Fe Limitation Research Articles

Automatic fragility curve evaluation of masonry churches accounting for partial collapses by means of 3D FE homogenized limit analysis

A novel 3D homogenized FE limit analysis software is proposed for the fragility curve evaluation of entire existing masonry churches. The approach is a two-step one. In the first phase homogenized masonry strength domains are obtained using a simplified kinematic procedure applied on a three-leaf unitary cell. In the second phase, homogenized domains are implemented at a structural level on a plate and cell kinematic FE software. The procedure is tested on two real scale existing churches located in Italy and compared to results provided by means of a conventional elastic approach performed by means of a standard commercial software. The global behavior at failure and the overall strength of the buildings are assessed through Monte Carlo simulations varying both masonry mechanical properties and direction of the equivalent seismic load.

Phenazine-1-Carboxylic Acid Promotes Bacterial Biofilm Development via Ferrous Iron Acquisition

The opportunistic pathogen Pseudomonas aeruginosa forms biofilms, which render it more resistant to antimicrobial agents. Levels of iron in excess of what is required for planktonic growth have been shown to promote biofilm formation, and therapies that interfere with ferric iron [Fe(III)] uptake combined with antibiotics may help treat P. aeruginosa infections. However, use of these therapies presumes that iron is in the Fe(III) state in the context of infection. Here we report the ability of phenazine-1-carboxylic acid (PCA), a common phenazine made by all phenazine-producing pseudomonads, to help P. aeruginosa alleviate Fe(III) limitation by reducing Fe(III) to ferrous iron [Fe(II)]. In the presence of PCA, a P. aeruginosa mutant lacking the ability to produce the siderophores pyoverdine and pyochelin can still develop into a biofilm. As has been previously reported (P. K. Singh, M. R. Parsek, E. P. Greenberg, and M. J. Welsh, Nature 417:552-555, 2002), biofilm formation by the wild type is blocked by subinhibitory concentrations of the Fe(III)-binding innate-immunity protein conalbumin, but here we show that this blockage can be rescued by PCA. FeoB, an Fe(II) uptake protein, is required for PCA to enable this rescue. Unlike PCA, the phenazine pyocyanin (PYO) can facilitate biofilm formation via an iron-independent pathway. While siderophore-mediated Fe(III) uptake is undoubtedly important at early stages of infection, these results suggest that at later stages of infection, PCA present in infected tissues may shift the redox equilibrium between Fe(III) and Fe(II), thereby making iron more bioavailable.

Whole‐genome expression analysis reveals a role for death‐related genes in stress acclimation of the diatom Thalassiosira pseudonana

Low iron (Fe) availability critically limits diatom distribution and productivity in vast regions of the modern ocean, such as open-ocean, high nutrient low chlorophyll areas and coastal regimes characterized as Fe limitation 'mosaics'. While unique strategies of Fe uptake and storage confer competitive advantages to pennate diatoms, the molecular determinants of low Fe acclimation are largely unknown in centric diatoms. We combined genome-wide and targeted comparative transcriptomic analysis with diagnostic biochemistry and in vivo cell staining as a platform to identify the suite of genes involved in acclimation to Fe and associated oxidative stress in Thalassiosira pseudonana. A total of 1312 genes, nearly 12% of the total genome content, responded to Fe starvation in growing cells characterized by low photosynthetic efficiency and enhanced oxidative stress, caspase activity and metacaspase expression. While 82% of the most highly upregulated genes were also represented in EST libraries derived from diverse diatoms grown under various stress conditions (e.g. silicon, CO(2) and nitrogen limitation), our analysis suggests that T. pseudonana mounts a unique molecular response to Fe starvation that includes a number of genes distinct from those of the model pennate diatom, Phaeodactylum tricornutum, which diverged ~90 million years ago. Homologues to ~50% of the upregulated genes were also identified in a metatranscriptome of eukaryotic phytoplankton communities from a chronically Fe-limited region in the Northeast Pacific. Furthermore, we provide experimental evidence that a subset of putative death-related genes participate in the cellular acclimation to low Fe and associated oxidative damage, suggesting that they co-evolved with other metabolic pathways and play adaptive roles in the success of diatoms.

Simple lower bound limit analysis homogenization model for in- and out-of-plane loaded masonry walls

A simple homogenized equilibrated and admissible model for the limit analysis of masonry walls in- and out-of-plane loaded is presented. A rectangular running bond elementary cell (RVE) is discretized by means of a few constant stress triangular elements (CST) and interfaces. Non linearity is concentrated on brick–brick interfaces and joints reduced to interface, exhibiting a frictional behavior with limited tensile and compressive strength. When dealing with the flexural behavior, a Kirchhoff–Love homogenized model is derived, subdividing the REV into several layers along the thickness and in discretizing each layer analogously to the in-plane case. When dealing with the in-plane behavior, a linear programming problem with very few variables is obtained, which can be handled even manually for some cases of technical interest. The lower bound model proposed is validated at a cell level through literature kinematic models, to show that the lower bound approach provides almost identical results with respect to upper bound approaches, meaning that the actual solution is well approximated. Homogenized masonry behavior is then implemented at a structural level in consolidated FE limit analysis codes for the evaluation of collapse loads and failure mechanisms of real scale masonry structures.

Influence of iron on fatty acid and sterol composition of marine phytoplankton and copepod consumers

We conducted laboratory experiments to determine the influence of iron on the fatty acid (FA) and sterol composition of the diatom Thalassiosira oceanica, the cryptophyte Rhodomonas salina, the prymnesiophyte Isochrysis galbana, and the copepod Acartia tonsa that fed on these algae. Algal cells were cultured with no added Fe or with Fe added at 100 nmol L−1. Fe‐deficient culture medium resulted in lower growth rates, cell volumes, and photosynthetic efficiencies for all algal species. The total mass of FA per cell was 2.2–3.1 times as high in Fe‐replete cells as in Fe‐deficient cells; on a carbon–normalized basis, the FA concentrations of Fe‐replete algal cells were 1.4–1.5 times as high. Differences were noted between Fe treatments for specific saturated, monounsaturated, and polyunsaturated FAs within each algal species. Total C‐normalized 20:5ω3 + 22:6ω3 concentrations declined 18%, 25%, and 35% in Fe‐deficient T. oceanica, R. salina, and I. galbana, respectively. Across all species, another consistent difference in FA composition between Fe treatments was for 18:4ω3, which had a C‐normalized concentration that was ∼ twofold higher in Fe‐replete algae. No consistent pattern for all algal species was evident for Fe effects on algal sterol concentration and composition, and in response to Fe limitation sterols were conserved more than C and FAs in these cells. Copepods feeding on Fe‐replete and Fe‐deficient algae displayed smaller differences in FA and sterol concentrations than in their algal diets. Differences in lipid concentration and composition between Fe‐replete and Fe‐deficient algae may partly explain the lower egg production rate of copepods fed Fe‐deficient algae.

Early Recruitment of AU-Rich Element-Containing mRNAs Determines Their Cytosolic Fate during Iron Deficiency

The yeast Cth2 protein is a CX(8)CX(5)CX(3)H tandem zinc finger protein that binds AU-rich element (ARE)-containing transcripts to enhance their decay in response to iron (Fe) deficiency. Mammalian members of this family of proteins are known to undergo nucleocytoplasmic shuttling, but little is known about the role of shuttling in the mechanism of ARE-dependent mRNA decay. Here we demonstrate that, like its mammalian homologues, Cth2 is a nucleocytoplasmic shuttling protein whose nuclear export depends on mRNA transport to the cytosol. The nuclear import information of Cth2 is contained within its tandem zinc finger domain, but it is independent of mRNA-binding function. Moreover, we also demonstrate that nucleocytoplasmic shuttling of Cth2 requires active transcription and that disruption of shuttling leads to defects in Cth2 function in mRNA decay under Fe deficiency. Taken together, our data suggest that under conditions of Fe deficiency Cth2 travels into the nucleus to recruit target mRNAs, perhaps cotranscriptionally, that are destined for cytosolic degradation as part of the mechanism of adaptation to growth under Fe limitation. These data also suggest an important role for nucleocytoplasmic shuttling in this conserved family of proteins in the mechanism of ARE-mediated mRNA decay.

Quantification of nitrogenase in Trichodesmium IMS 101: implications for iron limitation of nitrogen fixation in the ocean

Iron is widely thought to limit nitrogen fixation in the open, oligotrophic ocean due to the low solubility of Fe in oxic seawater and the high Fe demand for the nitrogenase holozyme. However, empirical evidence for Fe limitation of field populations of Trichodesmium based on either incubation experiments or molecular and physiological indicators has not quantitatively related Fe supply to the cellular Fe quotas for nitrogenase. Rather, the Fe required for N2 fixation has been inferred from in vivo catalytic activity. Using a pet14b expression vector, we cloned the nif H gene (encoding the Fe-protein, which contains 4Fe atoms per subunit) from Trichodesmium IMS 101, and purified the His-tagged apoprotein with which we derived a primary standard based on quantitative Western blots. Using a standard curve derived from the cloned Trichodesmium Fe apoprotein, we measured the absolute abundance of the Fe-protein in iron-replete cultures of this marine diazotroph. At peak expression, we calculate 0.04 mg nitrogenase mg(-1) C. Assuming a conservative stoichiometry of two Fe-protein subunits per MoFe protein (which contains 15 Fe atoms per subunit, or a total of 38 atoms of Fe per holozyme), we estimate 236 µmol Fe is bound to nitrogenase per mol cellular C. This estimate is about 10 times greater than the Fe previously calculated to support diazotrophic growth under these conditions. Our results suggest that under bloom conditions in the subtropical North Atlantic and North Pacific, as much as ∼2.22 and 0.06 µmol m(-3) of Fe is bound to Trichodesmium nitrogenase respectively. Such a high quota represents between ∼50% and > 100% summer-time average particulate Fe in surface waters, suggesting the importance of this taxon for the retention and biogeochemical cycling of Fe. Moderate growth (0.10 day(-1) ) towards the end of these blooms would require a vertical flux as high as ∼23 µmol Fe day(-1) m(-2) into the mixed layer.

Characterization of putative iron responsive genes as species-specific indicators of iron stress in thalassiosiroid diatoms.

Iron (Fe) availability restricts diatom growth and primary production in large areas of the oceans. It is a challenge to assess the bulk Fe nutritional health of natural diatom populations, since species can differ in their physiological and molecular responses to Fe limitation. We assayed expression of selected genes in diatoms from the Thalassiosira genus to assess their potential utility as species-specific molecular markers to indicate Fe status in natural diatom assemblages. In this study, we compared the expression of the photosynthetic genes encoding ferredoxin (a Fe-requiring protein) and flavodoxin (a Fe-free protein) in culture experiments with Fe replete and Fe stressed Thalassiosira pseudonana (CCMP 1335) isolated from coastal waters and Thalassiosira weissflogii (CCMP 1010) isolated from the open ocean. In T. pseudonana, expression of flavodoxin and ferredoxin genes were not sensitive to Fe status but were found to display diel periodicities. In T. weissflogii, expression of flavodoxin was highly responsive to iron levels and was only detectable when cultures were Fe limited. Flavodoxin genes have been duplicated in most diatoms with available genome data and we show that T. pseudonana has lost its copy related to the Fe-responsive copy in T. weissflogii. We also examined the expression of genes for a putative high affinity, copper (Cu)-dependent Fe uptake system in T. pseudonana. Our results indicate that genes encoding putative Cu transporters, a multi-Cu oxidase, and a Fe reductase are not linked to Fe status. The expression of a second putative Fe reductase increased in Fe limited cultures, but this gene was also highly expressed in Fe replete cultures, indicating it may not be a useful marker in the field. Our findings highlight that Fe metabolism may differ among diatoms even within a genus and show a need to validate responses in different species as part of the development pipeline for genetic markers of Fe status in field populations.

The root-hairless barley mutant brb used as model for assessment of role of root hairs in iron accumulation

Main components of Strategy II mechanism for Fe uptake are secretion of chelating compounds, phytosiderophores, and specific uptake of Fe(III)-phytosiderophores complex. Since the amount of phytosiderophores secreted correlates positively with plant ability to cope with Fe shortage, a role of root hairs in enhancing root capability to store phytosiderophores under Fe stress might be envisaged. In this study the root-hairless mutant of barley (Hordeum vulgare L.) brb (bald root barley) and the wild-type genotype (cv. Pallas) were compared with respect to their capacity to respond to Fe shortage in nutrient solution. Plants were grown with Fe(III)-EDTA at 0, 0.02 and 0.08 mM, in order to reproduce severe or moderate Fe deficiency, and adequate Fe nutritional status, respectively. Analysis was performed after 11 and 14 days considering leaf Fe content, phytosiderophores release and accumulation in root tips, and ⁵⁹Fe uptake. Biomass accumulation and chlorophyll content were not reduced in mutant plants as compared to wild-type ones; leaf Fe content was similar in both genotypes after 14 days of growth. Accumulation and release of phytosiderophores showed a similar trend in both genotypes when subjected to Fe limitation. Furthermore, no significant difference between the two genotypes was observed when ⁵⁹Fe uptake was measured. Results seem to support the idea that the presence of root hairs and their increased production in response to low-Fe availability, while causing major modifications of root geometry, did not necessarily lead neither to an effect on growth nor on Fe uptake and accumulation in barley plants.

Ethylene is involved in the regulation of iron homeostasis by regulating the expression of iron-acquisition-related genes in Oryza sativa

Plants employ two distinct strategies to obtain iron (Fe) from the soil. In Strategy I but not Strategy II plants, Fe limitation invokes ethylene production which regulates Fe deficiency responses. Oryza sativa (rice) is the only graminaceous plant described that possesses a Strategy I-like system for iron uptake as well as the classic Strategy II system. Ethylene production of rice roots was significantly increased when grown under Fe-depleted conditions. Moreover, 1-aminocyclopropane-1-carboxylic acid (ACC) treatment, a precursor of ethylene, conferred tolerance to Fe deficiency in rice by increasing internal Fe availability. Gene expression analysis of rice iron-regulated bHLH transcription factor OsIRO2, nicotianamine synthases 1 and 2 (NAS1 and NAS2), yellow-stripe like transporter 15 (YSL15) and iron-regulated transporter (IRT1) indicated that ethylene caused an increase in transcript abundance of both Fe (II) and Fe (III)-phytosiderophore uptake systems. RNA interference of OsIRO2 in transgenic rice showed that ethylene acted via this transcription factor to induce the expression of OsNAS1, OsNAS2, OsYSL15, and OsIRT1. By contrast, in Hordeum vulgare L. (barley), no ethylene production or ethylene-mediated effects of Fe response could be detected. In conclusion, Fe-limiting conditions increased ethylene production and signalling in rice, which is novel in Strategy II plant species.

Dissolved manganese in the Atlantic sector of the Southern Ocean

The first comprehensive dataset (492 samples) of dissolved Mn in the Southern Ocean shows extremely low values of 0.04 up to 0.64 nM in the surface waters and a subsurface maximum with an average concentration of 0.31 nM ( n=20; S.D.=0.08 nM). The low Mn in surface waters correlates well with the nutrients PO 4 and NO 3 and moderately well with Si(OH) 4 and fluorescence. Furthermore, elevated concentrations of Mn in the surface layer coincide with elevated Fe and light transmission and decreased export ( 234Th/ 238U deficiency) and fluorescence. It appears that Mn is a factor of importance in partly explaining the HNLC conditions in the Southern Ocean, in conjunction with significant controls by the combination of Fe limitation and light limitation. No input of Mn from the continental margins was observed. This is ascribed to the protruding continental ice sheet that covers the shelf and shuts down the usual biological production, microbial breakdown and sedimentary geochemical cycling. The low concentrations of Mn in the deep ocean basins (0.07–0.23 nM) were quite uniform, but some elevations were observed. The highest deep concentrations of Mn were observed at the Bouvet Triple Junction region and coincided with high concentrations of Fe and are deemed to be from hydrothermal input. The deep basins on both sides of the ridge were affected by this input. In the deep Weddell Basin the input of Weddell Sea Bottom Water appears to be the source of the slightly elevated concentrations of Mn in this water layer.

COPPER‐UPTAKE KINETICS OF COASTAL AND OCEANIC DIATOMS1

We investigated copper (Cu) acquisition mechanisms and uptake kinetics of the marine diatoms Thalassiosira oceanica Hasle, an oceanic strain, and Thalassiosira pseudonana Hasle et Heimdal, a coastal strain, grown under replete and limiting iron (Fe) and Cu availabilities. The Cu‐uptake kinetics of these two diatoms followed classical Michaelis–Menten kinetics. Biphasic uptake kinetics as a function of Cu concentration were observed, suggesting the presence of both high‐ and low‐affinity Cu‐transport systems. The half‐saturation constants (Km) and the maximum Cu‐uptake rates (Vmax) of the high‐affinity Cu‐transport systems (∼7–350 nM and 1.5–17 zmol · μm−2 · h−1, respectively) were significantly lower than those of the low‐affinity systems (>800 nM and 30–250 zmol · μm−2 · h−1, respectively). The two Cu‐transport systems were controlled differently by low Fe and/or Cu. The high‐affinity Cu‐transport system of both diatoms was down‐regulated under Fe limitation. Under optimal‐Fe and low‐Cu growth conditions, the Km of the high‐affinity transport system of T. oceanica was lower (7.3 nM) than that of T. pseudonana (373 nM), indicating that T. oceanica had a better ability to acquire Cu at subsaturating concentrations. When Fe was sufficient, the low‐affinity Cu‐transport system of T. oceanica saturated at 2,000 nM Cu, while that of T. pseudonana did not saturate, indicating different Cu‐transport regulation by these two diatoms. Using CuEDTA as a model organic complex, our results also suggest that diatoms might be able to access Cu bound within organic Cu complexes.

Two-Step Pushover Analysis of an Ancient Masonry Oil-Mill in the Southern Italy

Historical masonry buildings located in the Southern Italy are usually built with irregular stones joined with mortar with poor mechanical properties. Therefore, piers and spandrels ultimate resistance is not always well predicted by simplified formulas suggested by codes of practice, which typically are tailored to regular patterns. In this framework, we present a two-step numerical model –within the equivalent frame approach assumption– for the pushover analysis of in-plane loaded historical masonry walls constituted by an irregular assemblage of stones. In Step I, ultimate bending moment-shear force strength domains of piers and spandrels are derived by means of a heterogeneous upper bound FE limit analysis and the results are stored in a database. Assessing the capacity of both piers and spandrels is crucial for correctly predicting the ultimate resistance of masonry walls acted upon by in-plane loads. Heterogeneous limit analysis is particularly suitable for computing failure loads, since it permits a distinct modeling of stones and mortar joints. Appropriate static and kinematic boundary conditions are set to account for the complex interaction of internal forces and deformed shapes of single elements. At Step II, a frame model of the masonry wall is assembled, where piers and spandrels are modeled as elastic Timoshenko beams. At each analysis step it is checked that the internal forces in each structural element are smaller than the failure loads stored in the database created at Step I. If the capacity is exceeded, suitable flexural hinges are introduced at the end of the structural elements. The resistance of the element is then set to zero when a limit chord rotation is exceeded. With the numerical tool developed, a real scale old masonry oil-mill located in the Southern Italy is analyzed in the inelastic range under increasing static loads.

Kinematic FE limit analysis homogenization model for masonry walls reinforced with continuous FRP grids

A homogenization model for periodic masonry structures reinforced with continuous FRP grids is presented. Starting from the observation that a continuous grid preserves the periodicity of the internal masonry layer, rigid-plastic homogenization is applied directly on a multi-layer heterogeneous representative element of volume (REV) constituted by bricks, finite thickness mortar joints and external FRP grids. In particular, reinforced masonry homogenized failure surfaces are obtained by means of a compatible identification procedure, where each brick is supposed interacting with its six neighbors by means of finite thickness mortar joints and the FRP grid is applied on the external surfaces of the REV. In the framework of the kinematic theorem of limit analysis, a simple constrained minimization problem is obtained on the unit cell, suitable to estimate – with a very limited computational effort – reinforced masonry homogenized failure surfaces. A FE strategy is adopted at a cell level, modeling joints and bricks with six-noded wedge shaped elements and the FRP grid through rigid infinitely resistant truss elements connected node by node with bricks and mortar. A possible jump of velocities is assumed at the interfaces between contiguous wedge and truss elements, where plastic dissipation occurs. For mortar and bricks interfaces, a frictional behavior with possible limited tensile and compressive strength is assumed, whereas for FRP bars some formulas available in the literature are adopted to reproduce the delamination of the truss from the support. Two meaningful structural examples are considered to show the capabilities of the procedure proposed, namely a reinforced masonry deep beam (0°/90° continuous reinforcement) and a masonry beam in simple flexion for which experimental data are available. Good agreement is found between present model and alternative numerical approaches.

Spatial patterns of nitrogen uptake and phytoplankton in the equatorial upwelling zone (110°W–140°W) during 2004 and 2005

The equatorial Pacific plays a central role in the global carbon cycle as the largest oceanic source of CO 2 to the atmosphere. The region has been described as a HNLC (High Nutrient, Low Chlorophyll) region with low chlorophyll being explained by either Fe limitation, Si(OH) 4 limitation, or a combination of the two. The autotrophic community in the equatorial upwelling zone (EUZ) is dominated numerically by small cell-sized picoplankton but also contains larger cells, including diatoms that require Si(OH) 4 and have been hypothesized to drive NO 3 uptake. The 1-D CoSiNE (Carbon, Silicon, Nitrogen Ecosystem) model was developed for the EUZ to evaluate the potential drivers of the HNLC condition. The model assumes two phytoplankton groups (small non Si(OH) 4 users and larger Si(OH) 4 requiring cells). Many of its predicted variables have not been measured in the field -- specifically, uptake of NO 3, NH 4 and Si(OH) 4 by the larger cells versus the smaller picoplankton and biomass values (as particulate nitrogen, PN) of the two size classes. Two cruises to the EUZ were undertaken in December 2004 (EB04) and September 2005 (EB05) to collect these data, in part to test the conceptual basis and functioning of the CoSiNE model. These EUZ cruises were unique in having size-fractionated uptake rates and phytoplankton community composition at the same depths throughout the euphotic zone and at many stations representing a range of environmental conditions and community composition. Here, we report results pertaining to NO 3 and NH 4 uptake, PN, and f-ratio by different size fractions of the phytoplankton community collected during zonal (0°Eq and 0.5°N) and meridional surveys at 110°W and 140°W. Larger sized (>5 μm) phytoplankton contributed more to NO 3 uptake (up to 85%) than cells <5 μm and so had higher f-ratios. Several autotrophic functional groups were likely responsible for NO 3 uptake in the >5-μm size class, including both diatoms and dinoflagellates. The importance of diatoms in NO 3 uptake appears to be variable, with biological “hotspots” around regions of locally enhanced upwelling associated with tropical instability waves (TIWs). In some cases, TIW activity favored diatoms, and Si(OH) 4 and NO 3 uptake were elevated strongly during TIW activity. Largely as a result of increased diatom productivity, integrated NO 3 uptake rates during the passage of these waves reached 6.74 mmol m −2 d −1, exceeding the mean values previously reported for the equator by >2-fold. NH 4 concentrations and uptake were also dynamic, with lower values south of the equator and higher values to the north. The major contributors to NH 4 uptake were the smaller (<5 μm) phytoplankton that resulted in low total community f-ratios of 0.26 over both years. A >2-fold increase in NH 4 uptake rates in 2005 was consistent with the observed roughly 2-fold increase in Prochlorococcus biomass. Our results highlight a potential area for improvement to the CoSiNE model, to include the role of autotrophic dinoflagellates.

The regulation of equatorial Pacific new production and pCO 2 by silicate-limited diatoms

Modeling and data from the JGOFS EqPac program suggested that the eastern equatorial Pacific upwelling ecosystem includes a quasi-chemostat culture system dominated by diatoms and limited by Si(OH) 4 due to a low ratio of Si(OH) 4 to NO 3 in the upwelling source water, the Equatorial Undercurrent. Diatoms were hypothesized to be the major users of NO 3 in this system and the amount assimilated limited by the low amount of Si(OH) 4 available. As a consequence NO 3 is left in the surface waters along with unused CO 2. Two cruises to the eastern equatorial Pacific (EB04 and EB05) were made to test the existing hypothesis of Si(OH) 4 limitation, and study the roles of source concentrations of Si(OH) 4 and Fe, and nutrient uptake kinetics for comparison with model predictions. Fractionated nitrogen uptake measurements showed that diatoms at times take up the major portion of the NO 3. Picoplankton and some phytoplankton in the >5-μm size group carried out primarily regenerated production, i.e. NH 4 uptake in a grazing dominated system. Equatorial diatoms followed uptake kinetics for Si(OH) 4 and NO 3 uptake as observed in laboratory investigations of diatoms under Si(OH) 4 and Fe limitations. Si(OH) 4 uptake responded to additions of Si(OH) 4 on a time scale of hours in uptake kinetic experiments while NO 3 uptake was unaffected by added NO 3. The uptake of Si(OH) 4 varied in a narrow range on a Michaelis-Menten hyperbola of Si(OH) 4 uptake vs. Si(OH) 4 concentration, with a maximal Si(OH) 4 uptake rate, V′ maxSi set to a relatively low value by some factor(s) other than Fe on a longer time scale, i.e., days in shipboard enclosures. Simply enclosing water collected from the mid euphotic zone and incubating for some days on deck at 50% surface irradiance increased V′ maxSi in accordance with V′ maxSi being a function of incident irradiance. Fe additions to the enclosures also increased V′ maxSi but not to the same extent as only enclosing the water and incubating on deck. The values of V′ maxSi and V′Si showed no relation to ambient Fe concentrations. The study was carried out in a region relatively rich in Fe, from 140°W eastward. These results call into question conclusions that Fe and Si(OH) 4 co-limit production based upon enclosure experiments amended with Fe and incubated at near surface irradiance without first considering what causes the initial large increase in V′ maxSi in the on-deck control enclosures. Some evidence for an Fe effect was seen at the eastern end of the EB04 equatorial section, where Fe concentration generally declined in the eastward direction and at about 118°W reached a low level that may have resulted in the reduction of the V′ maxSi. Data from the EB04 and EB05 cruises showed a close correlation between surface TCO 2 and NO 3 concentration as expected from the demonstrated limitation of diatom NO 3 uptake by Si(OH) 4, highlighting the important role of equatorial diatoms in the global carbon cycle.

Co-limitation of diatoms by iron and silicic acid in the equatorial Pacific

The relative roles of silicon (Si) and iron (Fe) as limiting nutrients in the eastern equatorial Pacific (EEP) were examined in a series of nine microcosm experiments conducted over two years between 110°W and 140°W longitude. Si and Fe additions had consistently different but synergistic effects on macronutrient use, phytoplankton biomass and phytoplankton community structure. Silicon addition increased silicic acid use and biogenic silica production, but had no significant effect on the use of inorganic nitrogen or orthophosphate, chlorophyll accumulation, particulate inorganic (PIC) carbon accumulation, or plankton community composition relative to controls. That result, together with observations that Si addition increased the cellular Si content of the numerically dominant diatom by ∼50%, indicates that the main effect of Si was to regulate diatom silicification. Like the effect of Si, Fe addition increased the rate of silicic acid use and biogenic silica production and had no effect on PIC production. Unlike the effect of Si, Fe addition also enhanced rates of organic matter production, had no effect on cellular Si content of diatoms, and resulted in the growth of initially rare, large (>40 μm) diatoms relative to controls, indicating that Fe limitation acts mainly through its effects on growth rate and phytoplankton community composition. A pennate diatom of the genus Pseudo-nitzschia dominated the diatom assemblage in situ, grew readily in the controls and did not show a strong growth response to either Fe or Si addition suggesting that its growth was regulated by other factors such as grazing or light. Addition of germanium, an inhibitor of diatom cell division, eliminated the effects of Fe on macronutrient use, biogenic silica production and chlorophyll accumulation and phytoplankton community composition, consistent with a predominantly diatom response to Fe addition. The lack of a response of PIC production to Fe suggests that coccolithophores were not Fe limited. Addition of Fe and Si together resulted in the greatest levels of nutrient drawdown and biomass accumulation through the effect of Fe in promoting the growth of large diatoms. The results suggest a form of co-limitation with Si regulating diatom silicification and the rate of biogenic silica production while Fe regulates the production of organic matter through limitation of phytoplankton growth rates, in particular those of large diatoms. The results argue against Si regulation of new production in the EEP under average upwelling conditions. Iron addition was necessary and sufficient to stimulate complete removal of nitrate within the equatorial upwelling zone suggesting that new production was restricted by low ambient dissolved Fe consistent with results from in situ Fe fertilization experiments conducted to the south of the equator outside of the equatorial upwelling zone.

SIGNIFICANCE OF NON APPLIED ELEMENTS IN COCONUT PRODUCTIVITY

A critical evaluation of nutritional factors involved in coconut productivitywas carried out during 1991-1994. West Coast Tall palms aged 25to 35 years varying in yield from 6 to 163 nuts per year growing in threelocations viz. Regional Agricultural Research Station, Pilicode, AgriculturalResearch Station, Mannuthy and Coconut Research Station, Balaramapuramunder regular management constituted the experimental material. The relationshipof individual nutrient contents in respect of 10 elements were workedout. First order nutrient ratios in respect of these elements were also studied.The results showed that continuous use of conventional fertilizers have leadto a stage of negative response for them. It also indicated that deficiency ofnon recommended and non applied elements like Mg and S and excess ofCa, Fe and Mn limits coconut yield. This points to importance of balancednutrition for problem management and better productivity in coconut.

Monte Carlo homogenized limit analysis model for randomly assembled blocks in-plane loaded

A simple rigid-plastic homogenization model for the limit analysis of masonry walls in-plane loaded and constituted by the random assemblage of blocks with variable dimensions is proposed. In the model, blocks constituting a masonry wall are supposed infinitely resistant with a Gaussian distribution of height and length, whereas joints are reduced to interfaces with frictional behavior and limited tensile and compressive strength. Block by block, a representative element of volume (REV) is considered, constituted by a central block interconnected with its neighbors by means of rigid-plastic interfaces. The model is characterized by a few material parameters, is numerically inexpensive and very stable. A sub-class of elementary deformation modes is a-priori chosen in the REV, mimicking typical failures due to joints cracking and crushing. Masonry strength domains are obtained equating the power dissipated in the heterogeneous model with the power dissipated by a fictitious homogeneous macroscopic plate. Due to the inexpensiveness of the approach proposed, Monte Carlo simulations can be repeated on the REV in order to have a stochastic estimation of in-plane masonry strength at different orientations of the bed joints with respect to external loads accounting for the geometrical statistical variability of blocks dimensions. Two cases are discussed, the former consisting on full stochastic REV assemblages (obtained considering a random variability of both blocks height an length) and the latter assuming the presence of a horizontal alignment along bed joints, i.e. allowing blocks height variability only row by row. The case of deterministic blocks height (quasi-periodic texture) can be obtained as a subclass of this latter case. Masonry homogenized failure surfaces are finally implemented in an upper bound FE limit analysis code for the analysis at collapse of entire walls in-plane loaded. Two cases of engineering practice, consisting on the prediction of the failure load of a deep beam and a shear wall arranged with random texture are critically discussed. In particular, homogenization results are compared with those provided by a heterogeneous approach. Good agreement is found both on the failure mechanism and on the distribution of the collapse load.

Effects of an iron-light co-limitation on the elemental composition (Si, C, N) of the marine diatoms &amp;lt;I&amp;gt;Thalassiosira oceanica&amp;lt;/I&amp;gt; and &amp;lt;I&amp;gt;Ditylum brightwellii&amp;lt;/I&amp;gt;

Abstract. We examined the effect of iron (Fe) and Fe-light (Fe-L) co-limitation on cellular silica (BSi), carbon (C) and nitrogen (N) in two marine diatoms, the small oceanic diatom Thalassiosira oceanica and the large coastal species Ditylum brightwellii. We showed that C and N per cell tend to decrease with increasing Fe limitation (i.e. decreasing growth rate), both under high light (HL) and low light (LL). We observed an increase (T. oceanica, LL), no change (T. oceanica, HL) and a decrease (D. brightwellii, HL and LL) in BSi per cell with increasing degree of limitation. The comparison with literature data showed that the trend in C and N per cell for other Fe limited diatoms was similar to ours. Interspecific differences in C and N quotas of Fe limited diatoms observed in the literature seem thus to be mostly due to variations in cell volume. On the contrary, there was no global trend in BSi per cell or per cell volume, which suggests that other interspecific differences than Fe-induced variations in cell volume influence the degree of silicification. The relative variations in C:N, Si:C and Si:N versus the relative variation in specific growth rate (i.e. μ:μmax) followed the same patterns for T. oceanica and D. brightwellii, whatever the irradiance level. However, the variations of C:N under Fe limitation reported in the literature for other diatoms are contrasted, which may thus be more related to growth conditions than to interspecific differences. As observed in other studies, Si:C and Si:N ratios increased by more than 2-fold between 100% and 40% of μmax. Under more severe limitation (HL and LL), we observed for the first time a decrease in these ratios. These results may have important biogeochemical implications on the understanding and the modelling of the oceanic biogeochemical cycles, e.g. carbon and silica export.