Tumorous crown gall response to iron‐deficiency stress in sunflower

Tumorous crown gall tissue in sunflower (Helianthus annus L.) initiates a mechanism for making Fe available to itself as evidenced by its ability to reduce Fe³⁺ to Fe²⁺. The objective of this study was to determine if a limited Fe supply to the plant might affect the growth, nutrition and reduction of Fe³⁺ to Fe²⁺ by the tumorous crown gall. Healthy green 14‐day‐old sunflower plants (cv mammoth Russian) were either stem‐inoculated with Agrobacterium tumefaciens to induce tumorous crown gall tissue development or were left uninoculated for comparison. The plants were grown in a modified Hoagland nutrient solution with treatments containing 0.0, 0.15, 0.6 and 2.0 mg Fe L^‐1. The 0 mg Fe L^‐1 treatment induced maximum Fe chlorosis, and consequently there was a release of hydrogen ions and of a yellow pigment by the roots, but there was no measureable release of ‘reductants’ by the roots. Iron‐deficiency stress (0 mg Fe L^‐1) also resulted in reduced tumorous crown gall growth, less reduction of Fe³⁺ to Fe²⁺, and lower levels of Fe in the tumorous tissue compared to tumorous tissues adequately supplied with Fe. The tumorous crown gall tissue on the stem reduced much more Fe³⁺ to Fe²⁺ than the nontumorous stem tissue regardless of Fe level in the treatment. Tumor tissue contained more Fe, Cu and P than the nontumorous stem tissues which may indicate a modified metabolism in this tissue. An abundant supply of Fe seems to enhance the development and growth of the tumorous crown gall tissue and a deficient supply of Fe retards its growth.     

Differential response of groundnut genotypes to iron‐deficiency stress

Differential response of groundnut genotypes to iron‐deficiency stress was studied in soils containing high calcium carbonate. Genotypes differed significantly for some traits that appeared to be important in determining adaptation to low iron. The genotypes TCGS 273, TCGS 2, TCGS 37, and Kadiri 3 had higher total chlorophyll, total dry matter, and active iron (Fe²⁺) contents under iron‐deficiency stress conditions. Total chlorophyll followed by active iron were found to be sensitive parameters to Fe deficiency. Based on the visual deficiency symptoms (chlorosis score), the genotypes were classified into three groups. Efficient (no genotype was found efficient), moderately efficient (TCGS 273, TCGS 2, TCGS 3, and Kadiri 3), and inefficient (TCGS 1, TCGS 7, TCGS 11, TCGS 26, TCGS 28, TCGS 29, TCGS 30, TCGS 1518, TPT 1, TPT 2, ICGS 11, ICGS 44, Girnar, JL 24, ICGS(E) 21, and TMV 2).     

An iron chelating compound released by barley roots in response to Fe‐deficiency stress

The excretion of phytosiderophores by barley (Hordeum vulgare L.) has recently been documented and a major difference in the Fe‐stress response of gramineous species and dicotyledonous species proposed. However, currently used methods of quantifying and measuring phytosiderophore are tedious or require specialized equipment and a cultivar easily accessible to U.S. scientists is needed. The objectives of this study were (a) to determine if “Steptoe”; and “Europa”; (used as a control cultivar) barleys would release Fe³⁺ solubilizing compounds in response to Fe‐deficiency stress and (b) to develop a technique to determine the efficiency of solubilization of Fe(OH)₃ by the released chelating substances. Two cultivars of barley were place under Fe‐stressed (‐Fe) and nonstressed (+Fe) conditions in modified Hoagland solutions (14 L). The solutions were periodically monitored for H⁺ and reductant release from the roots and plants were rated daily for chlorosis development. Periodic (6 or 7 harvests) evaluation of the release of Fe³⁺ solubilizing substances was performed as herein described. Neither H⁺ nor reductant extrusion occurred with either cultivar during Fe stress. However, Fe³⁺ solubilizing substances were released by both cultivars at relatively high levels under Fe‐stress conditions compared to the nonstressed plants. A convenient technique was developed to measure the release of Fe solubilizing substances released by barley roots.     

Factors in iron‐stress response mechanism enhanced by Zn‐deficiency stress in Sanilac,but not Saginaw navy bean

Zinc‐inefficient Sanilac and Zn‐efficient Saginaw navy bean (Phaseolus vulgaris L.) differ in their susceptibility to Zn‐deficiency stress. Sanilac accumulates Fe under Zn‐deficiency stress and Saginaw does not. These two navy bean cultivars were grown at 0, 0.006 and 0.12 mg/L Zn in modified Hoagland nutrient solution. Various Fe‐stress response mechanisms were quantified periodically over a 12‐day experimental period to determine if known factors in the Fe‐stress response mechanism were enhanced by Zn‐deficiency stress. Visual Zn‐deficiency symptoms were more severe in Sanilac than Saginaw navy bean under equivalent Zn treatments. Sanilac contained lower leaf Zn than Saginaw when Zn was present in solution (0.006 and 0.12 mg/L Zn), but the two cultivars were similar in leaf Zn in the absence of Zn (0 mg/L Zn). Sanilac accumulated more leaf Fe than Saginaw when under Zn stress (0 and 0.006 mg/L Zn). The higher levels of leaf Fe in Sanilac than Saginaw were closely associated with enhanced release of reductants and increased reduction of Fe³⁺ to Fe²⁺ by roots of Sanilac. Saginaw navy bean roots reduced Fe³⁺ to Fe²⁺ similarly to Sanilac with adequate Zn present in solution (0.12 mg/L), but experienced minuscule levels of Fe³⁺ reduction under Zn deficiency. Zinc deficiency stimulated the initiation of the Fe‐stress response mechanism in Sanilac, but not Saginaw, which may have enhanced the development of Zn‐deficiency symptoms in Sanilac due to the increased uptake of Fe by this cultivar. The common Fe‐deficiency stress response associated primarily with grasses (release of phytosiderophore) was not found in either navy bean cultivar.     

Tumorous crown gall response to iron‐deficiency stress in Fe‐efficient T3238fer and Fe‐inefficeent T3238fer tomatoes

Abstract

Although sunflower (Helianthus annus L.) is an Fe efficient plant, tumorous crown gall tissue development and tissue ability to reduce Fe³⁺ to Fe²⁺ were both diminished by Fe‐deficiency stress. Crown gall also develops readily on Fe‐efficient and Fe‐inefficient tomato cultivars (Lycopersicon esculentum Mill.). The objective of this study was to determine if the effect of a limited Fe supply on the growth, nutrition and reduction of Fe³⁺ to Fe²⁺ by tumorous crown gall would differ between Fe‐efficient T3238FER and Fe‐inefficient T3238fer tomato. Healthy green 25‐day‐old plants were either stem‐inoculated with Agrobacterium tumefaciens to induce tumorous crown gall tissue development or were left uninoculated for comparison. Plants were grown in modified Hoagland nutrient solutions containing 0.0, 0.15, 0.6 and 2.0 mg Fe L^?1. Yield of tumorous crown gall tissue was not diminished by low solution Fe in T3238FER, but was in T3238fer. This was attributed to inability of the T3238fer tomato to make Fe available to itself. Tumor tissue from both cultivars contained more Fe, Cu and P than normal stem tissues, which confirms a modified metabolism in these tissues previously observed in sunflower. An abundant supply of Fe enhances the development and growth of the tumorous crown gall tissue, but a deficient supply of Fe retards its growth.     

Response to iron‐deficiency stress of pear and quince genotypes 1

Roots of iron (Fe)‐efficient dicots react to Fe‐deficiency stress by strongly enhancing the ferric (Fe³⁺)‐reductase system and by lowering the rhizo‐sphere pH. In this study, we tested whether such adaptation mechanisms characterize pear and quince genotypes known to have differential tolerance to calcareous and alkaline soils. Two trials were performed using micropagated plants of three quince rootstocks (BA29, CTS212, and MC), three Pyrus communis rootstocks (OHxF51 and two selections obtained at the Bologna University: A28 and B21) and of two pear cultivars (Abbé Fétel and Bartlett, own‐rooted). In the first trial, plants were grown in a nutrient solution with [Fe(+)] and without [Fe(‐)] Fe for 50 days. Their root Fe‐reducing capacity was determined colorimetrically using ferrozine and FeEDTA, and Fe uptake of Fe(+) plants was estimated. In the second trial, the rhizosphere pH of plants grown in an alkaline soil was measured by a micro‐electrode. With the only exception of pears OHxF51 and A28, whose Fe‐reduction rates were similar in Fe(+) and Fe(‐) plants, the Fe‐deficiency stress resulted in a significant decrease in Fe reduction. Among the Fe(‐) plants, the two pear cultivars, OHxF51 and A28, had a higher Fe‐reducing capacity than the quince rootstocks and the cv. Abb6 F. When plants were pre‐treated with Fe, reduction rate was highest in the P. communis rootstocks, intermediate in the own‐rooted cultivars, and lowest in the quinces. Root Fe‐reducing capacity of Fe(+) plants proved to be linearly and positively correlated with Fe uptake and root proton release. Rhizosphere pH was highest in quince MC, intermediate in the other two quinces and in the cv. Abbe F., and lowest in the pear rootstocks and in the cv. Bartlett. Our results indicate that roots of pear and quinces do not increase their ability to reduce the Fe under Fe‐deficiency stress. The genotypical differential tolerance to Fe chlorosis likely reflects differences in the standard reductase system and in the capacity of lowering the pH at the soil/root interface. The determination of the root Fe‐reducing capacity is a promising screening technique for selecting pear root‐stocks efficient in taking up Fe.     

Monocot‐dicot variability in response to an iron oxide‐metallic iron source

Abstract

Iron from a mixture of Fe oxide and metallic Fe was more available to corn (Zea mays L.) than it was to soybeans when the plants wore grown in calcareous soil or in nutrient solution. All this Fe, however, was DTPA (diethylene triamine pentaacetic acid) extractable. In solution culture the Fe was available to the soybean (Glycine max L.) plants unless CaCO₃ was included in the nutrient solution.     

Influences of ultra‐violet (UV)‐blue light radiation on the growth of cotton. I. Effect on iron nutrition and iron stress response

Cool white fluorescent (CWF) light reduces Fe³⁺ to Fe²⁺ while low pressure sodium (LPS) light does not. Cotton plants grown under CWF light are green, while those yrown under LPS light develop a chlorosis very similar to the chlorosis that develops when the plants are deficient in iron (Fe). It could be that CWF light (which has ultra violet) makes iron more available for plant use by maintaining more Fe²⁺ in the plant. Two of the factors commonly induced by Fe‐stress in dicotyledonous plants‐‐hydroyen ions and reductants released by the roots‐‐were measured as indicators of the Fe‐deficiency stress response mechanism in M8 cotton.

The plants were grown under LPS and CWF light in nutrient solutions containing either NO₃‐N or NH₄‐N as the source of nitrogen, and also in a fertilized alkaline soil. Leaf chlorophyll concentration varied significantly in plants grown under the two light sources as follows: CWF+Fe > LPS+Fe > CWF‐Fe ≥ LPS‐Fe. The leaf nitrate and root Fe concentrations were significantly greater and leaf Fe was generally lower in plants grown under LPS than CWF light. Hydrogen ions were extruded by Fe‐deficiency stressed roots grown under either LPS or CWF light, but “reductants”; were extruded only by the plants grown under CWF light. In tests demonstrating the ability of light to reduce Fe³⁺ to Fe²⁺ in solutions, enough ultra violet penetrated the chlorotic leaf of LPS yrown plants to reduce some Fe³⁺ in a beaker below, but no reduction was evident through a yreen CWF grown leaf.

The chlorosis that developed in these cotton plants appeared to be induced by a response to the source of liyht and not by the fertilizer added. It seems possible that ultra violet liyht could affect the reduction of Fe³⁺ to Fe²⁺ in leaves and thus control the availability of this iron to biological systems requiring iron in the plant.     

Soybean response to the application of several elements on a low‐manganese soil

Abstract

Certain soils in the Lower Atlantic Coastal Plain are Mn‐deficient and because of their inherent properties, deficiencies of other elements might be expected. The response of soybean [Glycine max (L.) Merr. cv. ‘Ransom'] to annual soil applications of B, Cu, Fe, Mo, S, and Zn with and without Mn was examined. The study was conducted on an Olustee‐Leefield sand (Ultic Haplaquod‐Arenic Plinthaquic Paleudult) in 1975, 1976, and 1977. Seed yields, and soil and plant tissue concentrations of certain of the applied elements were determined. Soil pH increased from 6.4 in 1975 to 7.0 in 1977 as a result of lime applications.

Of the seven elements studied, only Mn significantly increased soybean seed yield compared to the check. Leaf concentrations of the respective elements were higher where the element was added than they were in plants grown on the check plots. In 1977, after three years of elemental additions, plant concentrations of Mn, Cu, and Zn were higher, relative to checks, than in 1975. In 1977, with the higher soil pH levels, plants from check plots had lower concentrations of Mn, but unexpectedly equal levels of Cu and higher levels of Zn than in 1975. This unexplained increase in plant Zn was also found in Other experiments in the same field during the same time period. Double acid extracted more soil Mn, Cu, and Zn than did DTPA from similar treatments during all three years. The DTPA‐extractable soil Mn correlated much better with plant Mn than did double acid Mn values, especially over years where a change in soil pH occurred. Correlation coefficients comparing extractable soil versus plant values for Cu and Zn data combined over years were higher for double acid than DTPA, although coefficients for the Cu and Zn data were still generally much lower than for the Mn data.     

Fe‐deficiency tolerance in papaya (Carica papaya L.): Ph reduction and chlorosis recovery in response to stress

The phenomenon of Fe‐stress response mechanism was examined using papaya as a test fruit plant species. This plant behaved like the many Fe‐efficient crop cultivars, reduced the pH of the minus Fe nutrient medium, followed by recovery from chlorosis. The response mechanism was observed in both the 2 cultivars, CO‐1 and CO‐3. Evidence obtained with this plant supports the widespread occurrence of the mechanism in diverse crop species ‐ grain, vegetable and fruit crop plants too.     

Substitutes for benzidine as H‐donors in the peroxidase assay,for rapid diagnosis of iron deficiency in plants

Abstract

3,3'5,5’ Tetramethyl benzidine, 3‐Amino‐9‐ethylcarbazol and chlorpromazine are suggested as alternatives for benzidine as H‐donors in the peroxidase assay for diagnosing iron deficiency in plant leaves. Procedures with these reagents, which have been used successfully with citrus and other plants, are described.     

Phytosiderophore release from manganese‐induced iron deficiency in barley

An experiment was conducted in the phytotron with barley (Hordeum vulgare L. cv. Minorimugi) grown in nutrient solution to compare iron (Fe) deficiency caused by the lack of Fe with manganese (Mn)‐induced Fe deficiency. Dark brown spots on older leaves and stems, and interveinal chlorosis on younger leaves were common symptoms of plants grown in either Mn‐toxic or Fe‐deficient treatments. Dry matter yield was affected similarly by Fe deficiency and Mn toxicity. The Mn toxicity significantly decreased the translocation of Fe from roots to shoots, caused root browning, and inhibited Fe absorption. The rate of Fe translocated from roots to shoots in the 25.0 μM Mn (toxic) treatment was similar to the Fe‐deficient treatment. Manganese toxicity, based on the release of phytosiderophore (PS) from roots, decreased from 25.0>250>2.50 uM Mn. The highest release of PS from roots occurred 7 and 14 days after transplanting (DAT) to Mn‐toxic and Fe‐deficient treatments, respectively; but was always higher in the Fe‐deficient treatment than the Mn‐toxic treatments. The release of PS from roots decreased gradually with plant age and with severity of the Mn toxicity symptoms. The PS content in roots followed the PS release pattern.     

Pattern of iron distribution in the soil‐plant system and its possible relation to iron‐chlorosis

Abstract

The frequent concentration‐ranges of various nutrient elements in soils and in plants are compared. Iron is different from almost all other nutrient elements in the fact that its optimal concentration range in plants is much lower than its frequent concentration range in soils. It is suggested that this observation is related to a chemical‐physiological mechanism of control on the uptake of iron by plants which in turn may explain the situations in which iron deficiency conditions in plants arise.     

Root responses of sterile‐grown onion plants to iron deficiency 1

Onion (Allium sativum) plants grown without iron (Fe) in sterile nutrient solutions readily developed chlorosis symptoms. Iron deficiency in the sterile‐grown plants stimulated the rates of root extracellular reduction of Fe³⁺, copper (Cu²⁺), manganese (Mn⁴⁺), and other artificial electron acceptors. While rapid reduction occurred with the synthetic chelate Fe³⁺HEDTA, no short‐term reduction occurred with the fungal siderophore Fe³⁺ferrioxamine B (FeFOB). In addition to the increased rate of extracellular electron transfer at the root surfaces, the Fe‐deficient plants showed greater rates of Fe uptake and translocation than the onion plants grown with Fe. The rates of uptake and translocation of Fe were sharply higher for the Fe‐deficient plants supplied with FeHEDTA than for similar plants supplied with FeFOB. Inhibition by BPDS of the Fe uptake by the Fe‐deficient onion plants further supported the importance of Fe³⁺ chelate reduction for the uptake of Fe into the roots. Rates of Fe uptake and translocation by Fe‐deficient onion plants supplied with ⁵⁵FeFOB were identical to the rates of uptake of ferrated [14C]‐FOD; a result that gives evidence of the uptake and translocation of the intact ferrated siderophore, presumably by a mechanism not involving prior extracellular Fe³⁺ reduction. Differences in the rates of transport of other micronutrients into the roots of the Fe‐deficient onion plants were evident by the significantly higher Zn and Mn levels in the shoots of the Fe‐deficient onion.     

Manganese‐iron interactions in the plant‐soil system

Dicotyledonous plants had significantly higher Mn and Fe intake rates on a near neutral soil, had a significantly higher Mn intake rate on a slightly calcareous soil, but had lower Mn and Fe intake rates on a calcareous soil, than monocotyledonous plants. This dependency on soil reaction suggests that dicots utilize primarily a chemical reduction mechanism, whereas monocots utilize some less pH‐dependent mechanism (possibly Mn(III)‐, Fe(III)‐organic complexation) to mobilize soil Mn and Fe. Soluble soil Mn and Fe fractions in the rhizosphere were consistently positively correlated with each other, as were Mn and Fe intake rates. These results suggest that for soil‐grown plants, Mn and Fe uptake was positively interrelated because both Mn and Fe were mobilized by similar root processes.     

Iron‐stress response of inoculated and non‐inoculated roots of an iron inefficient soybean cultivar in a split‐root system

The objective of this study was to establish whether the iron‐stress responses observed in T203 soybean (Fe‐inefficient) with active nodules are products of the nodules or of the entire root system. A split‐root system was used in which half the roots of each plant were inoculated and actively fixing nitrogen and the other half were not. Soybean cultivar T203 is normally Fe‐inefficient and does not exhibit the Fe‐stress responses, however an iron‐stress response did occur during active N₂ fixation in earlier studies. This implies that the active nodules influenced the plant's ability to respond to Fe‐deficiency stress. In this study, the Fe‐stress response (H⁺ and reductant release) observed in T203 soybean was limited to the inoculated side of the split‐root system. The severe Fe chlorosis which developed in these plants was overcome in a manner similar to Fe‐efficient cultivars undergoing normal Fe‐stress response and the T203 plants completely regreened. Exudation of H⁺ ions was similar in both the presence and absence of Fe, and was generally limited to inoculated roots. Reductant release was nearly nonexistent from the non‐inoculated roots and was greater for the Fe‐stressed (‐Fe) plants than for non‐stressed (+Fe) plants. Thus, the response observed, which alleviated Fe chlorosis, appeared to be associated with N₂ fixation of the active nodules.     

Increases in phosphatase and β‐glucosidase activities in wheat seedlings in response to phosphorus‐deficient growth

The ability of plants to utilize P efficiently is important for crops growing in P‐deficient soils or on soils with a high P‐fixing capacity. The purpose of this work was to investigate early physiological changes which occur when wheat (Triticum aestivum L.) seedlings were grown under P‐deficient conditions. Wheat plants were grown in a greenhouse and watered with nutrient solution containing or lacking P. During the interval 12 to 18 days after planting, the dry weight of wheat seedlings was similar regardless of P treatment, although the P‐deficient plants had a greater proportion of the total plant weight in the roots. Sixteen days after planting, the roots and leaves of P‐deficient plants had only 20 to 30% the P content of P‐sufficient plants. After 16 days, plants grown under P stress had 41% more p‐nitrophenol phosphatase activity and 70% more β‐glucosidase activity in shoot homogenates than was found in P‐sufficient plants. Changes in both enzyme activities may be involved in the mobilization of plant resources during the early stages of P‐deficient growth.     

Effect of sewage sludge application to amelio‐rate iron deficiency of grain sorghum

Abstract

Recent research has indicated that land application of municipal sewage sludge to calcareous soils can be used to ameliorate iron (Fe) deficiency of grain sorghum [Sorghum bicolor (L.) Monech]. A greenhouse study was conducted to determine the response of grain sorghum grown on three different soils to application of sewage sludge. Sludge applied at rates of 0, 7.5, 15.0, and 25.0 g/kg soil did not completely ameliorate grain sorghum Fe deficiency. When FeEDDHA was soil applied, sewage sludge application significantly increased plant growth due to increases in soil phosphorus (P) availability. Application of sewage sludge at rates greater than 7.5 g/kg reduced dry matter production of grain sorghum in the FeEDDHA amended Orelia SC soil, the soil with the lowest total neutralizing potential. The decreases yield was possibly due to toxic levels of soil and plant copper (Cu) and zinc (Zn), and increased soil salinity.     

Chelation effects on the iron reduction and uptake by low‐iron stress tolerant and non‐tolerant citrus rootstocks

The relative rates of ferric‐iron (Fe³⁺) reduction and uptake by two citrus rootstocks were measured for a series of synthetic Fe³⁺ chelates and microbial siderophores. The rates of Fe³⁺ reduction by the citrus seedlings followed the order: FeHEDTA >> FeDPTA > FeCDTA. No reduction occurred for FeDFOB (ferrioxamine B) and FeTAF (ferric triacetylfusagen). Low rates of Fe³⁺ reduction occurred for Fe2RA3 (ferric rhodotorulic acid). The levels of ⁵⁵Fe taken up the citrus seedlings showed good correlations with the reduction rates. These results indicate the importance of Fe³⁺ reduction in the Fe uptake by citrus rootstocks. The immobility of a large percent of the ⁵⁵Fe taken up by the roots is attributed to the accumulation of Fe in the root apoplasts.     

Iron oxide as an iron source in potting‐soil mixtures

Abstract

A commercial product of Fe oxide‐metallic Fe at rates of 2 and 10 pounds per cubic yard (1.2 and 6 kg per cubic meter) in soil‐sawdust and soil‐bark mixtures was useful when Fe‐inefficient plants were grown in containers. A high Zn level in the Ys₁/Ys₁ Fe inefficient corn inbred (Zea mays L.) was associated with low Fe status. High Fe levels seemed to suppress Cd uptake.