Effect of sulphur nutrition on redistribution of sulphur in vegetative barley

Xenobiotics directed against sterol biosynthesis have proved to be useful tools in the determination of which sterol molecules are necessary for successful plant cell growth. However, the exact mode of action by which sterols are able to trigger cell growth remains to be elucidated. Previous studies using the triazole paclobutrazol, demonstrated that in Apium graveolens (cv. New Dwarf White) suspension cultures, sterol and phosphatidylcholine biosynthesis are co-ordinately regulated (C. E. Rolph and L. J. Goad 1991, Physiol. Plant. 83: 605–610). The studies presented herein, were designed to investigate the possible role of phosphatidylcholine in the stimulation of plant cell growth. Sterol biosynthesis, and hence cell growth, was inhibited by the use of the azole xenobiotic miconazole. Treatment of the cultures with miconazole lead to compositional changes in the free sterol content of the cells. For example, 30 μM miconazole treatment led to a reduction in the stigmasterol/sitosterol ratios from 1.53 to 1.24. In contrast, the phospholipid content of the cells remained relatively unchanged with phosphatidylcholine accounting for approximately 25% of the total phospholipids present in both control and miconazole-treated cells. The cytostatic effect of miconazole could be partially counteracted by supplementation of the growth medium with the phytosterol stigmasterol and/or the unsaturated fatty acids oleate and linoleate. The activity of CTP:cholinephosphate cytidylyltransferase (EC 2.7.7.15), a rate-limiting enzyme in phosphatidylcholine biosynthesis, was significantly reduced in cells whose growth had been arrested by miconazole treatment. In miconazole-treated cultures whose growth had been partially restored by supplementation with either specific sterols or unesterified fatty acids, the activity of this key enzyme was increased. In the case of stigmasterol, oleate and linoleate supplementation, the microsomal activity was found to be similar to that exhibited by control cultures. From these studies, it may be concluded that certain phytosterols and unsaturated fatty acids play key roles with respect to phosphatidylcholine biosynthesis and that phosphatidylcholine biosynthesis via the CDP-base pathway is an important pre- and/or co-requisite for successful culture growth.     

The role of soil microbes in plant sulphur nutrition

Chemical and spectroscopic studies have shown that in agricultural soils most of the soil sulphur (>95%) is present as sulphate esters or as carbon-bonded sulphur (sulphonates or amino acid sulphur), rather than inorganic sulphate. Plant sulphur nutrition depends primarily on the uptake of inorganic sulphate. However, recent research has demonstrated that the sulphate ester and sulphonate-pools of soil sulphur are also plant-bioavailable, probably due to interconversion of carbon-bonded sulphur and sulphate ester-sulphur to inorganic sulphate by soil microbes. In addition to this mineralization of bound forms of sulphur, soil microbes are also responsible for the rapid immobilization of sulphate, first to sulphate esters and subsequently to carbon-bound sulphur. The rate of sulphur cycling depends on the microbial community present, and on its metabolic activity, though it is not yet known if specific microbial species or genera control this process. The genes involved in the mobilization of sulphonate- and sulphate ester-sulphur by one common rhizosphere bacterium, Pseudomonas putida, have been investigated. Mutants of this species that are unable to transform sulphate esters show reduced survival in the soil, indicating that sulphate esters are important for bacterial S-nutrition in this environment. P. putida S-313 mutants that cannot metabolize sulphonate-sulphur do not promote the growth of tomato plants as the wild-type strain does, suggesting that the ability to mobilize bound sulphur for plant nutrition is an important role of this species.     

三江源高寒草地牧草营养时空分布

草地的产草量和牧草品质影响着草地载畜量和草食动物的营养状况、生命活动及生产性能。以三江源及其周边地区的天然草地不同生长期（返青期,盛草期,枯黄期）牧草为对象,分析草地产草量及牧草品质的时空格局。结果显示：从整个区域看,草地生产力、可食牧草产量以及粗蛋白产量(Crude Protein, CP)分布均具有较明显的地域差异性,三者的高值区大多分布在三江源东部及南部的高寒草甸区,三者的低值区主要在三江源中部和西部的草甸及高寒草原区;大部分地区总生产力和可食牧草产量盛草期显著高于枯黄期,枯黄期显著高于返青期。粗蛋白产量在大部分地区呈现出盛草期显著高于返青期和枯黄期,返青期和枯黄期差异不明显,少数地区返青期显著高于盛草期和枯黄期,盛草期显著高于枯黄期（如：可可西里）;CP含量及产量在三江源区空间上的分布,均为南部地区高于北部地区,东部农牧交错区高于西部无人区;所有区域CP含量的返青期为最高,枯黄期为最低,而其他养分含量在不同区域的季节波动并不一致;盛草期牧草CP含量随海拔升高先增加后减小,粗灰分随着海拔升高而显著增加,中性洗涤纤维、酸性洗涤纤维和酸性洗涤木质素含量随着海拔升高呈降低趋势。结论：三江源及周边地区牧草生产力、可食牧草产量以及CP产量较高,可食性较好;三江源多数地区高寒草地牧草营养质量相对较好,牧草在返青期营养价值最高（蛋白高,纤维低）,到盛草期,牧草产量及营养物质输出量达到最高峰,进入枯黄期的牧草营养质量低劣,此时应进行补饲,以提高家畜生产性能。三江源区可利用草地主要集中在东部、南部和东南部,该区牧草品质较高、利用潜力（粗蛋白产量）较大,可适度利用开发。     

Evaluation of a soil test method and plant analysis for determing the sulphur status of alluvial soils

Summary In a pot experiment with soils of Alfisol, Entisol, and Inceptisol orders, the relative yield of Egyptian clover (Trifolium alexandrinum L.) was significantly correlated with Morgan's reagent (N NaOAc+HOAc, pH 4.8)—extractable soil S (r=0.88), plant S (r=0.82), and plant N/S ratio (r=−0.77) suggesting suitability of these tests for diagnosing S deficiency. Total plant S lower than 0.21 per cent, plant N/S ratio wider than 17, and extractable soil S lower than 10 ppm were indicative of S deficiency, and were suggested therefore to be critical limits for these tests. Nitrogen and S in plant proteins were in near constant ratio of 16 and were significantly correlated (r=0.99). Sixty one per cent of 250 surface soil samples had less than 10 ppm extractable S and hence were deficient in S, suggesting a widespread S deficiency in soils under study. Extractable soil S in all soil series was significantly correlated with electrical conductivity and alkaline KMnO₄-extractable N, but not with pH, organic C, and CaCO₃.     

Interactions between selenium and sulphur nutrition in Arabidopsis thaliana

Selenium (Se) is an essential plant micronutrient, but is toxic at high tissue concentrations. It is chemically similar to sulphur (S), an essential plant macronutrient. The interactions between Se and S nutrition were investigated in the model plant Arabidopsis thaliana (L.) Heynh. Arabidopsis plants were grown on agar containing a complete mineral complement and various concentrations of selenate and sulphate. The Se/S concentration ratio in the shoot ([Se](shoot)/[S](shoot)) showed a complex dependence on the ratio of selenate to sulphate concentration in the agar ([Se](agar)/[S](agar)). Increasing [S](agar) increased shoot fresh weight (FW) and [S](shoot), but decreased [Se](shoot). Increasing [Se](agar) increased both [Se](shoot) and [S](shoot), but reduced shoot FW. The reduction in shoot FW in the presence of Se was linearly related to the shoot Se/S concentration ratio. These data suggest (i) that Se and S enter Arabidopsis through multiple transport pathways with contrasting sulphate/selenate selectivities, whose activities vary between plants of contrasting nutritional status, (ii) that rhizosphere sulphate inhibits selenate uptake, (iii) that rhizosphere selenate promotes sulphate uptake, possibly by preventing the reduction in the abundance and/or activity of sulphate transporters by sulphate and/or its metabolites, and (iv) that Se toxicity occurs because Se and S compete for a biochemical process, such as assimilation into amino acids of essential proteins.     

Effect of nitrogen nutrition on the export of sulphur from leaves in soybean

Soybean plants were grown in complete solution for 33 days and then transferred to medium containing inadequate sulphur (5 t M) and nitrogen at 15, 7.5, 2 or 0.25 mt M. In mature leaves (L1 and L2), and leaves that were 70% expanded at day 33 (L3), the net loss of sulphur over the ensuing 25 days was inversely related to the level of nitrogen nutrition. Leaf 5, which formed during the study period, exhibited complementary characteristics; the increase in the sulphur content was inversely related to the level of nitrogen nutrition even though low nitrogen nutrition supported less growth. L4, which was 31% expanded at day 33, exhibited intermediate characteristics. 35S-Labelled sulphate was supplied to all of the plants for 48 h at day 31 and was distributed principally to L3 at day 33. During early development, L5 became heavily labelled but, at low nitrogen nutrition, the massive import of total sulphur into L5 during the late stages of development was not accompanied by a commensurate increase in 35S-label, indicating that redistribution of soluble sulphur from mature leaves was not involved. The loss of sulphur from mature leaves was parallelled by similar changes in nitrogen at all levels of nitrogen nutrition. Collectively, the data suggest that a common mechanism, presumably proteolysis, is involved in the export of sulphur and nitrogen from mature leaves and that this process is inhibited at high levels of nitrogen nutrition, even under conditions of sulphur deficiency.     

Influence of plant sulphur nutrition on oviposition and larval performance of the diamondback moth

We measured adult oviposition preference, larval growth, and feeding behaviour of the crucifer specialist Plutella xylostella (L.) (Lepidoptera: Plutellidae) on plants of Brassica napus (L.) cv. Express (Brassicaceae), grown under three different sulphur regimes. The nutrient solutions used were the following: one sulphur‐free (S₀), one normal sulphur (S_n, normal field concentration), and one sulphur‐rich (S₊, double concentration of S_n). Females laid more eggs on S_n than on S₀ plants, while only a slight, non‐significant difference was observed between S_n and S₊ plants. Moreover, the development time from hatching to emergence was significantly shorter, and adults were heavier on S_n than on S₀ plants. Comparing these same two parameters from S_n and S₊ plants, we found a shorter development time on plants rich in sulphur, although this trend was not statistically significant. Larval feeding preferences were tested in a dual choice assay using leaf discs. A significantly higher number of larvae preferred leaf discs of S_n plants than those of S₀ plants. Furthermore, the larvae preferred S₊ to S_n discs. An optimal supply of sulphur to oilseed rape is necessary for a good seed harvest, and it also plays an important role in acceptance by P. xylostella of the host plant. Maintaining higher levels of sulphur in the plant nutrient solution benefits insect performance, both at the adult and larval stage.     

Impact of pedospheric and atmospheric sulphur nutrition on sulphur metabolism of Allium cepa L., a species with a potential sink capacity for secondary sulphur compounds

Onion (Allium cepa L.) was able to use atmospheric H(2)S as sole sulphur source for growth. The foliarly absorbed H(2)S was rapidly metabolized into water-soluble, non-protein thiol compounds, including cysteine, and subsequently into other sulphur compounds in the shoots. In H(2)S-exposed plants, the accumulation of sulphur compounds in the shoots was nearly linear with the concentration (0.15-0.6 microl l(-1)) and duration of the exposure. Exposure of onion to H(2)S for up to 1 week did not affect the sulphur content of the roots. Secondary sulphur compounds formed a sink for the foliarly absorbed sulphide, and the sulphur accumulation upon H(2)S exposure could, for a great part, be ascribed to enhancement of the content of gamma-glutamyl peptides and/or alliins. Furthermore, there was a substantial increase in the sulphate content in the shoots upon H(2)S exposure. The accumulation of sulphate originated both from the pedosphere and from the oxidation of absorbed atmospheric sulphide, and/or from the degradation of accumulated secondary sulphur compounds. From studies on the interaction between atmospheric and pedospheric sulphur nutrition it was evident that H(2)S exposure did not result in a down-regulation of the sulphate uptake by the roots.     

Preferential allocation of sulphur into y‐glutamylcysteinyl peptides in wheat plants grown at low sulphur nutrition in the presence of cadmium

Plants respond to Cd by synthesising phytochelatins (PC) and similar S‐rich peptides which are important in alleviating Cd toxicity. The hypothesis that S nutrition influences the sensitivity of plants to Cd was examined by measuring the growth, PC and Cd content of wheat plants ( Triticum aestivum cv. Condor) grown at 10–1000 µ M S with and without 30 µ M Cd. In the absence of Cd, 100 µ M S was marginally S‐limiting. Cd severely inhibited root growth at 100 µ M S but the concentrations of PC, Cd–PC complexes and Cd in the root were similar to those of plants grown at 1000 µ M S which exhibited no evidence of Cd toxicity. Plants grown at low S (10 and 30 µ M ) contained lower concentrations of Cd. However, they produced very low amounts of PC and were very sensitive to Cd. At 10–300 µ M S (but not at 1000 µ M ), Cd enhanced the concentration of S in the root. The data are consistent with a model in which plants preferentially allocate S to PC synthesis. When S was marginally limiting (100 µ M ), the S supply was sufficient for PC synthesis but not root growth, effectively causing Cd‐induced S deficiency. This did not occur at high S (1000 µ M ). Conversely, at low S (10 and 30 µ M ), the synthesis of PC₂–PC₄ was decreased by 87 and 66%, respectively, thereby resulting in decreased Cd uptake while also making the plants especially sensitive to Cd.     

Influence of sulphur plant nutrition on oviposition and larval performance of the cabbage root fly

1 Oilseed rape plants (Brassica napus) (L.) (Brassicaceae) were grown under different levels of sulphur supply and tested for the oviposition preference and larval performance of cabbage root flies Delia radicum (L.) (Diptera: Anthomyiidae). 2 Adult females laid more than three‐fold as many eggs on control S_n (normal field concentration) than on sulphur‐free S₀ plants. By contrast, no significant difference was observed between control and double normal concentration (S₊) plants. 3 The larval performance was evaluated using three additional, intermediate sulphur levels between S₀ and S_n, and the plants were infected with equal numbers of eggs. The percentage pupation at the end of larval feeding ranged from 6% (S₀) to 32% (S_n or S₊) and the average number of pupae, or of emerging flies, was significantly correlated with sulphur application. 4 The weight of emerging males and females was correlated with plant sulphur supply. 5 The duration of development from eggs to adult emergence was approximately 2 days longer in females than in males. Females originating from plants with a normal or higher sulphur supply tended to emerge 1–2 days earlier.     

Nutritional status of Ribeirinhos in Brazil and the nutrition transition

Anthropometric and household data (size, composition, economic activity) were collected from a population of Ribeirinhos living in a rural setting in the eastern Amazon. Data are compared to international reference standards and to other Amazonian populations with the goals of increasing our understanding of the Amazon's largest ethnic group and identifying the relationship between changes in subsistence strategies and nutritional status. Data on height, weight, skinfolds, and circumferences were collected from 471 adults and subadults. The population showed a high degree of stunting with an average HAZ below -2.0 for all age groups over 3 years, and 60% of adult men and 70% of adult women were stunted. Wasting was rare. Average skinfold thicknesses and upper-arm muscle area were near or below average but within the normal range compared to the reference standard, indicating adequate energy and protein stores. Thirty-one percent of males and 29% of females were overweight/obese, and the highest average BMIs were found among men and women in their 40s. Adult males who participated in wage labor had higher weights, BMIs, and UMA values, and were more likely to be overweight and obese compared with those who did not work in wage-labor jobs. Children of fathers who worked in wage labor had higher BMI and UMA values, but there was no significant effect on the nutritional status of other adults in these same households. Signs of the nutrition transition were most noticeable among adult males involved in wage labor because of changes in their diet and activity patterns.     

A sulphur deficiency-induced gene, sdi1, involved in the utilization of stored sulphate pools under sulphur-limiting conditions has potential as a diagnostic indicator of sulphur nutritional status

A sulphate deficiency-induced gene, sdi1 , has been identified by cDNA-amplified fragment length polymorphism (AFLP) analysis utilizing field-grown, nutrient-deficient wheat ( Triticum aestivum var. Hereward). The expression of sdi1 was specifically induced in leaf and root tissues in response to sulphate deficiency, but was not induced by nitrogen, phosphorus, potassium or magnesium deficiency. Expression was also shown to increase in plant tissues as the external sulphate concentration in hydroponically grown plants was reduced from 1.0 to 0.0 m m . On this basis, sdi1 gene expression has potential as a sensitive indicator of sulphur nutritional status in wheat. Genome-walking techniques were used to clone the 2.7-kb region upstream of sdi1 from genomic DNA, revealing several cis -element motifs previously identified as being associated with sulphur responses in plants. The Arabidopsis thaliana gene most highly homologous to sdi1 is At5g48850, which was also demonstrated to be induced by sulphur deficiency, an observation confirmed by the analysis of microarray data available in the public domain. The expression of Atsdi1 was induced more rapidly than previously characterized sulphur-responsive genes in the period immediately following the transfer of plants to sulphur-deficient medium. Atsdi1 T-DNA 'knockout' mutants were shown to maintain higher tissue sulphate concentrations than wild-type plants under sulphur-limiting conditions, indicating a role in the utilization of stored sulphate under sulphur-deficient conditions. The structural features of the sdi1 gene and its application in the genetic determination of the sulphur nutritional status of wheat crops are discussed.     

Evaluation of the nitrogen nutrition status of rice by plant analysis

Summary The results of a greenhouse and a field experiment to evaluate the nitrogen nutrition status of rice indicate that: (1) first and second most recently natured leaves are sensitive indicators of the nitrogen status of plants; (2) the total nitrogen concentration in these leaves are a better indicator of the nutrient status of plants than are the water-soluble and amide-plus-ammonium nitrogen fractions; and (3) that the best correlation between plant growth and total nitrogen is obtained when these leaves are sampled at the stage of growth between maximum tillering and panicle emergence.     

Effects of sulphur nutrition on growth and nitrogen fixation of pea (Pisum sativum L.)

A S-deficient soil was used in pot experiments to investigate the effects of S addition on growth and N₂-fixation in pea (Pisum sativum L.). Addition of 100 mg S pot⁻¹ increased seed yield by more than 2-fold. Numbers of pods formed were the most sensitive yield component affected by S deficiency. Sulphur addition also increased the concentration of N in leaves and stems, and the total content of N in the shoots. The amounts of N fixed by pea were determined at four growth stages from stem elongation to maturity, using the ¹⁵N dilution technique. Sulphur addition doubled the amount of N fixed at all growth stages. In contrast, leaf chlorophyll content and shoot dry weight were increased significantly by S addition only after the flowering and pod fill stage, respectively. Pea roots were found to have high concentrations of S, reaching approximately 10 mg g⁻¹ dry weight and being 2.6–4.4 times the S concentration in the shoots under S-sufficient conditions. These results suggest that roots/nodules of pea have a high demand for S, and that N₂-fixation is very sensitive to S deficiency. The effects of S deficiency on pea growth were likely to be caused by the shortage of N, due to decreased N₂-fixation. This revised version was published online in June 2006 with corrections to the Cover Date.     

Separation, subcellular location and influence of sulphur nutrition on isoforms of cysteine synthase in spinach

Cysteine synthase (O-acetylserine (thiol) lyase, EC 4.2.99.8) and -cyanoalanine synthase (EC 4.4.1.9) have been isolated from leaves of Spinacea oleracea L. and separated by anion-exchange chromatography. Further separation of one minor and two major isoforms of cysteine synthase was achieved by hydrophobic interaction chromatography and high resolution native electrophoresis (PAGE). Analysis of root material indicated that amongst the multiple isoforms present, a single isoform predominated. Subcellular fractionation studies indicated that one of the major leaf forms, cysteine synthase B, was located in the chloroplast and the other, cysteine synthase B, occurred in the cytoplasm. No specific isoform of cysteine synthase was resolved in the mitochondria, while cyanoalanine synthase was predominantly located in the mitochondrial fraction. Sulphur deprivation decreased cysteine synthase activity, but not cyanalanine synthase activity in both young and mature leaves, although cysteine synthase activity in the roots increased slightly. A selective decrease in cystein synthase B (chloroplastic abundance was observed in mature leaves. Patterns of expression of cysteine synthase in response to S-availability are discussed in relation to possible roles for this enzyme in controlling S-flux through the S-assimilatory pathway.Key words: Cysteine synthase isoforms (expression of), Spinacea oleracea L., sulphur deficiency.