Differential responses ofAzolla microphylla Kaulf andAzolla filiculoides Lam. to lead nitrate

The aquatic fernAzolla is used as a green manure for rice production systems in warm temperate and tropical regions throughout the world. We used lead nitrate in nutrient media to assess the tolerance/response of twoAzolla microphylla andAzolla filiculoides to heavy metals. While both species showed negative responses in growth with an increase in lead concentrations, They were distinctly different in their tolerance for higher concentration of the heavy metal. This was apparent in their growth and biochemical characteristics. Amicrophylla was more tolerant of the higher concentrations of lead nitrate (25 and 50 ppm), whereasA. filiculoides had a higher rate of lead uptake.     

Differential response of antioxidant enzymes to salinity stress in two varieties of Azolla (Azolla pinnata and Azolla filiculoides)

The activity and modulation of antioxidant components were comparatively analyzed in two varieties of Azolla (Azolla pinnata and Azolla filiculoides) under different concentrations of NaCl. The total superoxide dismutase (SOD) and ascorbate peroxidase (APX) activity increased in A. pinnata, whereas both enzyme activities decreased in A. filiculoides. The plants of A. pinnata exposed to 30 mM NaCl contained a lower amount of Na⁺ ions and showed a lower electrolyte leakage than A. filiculoides. Our studies indicate a differential response of antioxidant enzymes in relation to salt tolerance in Azolla plants. On the basis of our comparative analysis, A. pinnata has been ranked salt tolerant as compared with A. filiculoides, which is salt sensitive.     

Effects of sodium dodecyl sulphate on the aquatic macrophytes Azolla and Lemna

Abstract

In this work, we determined the tolerance to sodium dodecyl sulphate (SDS) of Azolla filiculoides and of Lemna minor. The presence of the detergent in the media affected growth more than the chlorophyll content. On the basis of the effect indices, Lemna is more tolerant to SDS than Azolla. The fern had a better capacity, compared with duckweed, to remove and accumulate the pollutant.     

Taxonomic re-evaluation of the Azolla genus in Portugal

ABSTRACT

The Portuguese and the European Flora refer to the presence of two or three Azolla species in Portugal: A. filiculoides Lam., A. caroliniana Willd. and/or Azolla mexicana Presl., the latter included in the last edition of Flora Europaea. In the present work, the taxonomy of Azolla species is reviewed using the two most important characters that can distinguish between these two/three species: papillae in the dorsal leaf lobe and perine architecture of the megaspore apparatus. Other characteristics, such as the hyaline border cells of the dorsal leaf lobes and the number of glochidia septa in microsporangium massulae, are also used. All the Azolla specimens, collected from several locations in Portugal, were identified as Azolla filiculoides Lam. This identification disagrees with previous published reports on Azolla taxonomy in Portugal as well as with herbarium identification.     

Interacting effects of atmospheric CO2 enrichment and solar radiation on growth of the aquatic fern Azolla filiculoides

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Use of a natural aquatic fern, Azolla microphylla, as a main component in food for the omnivorous–phytoplanktonophagous tilapia, Oreochromis niloticus L.

An aquatic fern, Azolla microphylla (strain 175 MI, Catholic University of Louvain, Belgium), a natural source of protein, was used in this study to produce low‐cost feeds for the omnivorous–phytoplanktonophagous tilapia, Oreochromis niloticus L. Fish were grown in a recirculating system and fed with six different diets in triplicate groups. Diets were formulated with approximately similar total protein, ranging from 27.25 to 27.52% dry weight (dw), gross energy content ranging from 85.1 to 96.5 MJ kg^?1 dw, and with different levels of dry meal Azolla (0, 15, 20 30 40, 45% diet dw). All diet levels with incorporated Azolla meal exhibited weight gain, thus it can be assumed that Azolla in good combination with local products can be used to promote fish culture development. The Azolla‐free diet and the diet containing 15%Azolla produced the same growth performance. However, the least expensive diet containing 45%Azolla also exhibited growth and can be used as a complementary diet for tilapia raised in fertilized ponds.     

Fructose supports glycogen accumulation,heterocysts differentiation,N2 fixation and growth of the isolated cyanobiont Anabaena azollae

Cells of the cyanobiont Anabaena azollae isolated from the water fern Azolla filiculoides were found to take up and utilize fructose in the light for mixotrophic growth. Fructose was favored by the cyanobiont as a substrate over sucrose and glucose. Cell growth in the presence of 8 mM fructose led to glycogen accumulation in the cells which approached 20% of the cell dry weight within 2 to 3 days, followed by reduction of glycogen content during the fourth day. Glucose-6-phosphate dehydrogenase activity was increased 5–6-fold in the fructose grown cells from the third day of growth onwards. The frequency of heterocysts in fructose-grown cells increased from 6 to 18%, and acetylene reduction by nitrogenase was increased 3-fold in the presence of fructose as compared with control cells, with maximum values observed between the third and fifth day of mixotrophic growth. Fructose-supported growth yielded a 2–4-fold increase in cell dry weight over controls.It is suggested that fructose-supported development and growth of the cyanobiont in batch cultures may resemble its mixotrophic growth and development in situ in the leaf cavity of the host fern Azolla.Abbreviation G6PDH glucose-6-phosphate dehydrogenase     

Adaptation of exotic Azolla to tropical environment of Thailand

There are seven known species of Azolla, two of which have been used in cultivated systems, the tropical speciesA. pinnata, and the temperate speciesA. filiculoides. OnlyA. pinnata is indigenous in Thailand. In this study the two exotic species,A. caroliniana andA. microphylla, were evaluated under the various tropical field conditions in Thailand. When compared with seven selected strains ofA. pinnata under three field conditions,A. caroliniana andA. microphylla were similar to the indigenous species, in terms of growth performance, N₂ fixation, and yield. This study suggests thatA. caroliniana andA. microphylla can be successfully cultivated as a nitrogen fixing green manure for rice production in the tropics.     

Sub-cellular distribution of nitrogen compounds inAzolla andAnabaena by ESI and EELS analysis

Summary The sub-cellular localization of some nitrogen compounds within the leaf cavities ofAzolla filiculoides Lam. was obtained by means of electron spectroscopic imaging (ESI) and electron energy loss spectroscopy (EELS). The analyses were performed on ultrathin unstained sections of differentAzolla leaf cavities which contain epidermal hairs,Anabaena azollae Strasb. and bacteria. Net nitrogen distributions were visualized by image analysis, and nitrogen peaks were evidenced in spectra recorded in the same areas. Different distributions of nitrogen compounds were observed within the leaf cavities along the stem, in particular inside the epidermal hairs ofAzolla and the vegetative cells and heterocysts ofA. azollae.     

A comparative study of nitrogen fixation by the Anabaena-Azolla symbiosis and free-living populations of Anabaena spp. in Lake Ngahawa,New Zealand

Summary The symbiotic fern Azolla filiculoides var. rubra, which contains a blue-green nitrogen fixing alga Anabaena azollae, fixed 164 Kg N·ha^-1·ann^-1 in the littoral zone of a small eutrophic lake. Associated planktonic Anabaena spp. blooms, dominated by Anabaena spiroides, fixed 29.5Kg N·ha^-1·ann^-1. Nitrogen fixation in both organisms was not obviously related to ambient dissolved inorganic nitrogen levels. By comparing ¹⁵N–N₂ and acetylene reduction techniques, we determined a ratio of 3 moles C₂H₂ reduced to 1 mole of N₂ fixed. Combining this with results from one diurnal investigation, it was estimated that 24% of the total daily fixation by Azolla occurred at night. Highest nitrogen fixation rates in Azolla occurred when plant density was lowest. Nitrogen fixation by planktonic Anabaena spp. generally paralleled changes in biomass. Frond breakage due to wind caused a decrease in Azolla nitrogen fixation and growth which was followed by a bloom of planktonic Anabaena spp. A second Anabaena spp. bloom was instrumental in the summer decline of Azolla. Maximum growth and nitrogen fixation of both organisms did not occur simultaneously. If physical disruption to the Azolla mat does not occur, it is likely that growth of the population would continue throughout the year.This work was completed at the Department of Scientific and Industrial Research, Freshwater Section, PO Box 415, Taupo, New Zealand, with partial assistance of N.S.F. Grant BMS-74-20745 to C.R. Goldman     

A Cobalt Requirement for Symbiotic Growth of Azolla filiculoides in the Absence of Combined Nitrogen

Cobalt was shown to be essential for the symbiotic growth of Azolla filiculoides and Anabacna azollac in the absence of fixed nitrogen. Addition of 0.01 μg/liter cobalt resulted in large increases in yield, chlorophyll content and nitrogen fixation as compared to control cultures without cobalt. Cobalt was not required for the growth of Azolla when nitrate nitrogen was supplied. The number of Anabaena azollae cells in the fronds of Azolla appeared to be decreased by ommission of cobalt from the culture medium containing nitrate nitrogen. It is concluded that cobalt is essential for the symbiotic growth of Azolla in the absence of combined nitrogen and it is suggested that the cobalt requirement is associated with the growth of Anabaena azollae.     

Biotechnological potential of Azolla filiculoides for biosorption of Cs and Sr: Application of micro-PIXE for measurement of biosorption

The presence of Cs and Sr in culture medium of Azolla filiculoides caused about 27.4% and 46.3% inhibition of biomass growth, respectively, in comparison to A. filiculoides control weight which had not metals. Biosorption batch experiments were conducted to determine the Cs and Sr binding ability of native biomass and chemically modified biosorbents derived from Azolla namely ferrocyanide Azolla sorbents type 1 and type 2 (FAS1 and FAS2) and hydrogen peroxide Azolla sorbent (HAS). The best Cs and Sr removal results were obtained when A. filiculoides was treated by 2 M MgCl₂ and 30 ml H₂O₂ 8 mM at pH 7 for 12 h and it was then washed by NaOH solution at pH 10.5 for 6 h. Pretreatment of Azolla have been suggested to modify the surface characteristics which could improve biosorption process. The binding of Cs and Sr on the cell wall of Azolla was studied with micro-PIXE and FT-IR.     

High Concentrations of Ammonium Ions at Low pH Decrease the Number of Cyanobionts in Apical Portions of Azolla

When Azolla plants (Azolla filiculoides vax.japonica) were grownin media with high concentrations of ammonium ions at pH 3.5,numbers of cyanobionts in apical portions were very much reduced.Microscopic observations revealed that cyanobionts in theseplants were preferentially damaged, with resultant death andlysis. (Received August 2, 1991; Accepted December 27, 1991)     

Biomass and Pigment Production in Three Isolates of Azolla I. Response to Water Stress

Two isolates of Azolla Lam. from Colombia, South America, andone from the USA (Florida) were selected for characterizationof their growth potentials in certain defined stress environments.These isolates were Amazonian (RAR) and Atlantic coastal (M-3)specimens of Azolla caroliniana Willd and a Floridian variety(LA) of Azolla filiculoides Lam. Osmotic stress (from polyethylene glycol-amended media) andosmotic/salt stress (from seawater-amended media) were the chosendeterminants in these trials. Their effects on growth responseswere monitored. Initial growth rates of LA and M-3 were stimulatedat 1 and 5 ppt seawater salts, relative to growth under controlconditions. RAR, however, was generally the most stable isolatein the experiments with seawater-induced ionic stress. LA wasmost stable in its biomass growth in experiments with polyethyleneglycol 6000, but chlorophyll accumulation data were again leastaffected in RAR. LA survived the longest of the three isolatesat a water potential of approximately —10 bars with eitherseawater (13 ppt salts) or polyethylene glycol (26 per cent).The latter amendment was usually more inhibitory to acetylenereduction rates in RAR and consistently more inhibitory in LA.RAR was tentatively judged by its reactions to water stressto be the superior isolate of the three. Azolla caroliniana Willd., Azolla filiculoides Lam., biomass and chlorophyll accumulation, acetylene reduction, osmotic/salt stress     

Improving nitrogen-fixing systems and integrating them into sustainable rice farming

This paper summarizes recent achievements in exploiting new biological nitrogen fixation (BNF) systems in rice fields, improving their management, and integrating them into rice farming systems. The inoculation of cyanobacteria has been long recommended, but its effect is erratic and unpredictable. Azolla has a long history of use as a green manure, but a number of biological constraints limited its use in tropical Asia. To overcome these constraints, the Azolla-Anabaena system as well as the growing methods were improved. Hybrids between A. microphylla and A. filiculoides (male) produced higher annual biomass than either parent. When Anabaena from high temperature-tolerant A. microphylla was transferred to Anabaena-free A. filiculoides, A. filiculoides became tolerant of high temperature. Azolla can have multiple purposes in addition to being a N source. An integrated Azolla-fish-rice system developed in Fujian, China, could increase farmers' income, reduce expenses, and increase ecological stability. A study using Azolla labeled with ¹⁵N showed the reduction of N losses by fish uptake of N. The Azolla mat could also reduce losses of urea N by lowering floodwater-pH and storing a part of applied N in Azolla. Agronomically useful aquatic legumes have been explored within Sesbania and Aeschynomene. S. rostrata can accumulate more than 100kg N ha^-1 in 45 d. Its N₂ fixation by stem nodules is more tolerant of mineral N than that by root nodules, but the flowering of S. rostrata is sensitive to photoperiod. Aquatic legumes can be used in rainfed rice fields as N scavengers and N₂ fixers. The general principle of integrated uses of BNF in rice-farming systems is shown.     

Changes in elemental content in fronds of Azolla filiculoides due to arsenic accumulation

This study analyzed the accumulation of arsenic and its influence on the content of P, Mn, Fe, Cu, Zn, S, Ca, Cl, and K in fronds of Azolla filiculoides via X-ray fluorescence (XRF). Azolla was exposed to increased concentrations of sodium arsenate (0, 5, 10, 20, and 30 μg ml ^? 1) in Yoshida's nutrient solution. After 96 h, fronds were collected and taken to constant weight to calculate the relative growth rate (RGR), the biomass duplication rate, the bioaccumulation factor (BAF), and the elemental concentration via XRF analysis. By increasing As concentrations the RGR and BAF were reduced. The greatest accumulation of As in fronds was achieved at 20 μg As ml ^? 1, which did not show significant differences with the highest concentration. Accumulation of As in fronds diminished concentrations of Mn, Fe, Zn, S, Ca, and K, while the concentration of P remained low but unchanging. The P content was significantly lower than the As content, thus the As:P ratio in fronds enhanced as the As concentration increased. Concentrations of Cl and Cu were unaffected due to As. This is one of the first reports about the influence of As-accumulation on the elemental content in Azolla fronds.     

DNA Probes Show Genetic Variation in Cyanobacterial Symbionts of the Azolla Fern and a Closer Relationship to Free-Living Nostoc Strains than to Free-Living Anabaena Strains

Twenty-two isolates of Anabaena azollae derived from seven Azolla species from various geographic and ecological sources were characterized by DNA-DNA hybridization. Cloned DNA fragments derived from the genomic sequences of three different A. azollae isolates were used to detect restriction fragment length polymorphism among all symbiotic anabaenas. DNA clones were radiolabeled and hybridized against southern blot transfers of genomic DNAs of different isolates of A. azollae digested with restriction endonucleases. Eight DNA probes were selected to identify the Anabaena strains tested. Two were strain specific and hybridized only to A. azollae strains isolated from Azolla microphylla or Azolla caroliniana. One DNA probe was section specific (hybridized only to anabaenas isolated from Azolla ferns representing the section Euazolla), and five other probes gave finer discrimination among anabaenas representing various ecotypes of Azolla species. These cloned genomic DNA probes identified 11 different genotypes of A. azollae isolates. These included three endosymbiotic genotypes within Azolla filiculoides species and two genotypes within both A. caroliniana and Azolla pinnata endosymbionts. Although we were not able to discriminate among anabaenas extracted from different ecotypes of Azolla nilotica, Azolla mexicina, Azolla rubra and Azolla microphylla species, each of the endosymbionts was easily identified as a unique genotype. When total DNA isolated from free-living Anabaena sp. strain PCC7120 was screened, none of the genomic DNA probes gave detectable positive hybridization. Total DNA of Nostoc cycas PCC7422 hybridized with six of eight genomic DNA fragments. These data imply that the dominant symbiotic organism in association with Azolla spp. is more closely related to Nostoc spp. than to free-living Anabaena spp.     

Studies of the Azolla pinnata—Anabaena azollae symbiosis: Concurrent growth of Azolla with rice

The growth of Azolla pinnata R. Br. under a rice crop was monitored throughout the growing season. Growth rate and N₂-fixation of Azolla decreased markedly after a 4–6 week culture period, a consequence of the low nutrient level in the floodwater rather than shading from the rice plants. The study also showed that Azolla growing concurrently with rice, increased grain yield significantly. The total N released by the local strain of Azolla pinnata, under 60% moisture and flooded soil conditions, was 1 mg g^?1 fresh weight of Azolla.     

Germination characteristics and sporeling success of Azolla filiculoides Lamarck, an aquatic invasive fern, in a Mediterranean temporary wetland

Spore germination characteristics of Azolla filiculoides Lamarck, a recent invader of the Doñana Mediterranean wetlands, have been quantified experimentally. Batches of spores were subjected to different storage conditions, salinity concentrations, phosphorus availability and drought effects. Cold pre-treatment did not stimulate spore germination, whereas drought, a characteristic different from Azolla's native habitat, reduced it (40 ± 7 vs 13 ± 5). Germination declined with the increase of salinity (r = −0.50, notably above 5000 μS cm⁻¹), whereas phosphorus availability had no significant effect.     

Evaluation of some common aquatic macrophytes cultivated in enriched water as possible source of protein and biogas

The biomass production of three common aquatic macrophytes,viz. Azolla pinnata, Eichhornia crassipes andHydrilla verticillata, was high at the prevailing environmental conditions and by the enriched water of River Ganga. The biomass production ofAzolla andEichhornia was positively correlated with the orthophosphate phosphorus and nitrate-nitrogen concentrations of the enriched water. The biomass ofAzolla andHydrilla was positively correlated with the electrical conductivity of the water. The average yield of crude protein was highest in Azolla (8,520 kg.ha^–1.yr^–1), and somewhat lower inEichhornia (6,520 kg.ha^–1.yr^–1). The annual biogas production was highest inEichhornia (44,381 litres), and somewhat lower inAzolla (17,186 litres).