Effect of Sulfur Nutrition on the Redistribution of Sulfur in Vegetative Soybean Plants

Soybean (Glycine max L.) plants were grown with sulfate at 2 (S2) or 20 [mu]M (S20) and treated with [35S]sulfate between d 36 and 38. Growth was continued with or without 20 [mu]M sulfate (i.e. S2 -> S0, S2 -> S20, etc.). When the leaves of S20 -> S20 plants were 70% expanded, they exported S and 35S label from the soluble fraction, largely as sulfate, to new expanding leaves. However, 35S label in the insoluble fraction was not remobilized. Very little of the 35S label in the soluble fraction of the leaves of S20 -> S0 plants was redistributed; most was incorporated into the insoluble fraction. The low levels of S remobilization from the insoluble fraction were attributed to the high level of N in the nutrient solution (15 mM). Most of the 35S label in S2 plants at d 38 occurred in the soluble fraction of the roots. In S2 -> S0 plants the 35S label was incorporated into the insoluble fraction of the roots, but in S2 -> S20 plants 35S label was rapidly exported to leaves 3 to 6. It was concluded that the soluble fraction of roots contains a small metabolically active pool of S and another larger pool that is in slow equilibrium with the small pool.     

Allocation of S in Generative Growth of Soybean

Soybean plants (Glycine max L. Merr) were grown with 100 [mu]M S and 15 mM N and studied with respect to S allocation during grain development. The grains accounted for 87% of the S taken up after d 42, the balance coming from internal redistribution of S from leaves and pods. Detailed studies of the leaves, pods, and grains associated with leaf axils 6 and 7 showed that sulfate accumulated in the pods as they expanded to 50% of full length, ahead of grain enlargement, but declined to very low levels as grain growth commenced. Conversely, homoglutathione (hGSH), cysteine, and methionine increased. In developing grains, hGSH accounted for 60 to 90% of the soluble-S but sulfate was barely detectable. The data are consistent with a model in which, under S-limiting conditions, the pods act as sinks for sulfate and grain growth initiates the assimilation of sulfate into hGSH in the pods, and then into developing grains, where it is incorporated into grain proteins.     

Variation in cadmium accumulation potential and tissue distribution of cadmium in tobacco

Variation in Cd accumulation between Nicotiana species but not varieties has been observed in seedlings grown in solution culture with moderate-to-low levels of Cd. Nicotiana tabacum has been characterized as a leaf and root accumulator while Nicotiana rustica is shown to be primarily a root accumulator, having about half the leaf Cd per gram dry weight of N. tabacum. This phenotype is retained in the mature N. rustica plant. To characterize these two species which differ in their modes of Cd accumulation, tissue Cd distribution, partitioning of metal in soluble and insoluble fractions and the contribution of soluble Cd-binding proteins (peptides) to total plant Cd was assessed using mature solution cultured plants. Metal accumulation was highest in the most mature leaves and in young roots. The preponderance of young roots in N. rustica may, in part, account for low leaf/high root Cd accumulation in this species. While Cd-binding peptides appear to be a principal form of Cd in leaves and roots of seedlings and these also occur in mature leaves, Cd is equally distributed between soluble (about 80% as Cd-binding peptide) and uncharacterized insoluble forms in mature plant roots.     

Accumulation of Oxalate in Tissues of Sugar-beet, and the Effect of Nitrogen Supply

In field-grown sugar-beet concentration of insoluble oxalatewas low in roots and high (about 12 per cent of ethanol insolublematerial) in leaves, and for a particular leaf the concentrationincreased continuously during its life. Of the insoluble oxalate,15–30 per cent was present as the magnesium salt and theremainder as the calcium salt. Oxalate contents of plants grownin culture solutions with nitrate as nitrogen source were similarto those of plants grown in soil, but when nitrogen was suppliedas ammonium sulphate or ammonium nitrate both soluble and insolubleoxalate were low. Plants grown in soil with regular additionsof ammonium sulphate or ammonium nitrate also had very low concentrationsof soluble oxalate although insoluble oxalate was only slightlylower than with nitrate nitrogen. Disks of root or leaf tissuewashed for several days in distilled water lost insoluble oxalatebut when washed in tap water insoluble oxalate increased morethan twofold. Addition of calcium and nitrate to the distilledwater caused an increase of insoluble oxalate, while additionof potassium caused a decrease. Use of ¹⁴C labelled oxalateand washing experiments showed that oxalate can be metabolizedby tissue disks and so is not necessarily a final product ofmetabolism. The accumulation of oxalate appears to be connectedwith the assimilation of nitrate and the preservation of thecation-anion balance of the plant.     

Properties and localization of the homoglutathione synthetase from Phaseolus coccineus leaves

The synthesis of homoglutathione (hGSH) by several plants of the tribe Phaseoleae is shown to be catalysed by a β-alanine-specific hGSH synthetase, Properties of the enzyme from Phaseolus coccineus L. cv. Preisgewinner were studied, using ammonium sulfate precipitates of primary leaf extracts. The hGSH synthetase showed a broad pH optimum at pH 8–9, an absolute requirement for Mg²⁺, a stimulation by K⁺, and a high affinity for γ-glutamylcysteine [K_m(app.) 73 μ M ]. The enzyme exhibited a high specificity for β-alanine [K_m(app.) 1.34 m M ] compared to glycine [K_m(app.) 98 m M ]. Chloroplasts, isolated from the leaves of Phaseolus coccineus , contained about 17% of the hGSH synthetase activity in the leaf cells.     

Manganese accumulation in leaves of Hakea prostrata (Proteaceae) and the significance of cluster roots for micronutrient uptake as dependent on phosphorus supply

When grown at a low P supply, Hakea prostrata R.Br. (Proteaceae) develops dense clusters of determinate branch roots, termed 'proteoid' or 'cluster' roots and accumulates Mn in its leaves. The aim of this study was to vary the production of cluster roots and assess the relationship between Mn uptake and cluster-root mass. We collected native soil from a location inhabited by H . prostrata and amended this with 'high' and 'low' amounts of insoluble or soluble P. After 14 months, we measured the impact of the treatments on cluster-root development and the [P], [Mn], [Fe], [Zn] and [Cu] in young (expanding) and mature leaves. Dry mass and leaf area increased with increasing P availability in the soil, but growth decreased at the highest soluble [P], which caused symptoms of P toxicity. The [P] in young leaves (1.3–2.7 mg g⁻¹ DM) exceeded that in older leaves (0.28–0.85 mg g⁻¹ DM), except when plants were grown with soluble P (3.2–21 mg g⁻¹ DM). Cluster-root formation was inhibited when leaf [P] increased; [P] in young leaves, rather than that in old leaves, appeared to be the factor that determined the proportion of the root mass invested in cluster roots. Old leaves of all treatments had [Mn] from 90 to 120 µg g⁻¹ DM, except for plants grown at high levels of soluble P, when [Mn] decreased below 30 µg g⁻¹ DM. The [Mn] and [Zn] in old leaves and the [Cu] in young leaves were positively correlated with the fraction of roots invested in cluster roots. These findings support our hypothesis that cluster roots play a significant role in micronutrient acquisition, and also provide an explanation for Mn accumulation in leaves of H . prostrata , and presumably Proteaceae in general.     

Effects of Nitrate Nutrition on Nitrogen Metabolism in Cassava

Two experiments were conducted independently with plants of cassava (Manihot esculenta Crantz) growing in sand with nutrient solutions with four nitrate concentrations (0.5, 3, 6 or 12 mM). In leaves, nitrate-N was undetectable at the low nitrate applications; total-N, ammonium-N, amino acid-N, reduced-N and insoluble-N all increased linearly, while soluble proteins did it curvilinearly, with increasing nitrate supply. In contrast, soluble-N did not respond to N treatments. Total-N and soluble proteins, but not nitrate-N or ammonium-N, were much higher in leaves than in roots. Plants grown under severe N deficiency accumulated ammonium-N and amino acid-N in their roots. Further, plants were exposed to either 3 or 12 mM nitrate-N, and leaf activities of key N-assimilating enzymes were evaluated. Activities of nitrate reductase, glutamine synthetase, glutamate synthase and glutamate dehydrogenase were considerably lower in low nitrate supply than in high one. Despite the low nitrate reductase activity, cassava leaves showed an ability to maintain a large proportion of N in soluble proteins.     

Influence of nitrogen supply and sink strength on changes in leaf nitrogen compounds during senescence in two wheat cultivars

Changes in various nitrogen compounds during senescence of the fourth leaf were studied in two cultivars of spring wheat (Triticum aestivum L.). One of the cultivars (Yecora) was supplied with two N levels; the other (Tauro) was grown with the high N level and pruned above the fourth leaf, whereas the control was left intact. In both cultivars grown with high N supply, net nitrogen export from the fourth leaf did not occur until 35 days after sowing (DAS). Loss of leaf soluble proteins started earlier than that of chlorophylis, and coincided initially with an increase in insoluble protein. In N deficient plants the level of total N, soluble protein, and the activity of nitrate reductase (NRA. EC 1.6.6.1) started to decrease about 5 days earlier, and along with chlorophyll, continued to decrease at a faster rate, than in high N plants. Also, with low N supply, the large subunit (LSU, 58 kDa) of ribulose-1.5-bisphosphate carboxylase/oxygenase (Rubisco, EC 4.1.1.39) decreased in greater proportion than other soluble proteins, while with high N supply the decrease in Rubisco LSU was similar to that of other soluble proteins. Nitrogen deficiency caused a greater decrease in soluble proteins than in insoluble proteins, and NRA relative to soluble proteins. The faster senescing Tauro cultivar had lower levels of most parameters, especially NRA, soluble protein and, after 35 DAS. Rubisco LSU as a proportion of soluble protein. The decrease in sink strength due to shoot pruning did generally not affect the level of the various nitrogenous compounds until 35 DAS; thereafter the levels of most parameters, especially soluble protein, Rubisco LSU and, at late stages of senescence, insoluble protein, were higher in pruned than in control shoots. Thus, shoot pruning slows down senescence. The 56- and 78-kDa polypeptides increased, rather than decreased, with leaf age; the level of these two polypeptides showed a negative relationship with Rubisco LSU (r = -0.933 and r = -0.758, respectively).     

Distribution of Sulfur within Oilseed Rape Leaves in Response to Sulfur Deficiency during Vegetative Growth

The distribution of S to sulfate, glucosinolates, glutathione, and the insoluble fraction within oilseed rape (Brassica napus L.) leaves of different ages was investigated during vegetative growth. The concentrations of glutathione and glucosinolates increased from the oldest to the youngest leaves, whereas the opposite was observed for SO₄²⁻. The concentration of insoluble S was similar among all of the leaves. At sufficient S supply and in the youngest leaves, 2% of total S was allocated to glutathione, 6% to glucosinolates, 50% to the insoluble fraction, and the remainder accumulated as SO₄²⁻. In the middle and oldest leaves, 70% to 90% of total S accumulated as SO₄²⁻, whereas glutathione and glucosinolates together accounted for less than 1% of S. When the S supply was withdrawn (minus S), the concentrations of all S-containing compounds, particularly SO₄²⁻, decreased in the youngest and middle leaves. Neither glucosinolates nor glutathione were major sources of S during S deficiency. Plants grown on nutrient solution containing minus S and low N were less deficient than plants grown on solution containing minus S and high N. The effect of N was explained by differences in growth rate. The different responses of leaves of different ages to S deficiency have to be taken into account for the development of field diagnostic tests to determine whether plants are S deficient.     

Impact of nitrate supply in C and N assimilation in the parasitic plant Striga hermonthica (Del.) Benth (Scrophulariaceae) and its host Sorghum bicolor L

The threshold of tolerance for nitrate of the parasitic weed Striga hermonthica (Del.) Benth and the host plant Sorghum bicolor L. was determined by estimating the impact of increasing nitrate loads on plant growth and various parameters of C and N assimilation. Nitrate supply improved chlorophyll (Chl) content and photosystem II (PSII) photochemistry of infected S. bicolor that, in comparison to S. hermonthica, displayed a low imbalance between C and N assimilation when nitrate was supplied up to 1500 mg N per plant. Indeed, nitrate supplies increased strongly the leaf N:C ratio of the parasite. The higher nitrate load induced strong accumulation of nitrate, nitrite and ammonium, and consequently the death of S. hermonthica. Nevertheless, lower nitrate loads (up to 500 mg N per S. bicolor in this study) promoted leaf expansion, PSII photochemistry and N metabolism of S. hermonthica mature (M) plants, as attested by the significant rise in soluble protein and free amino-acid contents. Following these N supplies, the nitrate tolerance of S. hermonthica was correlated with an increase in PSII activity and a high incorporation of N excess into asparagine. This confirmed the central role of asparagine in the N metabolism of S. hermonthica, although this detoxification pathway was insufficient to limit ammonium accumulation under higher nitrate loads.     

Nitrate Reduction in Solanum tuberosum L.: Development of Nitrate Reductase Activity in Field-grown Plants

Nitrate reductase activity (in vivo method, substrate non-limiting)in unshaded leaves from the top of the canopy has been determinedfor field-grown potato plants over the course of the growingseason. The pattern of change was almost identical for plantsreceiving no added fertilizer and those receiving 24 g N m^–2.Activity increased to a peak at about 90 days after plantingand declined thereafter. On a fresh weight basis activity wasalways higher in fertilized plants. Nitrate reductase activitywas positively and significantly correlated with leaf proteincontent in high N plants (r² = 0.71; P = 0.05), but poorly correlatedwith both the nitrate content of the leaf lamina and the nitrateconcentration in petiole sap. Up until 90 days after planting(mid-July) there appeared to be a positive relationship betweenincreased activity of nitrate reductase and solar radiation.However, results obtained over two seasons showed that the declinein activity after this time was not consistently linked witha fall in the level of solar radiation. Remobilization of reduced-Nand stored nitrate from leaves and stems accompanied this declinein nitrate reductase activity and in the latter part of theseason appeared to account for all of the N gained by growingtubers. In unfertilized plants nitrate-N accounted for 5 per cent orless of total plant N. Fertilized plants contained up to 25per cent nitrate-N. While nitrate availability limited growthin unfertilized plants, sub-optimal rates of nitrate assimilationin fertilized plants, particularly during the early stages ofpost-emergence growth, may contribute to inefficient use ofacquired nitrate. The carbohydrate status of leaf lamina and petiole sap weremodified by N supply. The soluble sugar and starch contentsof low N leaves were higher than in their high N counterparts.By contrast, the concentration of soluble sugars in petiolesap increased to a higher value in high N samples. Althoughsap sugar levels declined in both treatments towards the endof the season, N application delayed this decline for severalweeks. Solanum tuberosum, nitrate reductase, nitrate assimilation, senescence     

Induction of leaf senescence by low nitrogen nutrition in sunflower (Helianthus annuus) plants

Different parameters which vary during the leaf development in sunflower plants grown with nitrate (2 or 20 mM) for a 42‐day period have been determined. The plants grown with 20 mM nitrate (N+) showed greater leaf area and specific leaf mass than the plants grown with 2 mM nitrate (N?). The total chlorophyll content decreased with leaf senescence, like the photosynthetic rate. This decline of photosynthetic activity was greater in plants grown with low nitrogen level (N?), showing more pronounced senescence symptoms than with high nitrogen (N+). In both treatments, soluble sugars increased with aging, while starch content decreased. A significant increase of hexose to sucrose ratio was observed at the beginning of senescence, and this raise was higher in N? plants than in N+ plants. These results show that sugar senescence regulation is dependent on nitrogen, supporting the hypothesis that leaf senescence is regulated by the C/N balance. In N+ and N? plants, ammonium and free amino acid concentrations were high in young leaves and decreased progressively in the senescent leaves. In both treatments, asparagine, and in a lower extent glutamine, increased after senescence start. The drop in the (Glu+Asp)/(Gln+Asn) ratio associated with the leaf development level suggests a greater nitrogen mobilization. Besides, the decline in this ratio occurred earlier and more rapidly in N? plants than in N+ plants, suggesting that the N? remobilization rate correlates with leaf senescence severity. In both N+ and N? plants, an important oxidative stress was generated in vivo during sunflower leaf senescence, as revealed by lipid peroxidation and hydrogen peroxide accumulation. In senescent leaves, the increase in hydrogen peroxide levels occurred in parallel with a decline in the activity of antioxidant enzymes. In N+ plants, the activities of catalase and ascorbate peroxidase (APX) increased to reach their highest values at 28 days, and later decreased during senescence, whereas in N? plants these activities started to decrease earlier, APX after 16 days and catalase after 22 days, suggesting that senescence is accelerated in N‐leaves. It is probable that systemic signals, such as a deficit in amino acids or other metabolites associated with the nitrogen metabolism produced in plants grown with low nitrogen, lead to an early senescence and a higher oxidation state of the cells of these plant leaves.     

Effects of nitrogen deficiency on leaf photosynthetic response of tall fescue to water deficit

Abstract. The objective of the present work was to study the effect of nitrogen deficiency on drought sensitivity of tall fescue plants. The authors compared photosynthetic and stomatal behaviour of plants grown at either high (8 mol m⁻³) or low (0.5 mol m⁻³) nitrogen levels during a drought cycle followed by rehydration. Other processes investigated were stomatal and non-stomatal inhibition of leaf photosynthesis, water use efficiency and leaf rolling. Plants were grown in pots in controlled conditions on expanded clay. A Wescor in situ hygrometer placed on the leaf base outside the assimilation chamber permitted, simultaneously to leaf gas exchange measurements, monitoring of leaf water potential. Drought was imposed by withholding water from the pot. CO₂ uptake and stomatal conductance decreased and leaves started to roll at a lower leaf water potential in the high-N than in the low-N grown plants. Stomatal inhibition of leaf photosynthesis seemed larger in the low-N than in the high-N plants. Water-use efficiency increased more in the high-N than in the low-N grown plants during the drought. The decrease of photosynthesis was largely reversible after rehydration in low-N but not in high-N leaves. The authors suggest that low-N plants avoid water deficit rather than tolerate it.     

Extraordinarily high leaf selenium to sulfur ratios define 'Se-accumulator' plants

BACKGROUND AND AIMS: Selenium (Se) and sulfur (S) exhibit similar chemical properties. In flowering plants (angiosperms) selenate and sulfate are acquired and assimilated by common transport and metabolic pathways. It is hypothesized that most angiosperm species show little or no discrimination in the accumulation of Se and S in leaves when their roots are supplied a mixture of selenate and sulfate, but some, termed Se-accumulator plants, selectively accumulate Se in preference to S under these conditions. METHODS: This paper surveys Se and S accumulation in leaves of 39 angiosperm species, chosen to represent the range of plant Se accumulation phenotypes, grown hydroponically under identical conditions. RESULTS: The data show that, when supplied a mixture of selenate and sulfate: (1) plant species differ in both their leaf Se ([Se](leaf)) and leaf S ([S](leaf)) concentrations; (2) most angiosperms show little discrimination for the accumulation of Se and S in their leaves and, in non-accumulator plants, [Se](leaf) and [S](leaf) are highly correlated; (3) [Se](leaf) in Se-accumulator plants is significantly greater than in other angiosperms, but [S](leaf), although high, is within the range expected for angiosperms in general; and (4) the Se/S quotient in leaves of Se-accumulator plants is significantly higher than in leaves of other angiosperms. CONCLUSION: The traits of extraordinarily high [Se](leaf) and leaf Se/S quotients define the distinct elemental composition of Se-accumulator plants.     

Covariation in leaf and root traits for native and non-native grasses along an altitudinal gradient in New Zealand

Across 30 grassland sites in New Zealand that ranged from native alpine grasslands to low elevation improved pastures, there were consistent patterns of leaf and root traits and significant differences between native and non-native grasses. Plants of high altitude sites have low N concentrations in both their leaves and roots, have thick leaves and roots, yet no differences in tissue density or photosynthetic water use efficiency when compared to plants of low altitude sites. Both the leaves and roots of the low altitude plants were enriched in (15)N relative to the plants of higher altitude, indicating that the low-N set of traits is associated with a more closed N cycle at high altitude. A second independent set of correlations shows that plants of wetter habitats have lower photosynthetic water use efficiency (more negative partial differential (13)C) and lower leaf and root tissue density than the plants of drier sites. For both leaves and roots, plants of native species consistently had traits associated with lower resource availability: lower N concentrations, denser tissues, more negative partial differential (15)N, and more positive partial differential (13)C than non-native species. If root %N is correlated with root longevity as has been shown in other systems, root longevity may be able to be predicted from simple measurements of leaf %N, though a hysteresis in the relationship between leaf and root N concentrations may make prediction of high longevity roots difficult.     

Sulfur deprivation and nitrogen metabolism in maize seedlings

The objective of this experiment was to elucidate the manner in which N metabolism is influenced by S nutrition. Maize (Zea mays L.) seedlings supplied with Hoagland solution minus SO₄²⁻ exhibited S deficiency symptoms 12 days after emergence. Prior to development of these symptoms, a decline in leaf blade nitrate reductase (NR, EC 1.6.6.1) activity was observed in S-deprived seedlings compared to normal seedlings. Twelve days after emergence, in vitro NR activity was diminished 50% compared to normal seedlings. Glutamine synthetase (EC 6.3.1.2) and NAD-glutamate dehydrogenase (EC 1.4.1.2) activities were less severely affected (19 and 13%, respectively, at day 12). NADP-glutamate dehydrogenase (EC 1.4.1.4) activity and leaf blade fresh weight were not altered by S deprivation. Concentrations of soluble protein and chlorophyll (a and b) in leaf blades were reduced 18 and 25%, respectively, at day 12. A significantly higher concentration of NO₃⁻-N was observed for leaf blade and stem (culms, leaf sheaths, and unfurled leaves) fractions (46 and 31%, respectively) in S-deprived plants. In contrast to the other parameters measured, NR activity in S-deprived seedlings could be readily restored to the normal level by addition of SO₄²⁻. The apparent preferential effect of S deprivation on NR activity could be causally related to the observed changes in NO₃⁻-N and soluble protein concentration.     

Interaction of water supply and N in wheat

The purpose of this study was to investigate effects of N nutrition and water stress on stomatal behavior and CO₂ exchange rate in wheat (Triticum aestivum L. cv Olaf). Wheat plants were grown hydroponically with high (100 milligrams per liter) and low (10 milligrams per liter) N. When plants were 38 days old, a 24-day water stress cycle was begun. A gradual increase in nutrient solution osmotic pressure from 0.03 to 1.95 mega Pascals was achieved by incremental additions of PEG-6,000. Plants in both N treatments adjusted osmotically, although leaf water potential was consistently lower and relative water content greater for low N plants in the first half of the stress cycle. Leaf conductance of high N plants appeared greater than that of low N plants at high water potentials, but showed greater sensitivity to reductions in water potential as indicated by earlier stomatal closure during the stress cycle. The apparent greater stomatal sensitivity of high N plants was associated with a curvilinear relationship between leaf conductance and leaf water potential; low N plants exhibited more of a threshold response. Trends in [CO₂]_INT throughout the stress cycle indicated nonstomatal effects of water stress on CO₂ exchange rate were greater in high N plants. Although estimates of [CO₂]_INT were generally lower in high N plants, they were relatively insensitive to leaf water potential-induced changes in leaf conductance. In contrast, [CO₂]_INT of low N plants dropped concomitantly with leaf conductance at low leaf water potentials. Oxygen response of CO₂ exchange rate for both treatments was affected less by reductions in water potential than was CO₂ exchange rate at 2.5% O₂, suggesting that CO₂ assimilation capacity of the leaves was affected more by reductions in leaf water potential than were processes related to photorespiration.     

低N胁迫下巴西橡胶实生苗叶片SPAD值、N素营养及农艺性状的变化

对3个品种(GT1、RRIM600、PR107)巴西橡胶(Hevea brasiliensis)实生苗进行不同N素(0%N,50%N,100%N)处理,分析比较叶片SPAD值、N素营养和农艺性状的变化及其关系。结果表明:(1)缺N胁迫明显降低了叶片SPAD值、叶片N含量和硝酸还原酶活性,缩小了叶蓬距、叶面积,且影响幅度随胁迫强度的加大而增大;(2)不同品种橡胶树实生苗对缺N胁迫的响应程度不同,总体上GT1较为敏感,PR107次之,RRIM600较不敏感;(3)橡胶树实生苗叶片SPAD值、叶蓬距、叶面积与叶片N含量、硝酸还原酶活性之间均呈显著或极显著正相关。因此,有可能采用测定叶片SPAD值、叶蓬距和叶面积等简易方法判断橡胶树新生叶蓬的N素营养状况。     

Distribution and Redistribution of Sulfur Supplied as [35S]Sulfate to Roots during Vegetative Growth of Soybean

Soybean (Glycine max L.) plants were grown in nutrient solution containing 10 [mu]M sulfate and were treated at various times with [35S]sulfate for 48 h. Growth was then continued in unlabeled solution. The sulfur content of each leaf increased rapidly until it was about 40% expanded; small, additional increases occurred until the leaf was about 70% expanded after which the sulfur content decreased by about 50%. Leaves that were about 60 to 70% expanded during the pulse were strongly labeled but then underwent a significant loss of 35S label. Leaves that were in the early stages of expansion imported little 35S label during the pulse but acquired 35S label during the chase period as they expanded (i.e. redistribution). Most of the redistributed 35S label was derived from other leaves. The rates of both sulfur import and sulfur export by a leaf were greatest at about 70% expansion. Leaves that acquired 35S label during early development retained a much higher proportion of their label than leaves that were more developed, suggesting that the sulfur acquired by leaves during early development is preferentially incorporated into a pool that is less mobile than the sulfur acquired in the later stages of leaf growth.