, Otago, New Zealand. The fronds are the first published Southern Hemisphere macrofossil record for the family and provide valuable palaeoclimate data supporting warm conditions in Early Miocene New Zealand. The matching of Davallia fronds to the form spore taxon Polypodiisporites radiatus shows that the genus has had a long, apparently continuous history throughout late Cenozoic New Zealand.     

Early Cretaceous Gleicheniaceae and Matoniaceae (Gleicheniales) from Alexander Island,Antarctica

The first Cretaceous representatives of Matoniaceae and Gleicheniaceae in the Antarctic are described from the Albian flora of Alexander Island. Two new species are recognized, one is assigned to Matonia, and another to the emended gleicheniaceous form-genus Gleicheniaceaephyllum. The taxonomy of fossil genera assigned to Gleicheniaceae is problematic, and as an alternative to the invalid genus Gleichenites, we propose that Microphyllopteris be utilised as a form-genus for species that have either an unknown branching pattern or pseudo-dichotomous branching without resting buds; Microphyllopteris is reserved for ferns that cannot be ascribed conclusively to Gleicheniaceae. Alternatively, Gleicheniaceaephyllum should be used for gleicheniaceous material that has a resting bud amongst two to four primary branches. Gleicheniaceaephyllum acutum sp. nov. is preserved as sterile and fertile fronds, pinnae and pinnules. The other new species, Matonia jeffersonii sp. nov., occurs as sterile and fertile pinnae and pinnules, and is aligned to Matoniaceae based on the presence of circular-oval sori with peltate indusia. The two taxa are inferred to have either a pedate or scrambling habit. These morphologies are relatively common at Alexander Island, compared to other high latitude fossil floras, and are interpreted as an adaptation to the high disturbance, polar conditions and unusual photoperiod experienced at Alexander Island during the Early Cretaceous. The theory that angiosperms caused the demise of ferns is discussed in relation to the Matoniaceae and Gleicheniaceae, and there does not appear to be association between angiosperm expansion and the decline of these two fern families.     

2-deoxy-3-epiecdysone from the fern Blechnum vulcanicum

2-Deoxy-3-epiecdysone and ecdysone were isolated from fronds of the fern Blechnum vulcanicum. The structure and stereochemistry of the epiecdystero     

The photosynthetic performance of sterile and fertile sporophytes in a natural population of the fern Dryopteris affinis

We studied the photosynthetic performance of sterile and fertile sporophytes in a natural population of the fern Dryopteris affinis growing within a riparian forest (Central Italy) using chlorophyll (Chl) a fluorescence transients, the OJIP phase, where O is for the minimum fluorescence, P is for the peak (the maximum), and J and I are inflections. The “vitality” of the samples was assessed by the maximum quantum yield of primary photochemistry obtained indirectly from the fluorescence data (F_v/F_m); in the same way, the so-called performance index (PI_ABS) was obtained from fluorescence data. The photosynthetic performance (inferred from PI_ABS) of D. affinis changed significantly with the seasonal development of the fronds. The highest photosynthetic performance was recorded in the summer, corresponding to the period of spore release. The photosynthetic performance decreased in the winter, down to the minimal values of senescent fronds reached at the end of the seasonal cycle (May-June). On the whole, during the seasonal development, sterile and fertile fronds had a similar photosynthetic behaviour, as inferred from fluorescence data. At the end of spore maturation and dispersal (September-October), the fertile fronds showed somewhat lower photosynthetic performance than the sterile fronds, as revealed by PI_ABS. Being a long-lived fern, confined to humid and undisturbed sites in the Mediterranean, D. affinis deserves to be further investigated as a potential indicator of ecological continuity in Mediterranean riparian forests.     

Twin shoots borne on a Botryopteris frond from the British upper carboniferous

A long rachis of Bottyopteris aff. hirsuta bearing twin opposite lateral shoots is described from a coal ball of Westphalian age collected in Lancashire. The simultaneous emission, in the middle region of a frond, of two short axes each bearing a large number of poorly developed leaves is a feature hitherto unrecorded for the genus. The specimen suggests that this Paleozoic fern consisted of long fronds whose branching pattern showed a certain capacity for morphological variation; it provides yet another example of the surprisingly limited development of shoots borne on Botryopteris fronds.     

Interactive effects of selenium and arsenic on their uptake by Pteris vittata L. under hydroponic conditions

The interactive effects of selenium (Se) and arsenic (As) on plant uptake of Se and As have rarely been documented. In this study, the interactive effects of As and Se on their uptake by Chinese brake fern (Pteris vittata), an As-hyperaccumulator and Se-accumulator, were explored in two hydroponic experiments based on a two-factor, five-level central composite design. At Se levels of less than 2.5 mg L^?1, increasing amounts of As stimulated the uptake of Se in Chinese brake fern roots, possibly because of the beneficial effects of Se. In contrast, at Se concentrations greater than 2.5 mg L^?1, As suppressed the uptake of Se in Chinese brake fern roots. Uptake of As by both fronds and roots of Chinese brake fern was suppressed by the addition of Se, indicating the antagonistic effects of Se on As. In addition, at Se concentrations of less than 2.5 mg L^?1, As stimulated the translocation of Se from roots to fronds; meanwhile, the addition of Se resulted in reduced translocation of As from roots to fronds. These findings demonstrate the interactive effects of As and Se on their uptake by Chinese brake fern.     

Kaempferol 3-sulphate in the fern Adiantum capillus-veneris

A new natural product isolated from the fronds of the fern Adiantum capillus-veneris has been shown to be kaempferol 3-sulphate by chemical and spectroscopic methods.     

Sulphate esters of hydroxycinnamic acid—sugar derivatives from Adiantum capillus-veneris

Two new and two known sulphate esters of hydroxycinnamic acid—sugar derivatives have been isolated from the fronds of the fern Adiantum capillus-veneris. The new compounds have been shown to be 1-p-coumarylglucose 2-sulphate and 1-caffeylgalactose 6-sulphate by chemical and spectroscopic methods.     

Volatile Emission in Bracken Fern Is Induced by Jasmonates but Not by Spodoptera littoralis or Strongylogaster multifasciata Herbivory

Jasmonate-mediated regulation of VOC emission has been extensively investigated in higher plants, however, only little is known about VOC production and its regulation in ferns. Here, we investigate whether the emission of VOCs from bracken fern Pteridium aquilinum is triggered by herbivory and if so - whether it is regulated by the octadecanoid signaling pathway. Interestingly, feeding of both generalist (Spodoptera littoralis) and specialist (Strongylogaster multifasciata) herbivores as well as application of singular and continuous mechanical wounding of fronds induced only very low levels of VOC emission. In contrast, treatment with jasmonic acid (JA) led to the emission of a blend of VOCs that was mainly comprised of terpenoids. Likewise, treatment with the JA precursor 12-oxo-phytodienoic acid (OPDA) and α-linolenic acid also induced VOC emission, albeit to a lower intesity than the JA treatment. Accumulation of endogenous JA was low in mechanically wounded fronds and these levels were unaffected by the application of oral secretions from both generalist or specialist herbivores. The emission of terpenoids upon JA treatment could be blocked with fosmidomycin and mevinolin, which are inhibitors of the MEP- and MVA pathways, respectively. These results indicate that similar to higher plants, terpenoid VOCs are produced via these pathways in bracken fern and that these pathways are JA-responsive. However, the very low amounts of terpenoids released after herbivory or mechanical damage are in stark contrast to what is known from higher plants. We speculate that S. multifasciata and S. littoralis feeding apparently did not induce the threshold levels of JA required for activating the MEP and MVA pathways and the subsequent volatile emission in bracken fern.     

Seasonal and interspecific variation in the biochemical composition of some British fern species and their effects on Spodoptera littoralis larvae

Fern-feeding insects in Britain are mainly found on mature fronds in late summer. Six fern species (Dtyopteris filix-mas, D. dilatata, D. borreri, Phyllitis scolopendrium, Polyslkhum setiferum and Polypodium vulgare) were analysed for fibre, lignin, cellulose, tannins, cyanogenesis and thiaminase activity in an attempt to determine the biochemical basis for this seasonal pattern of attack. A bioassay was also carried out, using frond material incorporated in the diet of Spodoptera littoralis, to determine the effects of seasonal changes in fern toxins on a non-adapted, insect herbivore. The young fronds of all six species had a high protein content and low fibre-lignocellulose; protein levels decreased and cell wall materials increased as the fronds matured. Tannin concentrations and thiaminase activity showed a less consistent seasonal pattern. Tannins were present in the highest concentration in the young fronds of D. borreri, Polystichum and Polypodium and decreased with season; D. dilatata, Polystichum and Polypodium showed a similar pattern of thiaminase activity. The other species showed no strong seasonality in the levels of either tannins or thiaminase. No cyanogenic activity was detected in any of the fern species. Frond material from all six species decreased survivorship and growth rates of Spodoptera larvae. Larval performance and growth on diets containing fern material from young immature fronds was generally poorer than on diets containing material from mature fronds, collected later in the season. Neither tannin concentrations nor thiaminase activity levels closely corresponded to the observed interspecific and seasonal patterns of larval development and mortality, and other toxins must be involved. It is concluded that ferns are highly toxic to non-adapted herbivorous insects but the effects on adapted species are unknown. The restricted seasonal occurrence of a small number of insect species exhibiting specialized feeding habits suggests, however, that biochemical properties of the ferns determine this pattern of attack and limit the number of species which are able to exploit ferns as food resources.     

Root exudates and arsenic accumulation in arsenic hyperaccumulating Pteris vittata and non-hyperaccumulating Nephrolepis exaltata

This study compared the roles of root exudates collected from two fern species, the As hyperaccumulating Chinese Brake fern (Pteris vittata L.) and the As-sensitive Boston fern (Nephrolepis exaltata L.), on As-mobilization of two As minerals (aluminum arsenate and iron arsenate) and a CCA (chromated copper arsenate)-contaminated soil as well as plant As accumulation. Chinese Brake fern exuded 2 times more dissolved organic carbon (DOC) than Boston fern and the difference was more pronounced under As stress. The composition of organic acids in the root exudates for both ferns consisted mainly of phytic acid and oxalic acid. However, Chinese Brake fern produced 0.46 to 1.06 times more phytic acid than Boston fern under As stress, and exuded 3–5 times more oxalic acid than Boston fern in all treatments. Consequently, root exudates from Chinese Brake fern mobilized more As from aluminum arsenate (3–4 times), iron arsenate (4–6 times) and CCA-contaminated soil (6–18 times) than Boston fern. Chinese Brake fern took up more As and translocated more As to the fronds than Boston fern. The molar ratio of P/As in the roots of Chinese Brake fern was greater than in the fronds whereas the reverse was observed in Boston fern. These results suggested that As-mobilization from the soil by the root exudates (enhancing plant uptake), coupled with efficient As translocation to the fronds (keeping a high molar ratio of P/As in the roots), are both important for As hyperaccumulation by Chinese Brake fern.     

Antimony accumulation and antioxidative responses in four fern plants

Antimony (Sb) toxicity and contamination has become a growing concern in recent years. Remediation of Sb contamination using plants may be an effective approach. This study aimed to investigate the potential of antimony (Sb) tolerance and accumulation by plants, as well as to understand the antioxidative responses to Sb. One set of hydroponic trials was set up using four species of fern plants, including Pteris cretica (PCA), Microlepia hancei (MH), Cyrtomium fortunei (CYF) and Cyclosorus dentatus (CYD). Ferns were grown for 2 weeks in nutrient solution containing a medium (5 mg L^?1) and a high (20 mg L^?1) rate of Sb, with no Sb added as the control. The biomass of fern PCA remained constant with Sb addition, whereas the biomass of ferns CYF, MH and CYD at the high Sb rate exposure decreased by 12.5%, 35.0% and 38.3%, respectively as compared with their controls. This suggests a high to low Sb tolerance order for these four fern plants. For all of these fern plants, more Sb was accumulated in the roots than in the fronds. Antimony concentration in the roots at the high rate of Sb addition was recorded, on average, as 358 mg kg^?1 for fern PCA, 224 mg kg^?1 for fern CYF, 124 mg kg^?1 for fern CYD and 123 mg kg^?1 for fern MH. A high rate of addition of Sb increased the contents of malondialdehyde (MDA) by 41.3% and 171.6% for ferns MH and CYD, respectively, as compared with their controls. No changes for MDA contents were observed in ferns PCA and CYF with Sb addition, indicating no lipid peroxidation reaction in these two plants. At a medium rate of Sb addition, the activities of peroxidase, catalase and ascorbate peroxidase in fern PCA were much higher than those in ferns CYF, CYD and MH, demonstrating the important role of these three enzymes in resisting Sb toxicity. The consistency in unchanged biomass, high accumulation of Sb in roots, lower MDA contents, as well as high enzyme production in fronds, indicated that fern PCA was more tolerant to Sb than the other three fern plants. Antioxidative enzymes (peroxidase, catalase and ascorbate peroxidase) might be involved in Sb toxicity resistance of fern PCA.     

Differential Response in the Water Status of Immature and Mature Fronds of the Ostrich Fern (Matteuccia struthiopteris [L.] Todaro) to a Mild Water Stress

Experiments were conducted in growth chambers to examine the effect of a mild water stress (−200 kilopascals polyethylene glycol) on frond elongation and water status of the ostrich fern (Matteuccia struthiopteris [L.] Todaro). Measurements were taken for two days, starting one day after the application of polyethylene glycol. Total water potential in control (well-watered) plants was always significantly higher in immature fronds than in mature fronds. The osmotic potential in mature fronds was always significantly lower (about 800 kilopascals) than in immature fronds in both control and stressed plants. In immature fronds, the stress decreased elongation and total water and pressure potentials, while in mature fronds it increased total water and pressure potentials. The decreases in total and pressure potentials in immature fronds were approximately equal to the increases in mature fronds.     

Effects of Se on the uptake of essential elements in Pteris vittata L.

Selenium has been proven to be an antioxidant in plants at low dosages. To understand better the mechanisms of Se toxicity and benefit to plants, more investigations about effects of Se on the uptake of essential elements in plants would be desirable. In this study, hydroponic (nutrient solution culture) and pot (soil culture) experiments were simultaneously conducted to investigate the effects of Se on the uptake and distribution of essential elements in Pteris vittata. L (Chinese brake fern), an arsenic (As)-hyperaccumulator and a selenium (Se)-accumulator. Chinese brake fern took up much more Se in nutrient solution culture than in soil culture, with the highest Se content recorded as 1,573 mg kg^?1 in the roots, demonstrating remarkable tolerance to Se. In soil culture, Chinese brake fern also accumulated high content of Se, with the highest content measured as 81 mg kg^?1 and 233 mg kg^?1, in the fronds and roots, respectively. In soil culture, the addition of Se suppressed the uptake of most measured elements, including magnesium (Mg), potassium (K), phosphorus (P), iron (Fe), copper (Cu) and zinc (Zn). In nutrient solution culture, when the Se content in the tissues of Chinese brake fern was relatively low, the supplementation of Se suppressed the uptake of most essential elements; however, with the increase of Se content, stimulation effects of Se on the uptake of Ca, Mg, K were observed. An initial decrease followed by a rapid increase of Fe content in the fronds of Chinese brake fern was found with Se addition and tissue Se content increasing in nutrient solution culture, suggesting antagonistic and synergic roles of Se on these elements under low and high Se exposure, respectively. We suggest that Ca, Mg, K may be involved in the tolerance mechanism of Se, and that the regulation of Fe accumulation by Se in the fronds might be partially due to the dual effects of Se on Chinese brake fern.     

Effects of Enhanced Ultraviolet-B (280–320 nm) Radiation on Growth and Photosynthetic Activities in Aquatic Fern Azolla Microphylla Kaulf

The effects of enhanced UV-B radiation on growth and photosynthetic activities were investigated in fronds of the aquatic fern Azolla microphylla Kaulf. The fronds were exposed to UV-B radiation intermittently once in 3 d during 12 d. Biomass and relative growth rate of UV-B treated Azolla plants and the heterocyst frequency of the UV-B treated symbiont decreased resulting in an increase in doubling time over the control. The doubling time was 3.08 d for control and 3.35 d for UV-B irradiated plants. Chl and carotenoid contents per unit fresh mass and photosystem 2 (PS2) activity also decreased under UV-B treatment. Measurements of photosynthetic activity in terms of fluorescence kinetics and PS2 mediated O₂ evolution showed that the aquatic fern Azolla is sensitive to UV-B damage.     

Gibberellin-induced Fertility in the Fern Ceratopteris thalictroides (L.) Brongn

Treatment of the homosporous fern Ceratopteris thalictroides L. sporophyte with gibberellic acid results in fertile fronds appearing 3 to 5 weeks after treatment. The hetero-blastic leaf sequence is markedly reduced, and the entire life cycle can be completed in 3 months on defined medium. Treatment with gibberellic acid also stimulates the accumulation of starch in mature leaves.     

Establishment and Control of Hay-scented Fern: A Native Invasive Species

Hay-scented fern (Dennstaedtia punctilobula (Michx.) Moore) is a native forest understory species that behaves as an invasive plant under certain conditions. Previous work has shown that both increased understory light intensity following overstory thinning and removal of competing plants by herbivores can lead to accelerated growth of hay-scented fern, allowing it to develop dense, nearly monospecific understories that inhibit tree seedling regeneration. To study the relationship between these two factors, we sampled 28 forest stands thinned at different times and subjected to different levels of browsing by white-tailed deer (Odocoileus virginianus), and concluded that more than 15 years of intensive browsing following thinning was necessary for ferns to form closed understory canopies with densities of >90 fronds/m² and canopy heights of 60–80 cm; thinning alone or intensive browsing alone was not sufficient to cause this level of fern invasion. We applied three treatments to dense fern understories to determine the relative importance of the fern canopy and the dense mat of roots, rhizomes, and dead fronds in the inhibition of tree seedling establishment. Results after two years were: (1) complete removal of the organic mat produced a large germination response of woody and herbaceous species; (2) mixing the organic mat into the mineral soil produced an initial germination response but poor seedling survivorship, as the fern canopy regrew to near pretreatment density; (3) repeatedly clipping the ferns for two years without disturbing the organic mat resulted in a lower germination response than the removal treatment, but rapid growth of seedlings.     

Multivariate analysis of elements in chinese brake fern as determined using neutron activation analysis

Pytoremediaton of arsenic (As) contamination using Chinese brake fern (Pteris vittata L.), an As hyperaccumulator has proven potential because of its cost-effectiveness and environmental harmonies. Aiming to investigate the elemental correlation in Chinese brake fern, 20 elements (As, Br, Ca, Ce, Co, Cr, Eu, Fe, Hf, La, Na, Nd, K, Rb, Se, Sm, Sr, Th, Yb and Zn) were measured in the fronds and roots of the fern by neutron activation analysis. The ferns were sampled from two sites with high geogenic As levels: Zimudang (ZMD) and Lanmuchang (LMC) in Guizhou Province, China. Multivariate statistic analysis was performed to explore the interrelationship between these elements, especially between As and other elements. As was found to be positively related to K, Na, La, and Sm in both the roots and the fronds, suggesting that these four elements might operate as synergies to As during uptake and transportation processes. Se was positively related to most of the other cations measured, except in the fronds of the fern at ZMD, where Br replaced Se as positively related to the other cations. The difference of As and Se in correlation with other cationic elements suggested that the two anionic elements play different roles in elemental uptake processes. Our findings of elemental correlation highlight the importance of the anioncation balance in Chinese brake fern.     

Azolla-Anabaena Relationship: VIII. Photosynthetic Characterization of the Association and Individual Partners

Photosynthesis in the Azolla-Anabaena association was characterized with respect to photorespiration, early products of photosynthesis, and action spectra. Photorespiration as evidenced by an O₂ inhibition of photosynthesis and an O₂-dependent CO₂ compensation concentration was found to occur in the association, and endophyte-free fronds, but not in the endophytic Anabaena. Analysis of the early products of photosynthesis indicated that both the fern and cyanobacterium fix CO₂ via the Calvin cycle. The isolated endophytic Anabaena did not release significant amounts of amino acids synthesized from recently fixed carbon. The action spectra for photosynthesis in the Azolla-Anabaena association indicated that the maximum quantum yield is between 650 and 670 nanometers, while in the endophyte the maximum is between 580 and 640 nanometers. Although the endophytic cyanobacterium is photosynthetically competent, any contribution it makes to photosynthesis in the intact association was not apparent in the action spectrum.     

New reconstruction of Weichselia reticulata (Stokes et Webb) Fontaine in Ward emend. Alvin, 1971 based on fertile remains from the middle Albian of Spain

Exceptionally well-preserved specimens from the middle Albian of Spain corresponding to a nearly complete fertile frond and fragments of pinnae containing soral clusters of the tree fern Weichselia reticulata have provided new data about the structure and arrangement of pinnae and their associated fertile reproductive structure. This new material has been compared with the previous studies, and recostructions of this fern and the new data indicate the segregation of vegetative and fertile fronds within the main stipe of this fern.