Structural and Functional Damage Caused by Boron Deficiency in Sunflower Leaves

Photosynthetica - Boron deficiency induced a dramatic inhibition in sunflower plant growth, shown by a reduction in dry mass of roots and shoots of plants grown for 10 d in nutrient solution...     

Structural and Functional Damage Caused by Boron Deficiency in Sunflower Leaves

Boron deficiency induced a dramatic inhibition in sunflower plant growth, shown by a reduction in dry mass of roots and shoots of plants grown for 10 d in nutrient solution supplied with 0.02 μM B. This low B supply facilitated the appearance of brown purple pigmentation on the plant leaves over the entire growth period. Compared to B-sufficient (BS) leaves, leakage from B-deficient (BD) leaves was 20 fold higher for potassium, 38 fold for sucrose, and 6 fold for phenolic compounds. High level of membrane peroxidation was detected by measuring peroxidase activities as well as peroxidative products in BD sunflower plants. Soluble and bound peroxidase activities measured in BD thylakoid membranes were accelerated two fold compared to those detected in BS-membranes. No detectable change in soluble peroxidase activity in roots whereas a 4 fold stimulation in bound peroxidase activity was detected. Thylakoid membranes subjected to low B supply showed enhancement in lipoxygenase activity and malondialdehyde (MDA) content in parallel with 40 and 30 % decrease of linoleic and linolenic acid contents (related to total unsaturated fatty acids). A slower rate of Hill reaction activity (40 %) and a suppressed flow of electron transfer of the whole chain (30 %) were detected in BD thylakoid membranes. This reduction was accompanied with a decline in the activity of photosystem 2 shown by a diminished rate of oxygen evolution (42 %) coupled with a quenching (27.5 %) in chlorophyll a fluorescence emission spectra at 685 nm (F₆₈₅). Thus B is an important element for membrane maintenance, protection, and function by minimizing or limiting production of free oxygen radicals in thylakoid membranes of sunflower leaves. This revised version was published online in June 2006 with corrections to the Cover Date.     

2,6-Dichlorobenzonitrile and Boron Deficiency

Symptoms of 2,6-dichlorobenzonitrile poisoning in plants aredescribed and compared with the symptoms produced by phenylboronicacid and boron deficiency. The effects on the macroscopic andmicroscopic appearance of plants and on their ability to translocategrowth regulators are so alike that it is thought that both2,6-dichlorobenzonitrile and boron deficiency affect the samebasic process.     

Distribution and Partitioning of Newly Taken-up Boron in Sunflower

Distribution and partitioning of newly acquired boron (B) in a mature sunflower (Helianthus annuus L., cv. 3101) plant was investigated. In leaf blades of sunflower plants grown under 0.93, 2.8, and 9.3 mmol B m⁻³, the level of cell-wall-bound B was rather uniform, irrespective of leaf position and B concentration. Boron concentration gradients among leaf positions were produced mainly by different levels of water-soluble B. To determine the distribution of newly taken-up B in plant parts, ¹⁰B-labeled boric acid at a concentration of 2.8 mmol B m⁻³ was applied. The majority of newly acquired B was delivered to the younger leaves, however, approximately one-fourth of the B in the top and second leaves was the older B which was taken up before the 6 d treatment period. In the root tissues, two-fifth of the water-soluble B was new B taken up in the last 6 d, however, within 6 h of the application new B contributed to approximately 80% of the xylem sap B, suggesting that newly taken-up B is preferentially transported to the shoots. When B was withdrawn from the culture solution, the B concentration per leaf area of the lower leaves decreased slightly over 9 d. However, there was an abrupt decrease in the younger leaves, even when taking into account the rapid expansion of the leaf blade, suggesting that B moves more rapidly from the younger leaves than from the older leaves     

Boron in Plants: Deficiency and Toxicity

Boron (B) is an essential nutrient for normal growth of higher plants, and B availability in soil and irrigation water is an important determinant of agricultural production. To date, a primordial function of B is undoubtedly its structural role in the cell wall; however, there is increasing evidence for a possible role of B in other processes such as the maintenance of plasma membrane function and several metabolic pathways. In recent years, the knowledge of the molecular basis of B deficiency and toxicity responses in plants has advanced greatly. The aim of this review is to provide an update on recent findings related to these topics, which can contribute to a better understanding of the role of B in plants.     

Characterization of Boron Deficiency in Soybeans

Boron-deficient soybean roots have a rosette appearance at the terminals caused by the death of the root primordium and the initiation of new root primordia. The root tips show some darkening, swelling, and then a collapse of the tissue. The petiole s of B-deficient plants are very brittle. Physiologically, B-deficiency in HA-soybeans does not appear to involve an interference in the transport of photosynthate (C¹⁴ distribution) or precursors required for citrate synthesis in the roots. More Ca 45 was found in the tops of B-sufficient than B-deficient plants, but the opposite trend existed in the root sap. Boron deficiency symptoms were accentuated by maintaining a relatively low pH of the prenutrient solution by the addition of K salts.     

Translocation of Organic Compounds in Sunflower IV. Effect of Temperature upon Translocation of Sugars

Apical halves of sunflower leaves were fed a solution of either¹⁴C-glucose or ^l4C-glucose and oligomycin at 5 and 25°C,respectively, and temperature, metabolism and translocationof ¹⁴C were measured at 10- to 30-min intervals. ¹⁴C-Labeledcompounds produced by control group leaves were compared tothose produced at 5°C and in the presence of oligomycin,at 25°C. In both experiments, similar small amounts of ¹⁴C-glucosemigrated into the basal half of the leaf. Sucrose translocationfrom the apical half was apparently affected by low temperatureand oligomycin. At least two types (affected and unaffectedby low temperature and oligomycin) of translocation seem tooccur simultaneously in sunflower. (Received July 4, 1980; Accepted November 27, 1980)     

Stem Anoxia Temporarily Interrupts Translocation of 11C-Photosynthate in Sunflower

Treatment of 5.0 cm of stem or petiole of sunflower with nitrogengas for 5 min or longer reduced phloem translocation of ¹¹C-photosynthateor stopped it entirely for a 15–20 min period after whichtime it slowly resumed When the N₂ was replaced by air the transportrestarted immediately and soon reached speeds higher than pre-anoxiarates. Thus anoxia, which we know inhibits export of photosynthateby preventing dark respiration in the leaf (Grodzinski, Jahnke,and Thompson, 1984), also inhibited transport in veins. It reducedthe speed of photosynthate movement eventually stopping it,probably by inhibiting the loading-unloading mechanisms alongthe sieve-tubes. These mechanisms are, apparently, themselves under control ofa metabolic respiratory mechanism (Fensom, 1981). Short-termanoxia of the stem did not appreciably affect the loading andtransport in the leaf; it probably did not destroy the integrityof membranes at the point of treatment nor cause a physicalblockage of sieve-plate pores nor change the viscosity of sieve-tubesap. A metabolic component which assists transport is indicatedin the mechanism of short-term translocation in sunflower. Key words: Sunflower, Petiole, Nitrogen gas, Phloem transport     

Rubidium Uptake and Boron Deficiency in Vicia faba L

Removal of boron from the nutrient solution reduced root elongationwithin 24 h in V. faba seedlings. This was accompanied by areduction in rubidium uptake per plant as well as an altereddistribution within the root. Although the uptake by the terminalcentimetre was reduced in boron deficiency, there was littleeffect on older tissue. The proportion of absorbed Rb transportedto the shoot, efflux rates, and other uptake characteristicswere similar for deficient and normal tissue. The reduced uptakeis considered to be a secondary effect associated with the requirementsof boron for root elongation and metabolism.     

Translocation and Complex Formation of Picloram and 2,4-D in Rape and Sunflower

Uptake, translocation and complex formation of ¹⁴C-labelled 4-amino-3,5,6-trichloropicolinic acid (picloram) and 2,4-dichlorophenoxyacetic acid (2,4-D) in seedlings of rape (Brassica napus L. cv. Nilla) and sunflower (Helianthus annuus L. var. uniflorus) were studied. Sunflower is susceptible both to 2,4-D and picloram, while rape is susceptible to 2,4-D but more tolerant to picloram. The uptake of the herbicides through the leaves was almost complete in both species. Translocation of 2,4-D into the roots took place more readily than that of picloram. In sunflower about 50 per cent of the applied 2,4-D was extruded through the roots into the nutrient solution after 9 days. In the picloram-treated sunflower most of the activity was found in the aerial parts, while in picloram-treated rape most of the activity still occurred in the treated leaf after 9 days. No activity at all was found in the roots or in the nutrient solution of the picloram-treated rape seedlings. While the major part of 2,4-D always was found in the state of free herbicide, a large fraction of picloram was rapidly bound into water-soluble complexes. This binding was especially pronounced in rape. Separation by paper chromatography showed that different radioactive compounds were formed. Most of these could be hydrolyzed, thereby releasing free herbicide. The results support the hypotheses that complex formation could counteract herbicide translocation and toxicity of auxin herbicides.     

Ammonia Gas Temporarily Interrupts Translocation of 11C Photosynthate in Sunflower

Exposure of a leaf of sunflower to ammonia gas for 1 to 2 sreduced translocation of ¹¹C-photosynthate or stopped it entirelyfor a 13 min period, after which time it slowly resumed. Treatmentof a stem of sunflower with NH₃ gas for 30 s to 90 s had a similareffect on ¹¹C translocation in the veins of the stem, by inhibitingloading from adjacent tissue. Unloading of ¹¹C from stem tissuecould, at times, be detected before normal transport resumed.Small applications of NH3 gas did not appear to damage the tissueand are thought to act by inhibiting transport primarily inplasmodesmata. Key words: Translocation, ammonia gas, ¹¹C photosynthate, sunflower     

14CO2 Assimilation and 14C-photosynthate Translocation in Different Leaves of Sunflower

Chlorophyll and nitrogen contents were highest in leaves of middle position, similarly as photosynthetic efficiency represented by ¹⁴C fixation (maxima in leaf 5 from the top). All the leaves lost ¹⁴C after 2 weeks of ¹⁴CO₂ exposure. However, the reduction in radioactivity was less in young upper leaves than in the mature lower leaves. Leaves exported ¹⁴C-photosynthates to stem both above and below the exposed leaf. Very little radioactivity was recovered from the seeds of plants in which only first or second leaves were exposed to ¹⁴CO₂ implying thereby that the carbon contribution of first two leaves to seed filling was negligible. The contribution of leaves to seed filling increased with the leaf position up to the sixth leaf from the top and after the seventh leaf their contribution to seed filling declined gradually.     

Profiles of Leaf Senescence During Reproductive Growth of Sunflower and Maize

We investigated the effect of reproductive growth on the profilesof leaf senescence in maize (Zea mays L.) and sunflower (Helianthusannuus L.). Leaf senescence after flowering was assessed usingboth structural (leaf chlorophyll, nitrogen and dry matter)and functional (photosynthesis) variables in undisturbed plants(+G) and in plants in which grain set was prevented (-G). Twoweeks after flowering, lack of grain accelerated senescencein maize and delayed senescence in sunflower as indicated byleaf chlorophyll; leaf nitrogen and dry matter were less sensitiveresponse variables. Lack of interaction between reproductivetreatment and leaf position indicates that the senescence signal,whatever its nature, was equally effective throughout the plantin both species. In both species, feedback inhibition of photosynthesiswas first detected 30–35 d after flowering; excess carbohydratein the leaves was therefore an unlikely trigger of acceleratedsenescence in maize. As reproductive development progressed,differences between +G and -G plants were more marked in sunflower,and tended to disappear or reverse in maize. In sunflower, interactionsbetween leaf position and reproductive treatment—attributableto the local effect of grain—were detected around 20–27d after flowering. Copyright 2000 Annals of Botany Company Helianthus annuus, Zea mays, chlorophyll, light, nitrogen, photosynthesis, reproductive growth, senescence, source-sink, SPAD.     

Boron Increases Synthesis of Glutathione in Sunflower Plants Subjected to Aluminum Stress

Ionic stress caused by high aluminum (Al) concentrations is one of the most widespread phytotoxicity problems globally in agricultural regions, greatly limiting crop yield in affected areas. The objective of this work was to examine a possible involvement of boron (B) in the detoxification of Al by stimulating glutathione (GSH) metabolism, a mechanism essential for the resistance of plants under stress conditions. Our results clearly demonstrate that increased application of B in the presence of high Al concentrations in the growth medium stimulates GSH biosynthesis, suggesting it could be an effective strategy to combat stress associated with the formation of active-oxygen species (AOS). In the specific case of Al toxicity, B reduces phytotoxicity by stimulating leaf biosynthesis of GSH and an increase in its concentration in the roots. Therefore, in this work, we also identify GSH metabolism as one of the key processes in Al detoxification. Finally, our results imply that greater B application leads to a greater resistance to Al toxicity, a fact that might be significant for higher productivity of agricultural plants grown in acid soils.     

Update on Boron in Higher Plants - Uptake, Primary Translocation and Compartmentation

Abstract: This review focuses on the uptake and primary translocation of boron (B), as well as on the subcellular compartmentation of B and its role in cell walls of higher plants. B uptake occurs via passive diffusion across the lipid bilayer, facilitated transport through major intrinsic proteins (MIPs), and energy-dependent transport through a high affinity uptake system. Whereas the first two represent passive uptake systems, which are constitutively present, the latter is induced by low B supply and is able to establish a concentration gradient for B between the root symplasm and the external medium. At high B supply, a substantial retention of B can be observed at xylem loading, and passive processes are most likely responsible for that. At low B supply, another energy-dependent high affinity transport system for B seems to be induced which establishes an additional concentration gradient between root symplasm and the xylem. The possible significance of all these processes at various B supplies is discussed. The role of soluble B complexes in uptake and primary translocation of B has been evaluated, but the few data available do not allow comprehensive conclusions to be drawn. In any case, there are no indications that soluble B complexes play a major role in either uptake or primary translocation of B. The subcellular compartmentation of B still remains a matter of controversy, but it is unequivocally clear that B is present in all subcellular compartments (apoplasm, cell wall, cytosol and vacuole). The relative distribution of B between these is dependent on plant species and experimental conditions and may vary greatly. Recent results on the well-established role of B in cell walls are summarized and their physiological significance discussed.     

The Micronutrient Boron Causes the Development of Adventitious Roots in Sunflower Cuttings

Three-day-old light-grown sunflower seedlings were de-rootedand incubated in nutrient solutions that either contained orlacked boric acid (B). In the absence of B, in the majorityof the seedlings, no adventitious roots were formed. The micronutrientB caused the development of numerous roots in the lower partof the hypocotyl. The effect of B occurred without the supplyof any phytohormones. A dose-response curve of B-induced rootingyielded an optimum concentration of 0.1 mM boric acid. Histologicalstudies revealed that cell divisions occurred in the controlbut no root primordia developed. In cuttings that were incubatedin B (0.1 m M) root primordia were observed that rapidly developedinto well-differentiated adventitious roots. Sunflower cuttingsthat were planted with their cut end in vermiculite that wasmoistened with nutrient solutions without B degenerated afterseveral weeks. In the presence of B the cuttings formed numerousadventitious roots that entirely replaced the tap root systemof intact seedlings. The rooted cuttings developed into sturdyadult sunflower plants. Our results are discussed with respectto the possible role of B in the evolution of vascular fromprevascular plants.Copyright 1999 Annals of Botany Company Adventitious roots, boron, cuttings, organogenesis, sunflower seedlings.