Physiological and proteomic analysis reveals the impact of boron deficiency and surplus on alfalfa (Medicago sativa L.) reproductive organs.

Boron (B), an essential element for increasing seed yield and germinability in alfalfa (Medicago sativa L.), plays a vital role in its reproductive processes. However, effects of B stress on physiological and proteomic changes in reproductive organs related to alfalfa seed yield and germinability are poorly understood. In order to gain a better insight into B response or tolerance mechanisms, field trials were designed for B deficiency (0 mg B L-1), B sufficiency (800 mg B L-1), and B surplus (1600 mg B L-1) application during alfalfa flowering to analyze the proteomics and physiological responses of alfalfa 'Aohan' reproductive organs. Results showed that B deficiency weakened the stress-responsive ability in these organs, while B surplus reduced the sugar utilization of 'Aohan' flowers and caused lipid membrane peroxidation in 'Aohan' seeds. In addition, four upregulated stress responsive proteins (ADF-like protein, IMFP, NAD(P)-binding Rossmann-fold protein and NAD-dependent ALDHs) might play pivotal roles in the response of 'Aohan' reproductive organs to conditions of B deficiency and B surplus. All of the above results would be helpful to understand the tolerance mechanisms of alfalfa reproductive organs to both B deficiency and B surplus conditions, and also to give insight into the regulatory role of B in improving seed yield and germinability in alfalfa seed production. In summary, B likely plays a structural and regulatory role in relation to lipid metabolism, carbohydrate metabolism, amino acid metabolism, and signal transduction, thus regulates alfalfa reproductive processes eventually affecting the seed yield and germinability of alfalfa seeds.

Boron: the essential element for vascular plants that never was.

Although a requirement for boron is a well-established feature of vascular plants, its designation, for almost a century, as essential is challenged and, instead, the proposal is made that it has never been so as conventionally defined. This is because an alternative interpretation of published evidence negates its compliance with one of the criteria for essentiality, that its effects are direct. The alternative, here postulated, is that boron is, and always has been, potentially toxic, a feature which, for normal growth, development and reproduction, needed to be nullified. This was enabled by exploitation of boron's ability to be chemically bound to compounds with cis-hydroxyl groups. Although particular cell wall carbohydrate polymers, glycoproteins and membrane glycolipids are among candidates for this role, it is here proposed that soluble phenolic metabolites of, or related to, the components of the pathway of lignin biosynthesis, themselves potentially toxic, are primarily used by vascular plants. When metabolic circumstances allow these phenolics to accumulate endogenously in the cytoplasm, their own inherent toxicity is also alleviated, partially at least, by formation of complexes with boron. This chemical reciprocity, enhanced by physical sequestration of the complexes in vacuoles and/or apoplast, thus achieves, in a flexible but indirect manner, a minimization of the inherent toxicities of both boron and relevant phenolics. In these ways, the multifarious outcomes of impairments, natural or experimental, to this interplay are responsible for the lack of consensus to explain the diverse effects observed in the many searches for boron's primary metabolic role, here considered to be nonexistent. In particular, since a toxic element cannot have 'deficiency symptoms', those previously so-called are postulated to be largely due to the expressed toxicity of phenylpropanoids. A principal requirement for the otherwise toxic boron is to nullify, by means of its indirect chemical and physical sequestration, such expression. In these ways, it is therefore neither an essential nor a beneficial element as currently strictly defined.

Mechanisms of organic acids and boron induced tolerance of aluminum toxicity: A review.

Aluminum is a major limiting abiotic factor for plant growth and productivity on acidic soils. The primary disorder of aluminum toxicity is the rapid cessation of root elongation. The root apex is the most sensitive part of this organ. Although significant literature evidence and hypothesis exist on aluminum toxicity, the explicit mechanism through which aluminum ceases root growth is still indefinable. The mechanisms of tolerance in plants have been the focus of intense research. Some plant species growing on acidic soils have developed tolerance mechanisms to overcome and mitigate aluminum toxicity, either by avoiding entry of Al3+ into roots (exclusion mechanism) or by being able to counterbalance toxic Al3+ engrossed by the root system (internal tolerance mechanism). Genes belonging to ALMT (Aluminum-activated malate transporter) and MATE (Multidrug and toxin compounds extrusion) have been identified that are involved in the aluminum-activated secretion of organic acids from roots. However, different plant species show different gene expression pattern. On the other hand, boron (B) (indispensable micronutrient) is a promising nutrient in the tolerance to aluminum toxicity. It not only hinders the adsorption of aluminum to the cell wall but also improves plant growth. This review mainly explains the critical roles of organic acid and B-induced tolerance to aluminum by summarizing the mechanisms of ALMT, MATE, internal detoxification, molecular traits and genetic engineering of crops.

Boron and the evolutionary development of roots.

Experimental work has shown that Boron (i.e., Boric acid, B) is an essential and multifunctional microelement for vascular plant development. In addition to its other functions, which include xylem development and lignin biosynthesis, we now know that B is involved in phytohormone-signaling and influences the mechanical properties of intercellular pectins. From these data, we conclude that B played an important role during the evolutionary development of lignified tissues, and that it may have been involved in the evolution of vascular plant roots, as hypothesized by D. H. Lewis in 1980. Herein, we review the data pertaining to Lewis' hypothesis, present experimental results on the role of B in root (vs. rhizoid) formation in sunflower vs. a liverwort, and describe the appearance of roots in the fossil record. Open questions are addressed, notably the lack of our knowledge concerning soil microbes and their interactive roles with the micronutrient B during root formation.

[FTIR Spectroscopic Characterization of Material Composition in Functional Leaf of Cotton under Stress of Potassium and Boron].

Potassium (K) and boron (B) are essential nutrient elements for plants, and the elements play an important role for plant growth, development and physiological metabolism. Cotton has a higher demand for K and B; K deficiency or B deficiency often occurs in cotton though. To reveal the component changes in functional leaf of cotton under K and B stress and investigate effects on material composition from K and B. A pot experiment was conducted at Huazhong Agricultural University. (1) the characteristic peaks at 1 546.86, 1 438.85, 1 153.39 and 1 024.17 cm(-1) disappeared due to B deficiency, and relative absorbance of other characteristic peaks was decreased compared with normal, which suggested that the structures of protein, fiber, soluble sugar and ribosome in cotton functional leaf changed and decreased in cotent when lack of K. (2) the relative absorbance of all characteristic peaks was increased in the B-deficient cotton leaves compared with normal, suggesting B deficiency leads to the accumulation in leaves of protein, and fiber, soluble sugar and other carbohydrates because of the hindered transportation. (3) lack of both potassium and boron, induced significant changes to both the locations and relative absorbance of characteristic peaks, and the content of protein, and soluble sugar and other carbohydrates increased, while the content of nucleic acids and polysaccharides dropped. K deficiency led to the structures of protein, fiber, soluble sugar and ribosome in cotton functional leaf changed and decreased in content; B deficiency gave rise to the accumulation in leaves of protein, and fiber, soluble sugar and other carbohydrates; the content of protein and soluble sugar and other carbohydrates increased, while the content of nucleic acids and polysaccharides dropped when K and B were all in short supply.

Effects of boron nutrition and water stress on nitrogen fixation, seed δ15N and δ13C dynamics, and seed composition in soybean cultivars differing in maturities.

Therefore, the objective of the current research was to investigate the effects of foliar B nutrition on seed protein, oil, fatty acids, and sugars under water stress conditions. A repeated greenhouse experiment was conducted using different maturity group (MG) cultivars. Plants were well-watered with no foliar B (W - B), well-watered with foliar B (W + B), water-stressed with no foliar B (WS - B), and water-stressed with foliar B (WS + B). Foliar B was applied at rate of 0.45 kg · ha(-1) and was applied twice at flowering and at seed-fill stages. The results showed that seed protein, sucrose, fructose, and glucose were higher in W + B treatment than in W - B, WS + B, and WS - B. The increase in protein in W + B resulted in lower seed oil, and the increase of oleic in WS - B or WS + B resulted in lower linolenic acid. Foliar B resulted in higher nitrogen fixation and water stress resulted in seed δ (15)N and δ (13)C alteration. Increased stachyose indicated possible physiological and metabolic changes in carbon and nitrogen pathways and their sources under water stress. This research is beneficial to growers for fertilizer management and seed quality and to breeders to use (15)N/(14)N and (13)C/(12)C ratios and stachyose to select for drought tolerance soybean.

Bone formation controlled by biologically relevant inorganic ions: role and controlled delivery from phosphate-based glasses.

The role of metal ions in the body and particularly in the formation, regulation and maintenance of bone is only just starting to be unravelled. The role of some ions, such as zinc, is more clearly understood due to its central importance in proteins. However, a whole spectrum of other ions is known to affect bone formation but the exact mechanism is unclear as the effects can be complex, multifactorial and also subtle. Furthermore, a significant number of studies utilise single doses in cell culture medium, whereas the continual, sustained release of an ion may initiate and mediate a completely different response. We have reviewed the role of the most significant ions that are known to play a role in bone formation, namely calcium, zinc, strontium, magnesium, boron, titanium and also phosphate anions as well as copper and its role in angiogenesis, an important process interlinked with osteogenesis. This review will also examine how delivery systems may offer an alternative way of providing sustained release of these ions which may effect and potentiate a more appropriate and rapid tissue response.

[Comparative study of the urinary excretion of boron, calcium, magnesium and phosphorus in postmenopausal women with and without osteoporosis]. / Estudio comparativo de la excreción urinaria de boro, calcio, magnesio y fósforo en mujeres posmenopáusicas con y sin osteoporosis.

In order to compare the possible relationship between urinary concentrations of boron, calcium, magnesium and phosphorus in serum and urine of postmenopausal women with and without osteoporosis, we selected 45 postmenopausal women over 47 years of age, divided into two groups: group I clinically healthy postmenopausal women and group II postmenopausal women with osteoporosis, without chronic kidney and hepatic diseases or diabetes mellitus. We determined the boron (B), phosphorus (P), total calcium (Ca) and total magnesium (Mg) in the urine of two hours, by atomic emission spectroscopy with induction-coupled plasma (ICPA-ES). Total calcium and total magnesium in serum were determined by atomic flame absorption spectroscopy (FAAS) and inorganic phosphorus in serum, and creatinine in serum and urine, by molecular absorption spectrometry. The preliminary results suggest the existence of a significant difference (p < 0.05) in boron and phosphorus concentrations in the urine of two hours between the groups. The model of linear regression analysis used showed a relationship between urinary concentrations of boron/creatinine index and calcium/ creatinine, magnesium/creatinine and phosphorus/creatinine indexes in the urine of postmenopausal women with osteoporosis.

Estudio comparativo de la excreción urinaria de boro, calcio, magnesio y fósforo en mujeres posmenopáusicas con y sin osteoporosis / Comparative study of the urinary excretion of boron, calcium, magnesium and phosphorus in postmenopausal women with and without osteoporosis

Con el propósito de comparar la posible relación entre las concentraciones urinarias de boro y las concentraciones de calcio, de magnesio y de fósforo en suero y orina de mujeres posmenopáusicas con y sin osteoporosis, seleccionamos 45 mujeres posmenopáusicas con más de 47 años de edad, divididas en dos subgrupos: grupo I mujeres posmenopáusicas clínicamente sanas y grupo II mujeres posmenopáusicas con osteoporosis, sin enfermedades renales, hepáticas o diabetes mellitus. Se determinó el boro (B), el fósforo (P), el calcio total (Ca) y el magnesio total (Mg) en la orina de dos horas por espectroscopia de emisión atómica con plasma acoplado por inducción (ICPA-ES), el calcio y el magnesio total en suero por espectroscopia de absorción atómica en llama (FAAS) y el fósforo inorgánico en suero y la creatinina en suero y orina por espectroscopia de absorción molecular. Los resultados obtenidos sugieren preliminarmente una diferencia significativa (p<0,05) en las concentraciones de boro y de fósforo en la orina de dos horas entre los grupos estudiados. El análisis de regresión lineal aplicado, sugiere relación entre el índice boro/creatinina y los índices calcio/creatinina, magnesio/creatinina y fósforo/creatinina en la orina de las mujeres posmenopáusicas con osteoporosis.

In order to compare the possible relationship between urinary concentrations of boron, calcium, magnesium and phosphorus in serum and urine of postmenopausal women with and without osteoporosis, we selected 45 postmenopausal women over 47 years of age, divided into two groups: group I clinically healthy postmenopausal women and group II postmenopausal women with osteoporosis, without chronic kidney and hepatic diseases or diabetes mellitus. We determined the boron (B), phosphorus (P), total calcium (Ca) and total magnesium (Mg) in the urine of two hours, by atomic emission spectroscopy with induction-coupled plasma (ICPA-ES). Total calcium and total magnesium in serum were determined by atomic flame absorption spectroscopy (FAAS) and inorganic phosphorus in serum, and creatinine in serum and urine, by molecular absorption spectrometry. The preliminary results suggest the existence of a significant difference (p <0.05) in boron and phosphorus concentrations in the urine of two hours between the groups. The model of linear regression analysis used showed a relationship between urinary concentrations of boron/creatinine index and calcium/ creatinine, magnesium/creatinine and phosphorus/creatinine indexes in the urine of postmenopausal women with osteoporosis.

Boron deficiency and transcript level changes.

Boron (B) is an essential element for plant growth whose deficiency causes an alteration in the expression of a wide range of genes involved in several physiological processes. However, our understanding of the signal transduction pathways that trigger the B-deficiency responses in plants is still poor. The aims of this review are (i) to summarize the genes whose transcript levels are affected by B deficiency and (ii) to provide an update on recent findings that could help to understand how the signal(s) triggered by B deficiency is transferred to the nucleus to modulate gene expression. In this contribution we review the effects of B deficiency on the transcript level of genes related to B uptake and translocation, maintenance of cell wall and membrane function, nitrogen assimilation and stress response. In addition, we discuss the possible mediation of calcium, arabinogalactan-proteins and other cis-diol containing compounds in the signaling mechanisms that transfer the signal of B deficiency to nuclei. Finally, we conclude that the advance in the knowledge of the molecular basis of B deficiency response in plants will allow improving the tolerance of crops to B deficiency stress.

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.

Is boron nutritionally relevant?

Evidence from numerous laboratories using a variety of experimental models, including humans, shows that boron is a bioactive beneficial element. Much evidence has come from studies that did not require nutritional or environmental stressors or fastidious methods in diet preparation or environmental control. The evidence includes deprivation studies showing that boron is necessary for some higher animals to complete the life cycle, and that realistic low boron intakes result in impaired bone health, brain function, and immune response. Thus, low boron intake is a relevant nutritional concern, which diets rich in fruits, vegetables, nuts, and pulses can prevent.

Physiological roles and transport mechanisms of boron: perspectives from plants.

Boron, an orphan of the periodic table of the elements, is unique not only in its chemical properties but also in its roles in biology. Its requirement in plants was described more than 80 years ago. Understandings of the molecular basis of the requirement and transport have been advanced greatly in the last decade. This article reviews recent findings of boron function and transport in plants and discusses possible implication to other organisms including humans.

Effect of thawing time, cooling rate and boron nutrition on freezing point of the primordial shoot in norway spruce buds.

BACKGROUND: Effects of cooling rates on bud frost hardiness have been studied but there is little information on bud responses to thawing. Since the cell wall pore size has been found to increase with boron (B) deficiency, B deficiency may affect the supercooling ability of buds in winter. METHODS: The effects of duration of thawing time and rate of cooling on bud frost hardiness of Norway spruce (Picea abies) were studied in a B fertilization trial in February 2003 and March 2005. Frost hardiness of apical buds was determined by differential thermal analysis (DTA) and visual scoring of damage. KEY RESULTS: In 2003, the freezing point of primordial shoots of buds (T(f)), i.e. the low-temperature exotherm (LTE), was, on average, -39 degrees C when buds were thawed for less than 3 h and the T(f) increased to -21 degrees C after 18 h of thawing. During the first 4 h of thawing, the rate of dehardening was 6 degrees C h(-1). In 2005, buds dehardened linearly from -39 degrees C to -35 degrees C at a rate of 0.7 degrees C h(-1). In 2003, different cooling rates of 1-5 degrees C h(-1) had a minor effect on T(f) but in 2005 with slow cooling rates T(f) decreased. In both samplings, at cooling rates of 2 and 1 degrees C h(-1), T(f) was slightly higher in B-fertilized than in non-fertilized trees. By contrast, at very short thawing times in 2003, T(f) was somewhat lower in B-fertilized trees. CONCLUSIONS: There was little evidence of reduced frost hardiness in trees with low B status. This study showed that buds deharden rapidly when exposed to above-freezing temperatures in winter, but if cooled again they reharden more slowly. According to this study, rapid dehardening of buds has to be taken into account in assessments of frost hardiness.

Involvement of auxin and CKs in boron deficiency induced changes in apical dominance of pea plants (Pisum sativum L.).

It has previously been shown that boron (B) deficiency inhibits growth of the plant apex, which consequently results in a relatively weak apical dominance, and a subsequent sprouting of lateral buds. Auxin and cytokinins (CKs) are the two most important phytohormones involved in the regulation of apical dominance. In this study, the possible involvement of these two hormones in B-deficiency-induced changes in apical dominance was investigated by applying B or the synthetic CK CPPU to the shoot apex of pea plants grown in nutrient solution without B supply. Export of IAA out of the shoot apex, as well as the level of IAA, Z/ZR and isopentenyl-adenine/isopentenyl-adenosine (i-Ade/i-Ado) in the shoot apex were assayed. In addition, polar IAA transport capacity was measured in two internodes of different ages using 3H-IAA. In B-deficient plants, both the level of auxin and CKs were reduced, and the export of auxin from the shoot apex was considerably decreased relative to plants well supplied with B. Application of B to the shoot apex restored the endogenous Z/ZR and IAA level to control levels and increased the export of IAA from the shoot apex, as well as the 3H-IAA transport capacity in the newly developed internodes. Further, B application to the shoot apex inhibited lateral bud growth and stimulated lateral root formation, presumably by stimulated polar IAA transport. Applying CPPU to the shoot apex, a treatment that stimulates IAA export under adequate B supply, considerably reduced the endogenous Z/ZR concentration in the shoot apex, but had no stimulatory effect on IAA concentration and transport in B-deficient plants. A similar situation appeared to exist in lateral buds of B-deficient plants as, in contrast to plants well supplied with B, application of CKs to these plants did not stimulate lateral bud growth. In contrast to the changes of Z/ZR levels in the shoot apex, which occurred after application of B or CPPU, the levels of i-Ade/i-Ado stayed more or less constant. These results suggest that there is a complex interaction between B supply and plant hormones, with a B-deficiency-induced inhibition of IAA export from the shoot apex as one of the earliest measurable events.

Boron deficiency increases putrescine levels in tobacco plants.

Polyamine concentrations were determined in leaves and roots of tobacco plants (Nicotiana tabacum L.) subjected to a short-term boron deficiency. A decrease in the growth of shoots and, especially, roots was found under this mineral deficiency. Boron deficiency did not lead to a significant decrease in leaf or root ion concentrations when compared to control treatment; however, as expected, leaf boron concentration was lower in boron-deficient plants in comparison to the control. In leaves, the levels of free putrescine and spermidine were similar in both treatments. In roots, a short-term boron deficiency caused an increase in free putrescine. Moreover, boron-deficient plants had higher conjugated polyamine concentration than boron-sufficient plants, which was especially evident for conjugated putrescine in leaves. A possible link between boron and polyamine levels is proposed and discussed.