Organic acids and Fe deficiency: a review

Organic acid concentrations often increase with iron deficiency in different plant parts such as roots, leaves and stem exudates. The review summarises data available on the changes in the concentrations of organic anions in plants with iron deficiency and the effects of these changes in plant metabolism. The paper reviews data available in the literature on the changes in xylem and apoplasmic fluid composition with iron deficiency, both in plants grown in controlled conditions and in the field, and discusses the possible ways of iron complexation and transport in these compartments. The characteristics of the iron reduction and uptake by the iron-deficient leaf mesophyll cells are also discussed, with especial emphasis in the possible roles of organic acids in these processes. Both the possible causes and functions of the organic acid concentration increases in iron-deficient plants are reviewed.     

Stomatal response to drying soil in relation to changes in the xylem sap composition of Helianthus annuus. I. The concentration of cations, anions, amino acids in, and pH of, the xylem sap

Sunflower plants (Helianthus annuus L.) were subjected to soil drying with their shoots either kept fully turgid using a Passioura-type pressure chamber or allowed to decrease in water potential. Whether the shoots were kept turgid or not, leaf conductance decreased below a certain soil water content. During the soil drying, xylem sap samples were taken from individual intact and transpiring plants. Xylem sap concentrations of nitrate and phosphate decreased with soil water content, whereas the concentrations of the other anions (SO₄² and Cl^?) remained unaltered. Calcium concentrations also decreased. Potassium, magnesium, manganese and sodium concentrations stayed constant during soil drying. In contrast, the pH, the buffering capacity at a pH below 5 and the cation/anion ratio increased after soil water content was lowered below a certain threshold. Amino acid concentration of the xylem sap increased with decreasing soil water content. The effect of changes in ion concentrations in the xylem sap on leaf conductance is discussed.     

Iron deficiency-induced changes in the photosynthetic pigment composition of field-grown pear (Pyrus communis L) leaves

In this work we characterize the changes induced by iron deficiency in the pigment composition of pear (Pyrus communis L.) leaves grown under high light intensities in field conditions in Spain. Iron deficiency induced decreases in neoxanthin and β-carotene concomitantly with decreases in chlorophyll a, whereas lutein and carotenoids within the xanthophyll cycle were less affected. Iron deficiency caused major increases in the lutein/chlorophyll a and xanthophyll cycle pigments/chlorophyll a molar ratios. The chlorophyll a/chlorophyll b ratio increased in response to iron deficiency. The carotenoids within the xanthophyll cycle in iron-deficient and in iron-sufficient (control) leaves underwent epoxidations and de-epoxidations in response to ambient light conditions. In control leaves dark-adapted for several hours, most of the xanthophyll cycle pigment pool was in the epoxidated form vio-laxanthin, whereas iron-deficient leaves had significant amounts of zeaxanthin. Iron-deficient leaves also exhibited an increased non-photochemical quenching, supporting the possibility of a role for pigments within the xanthophyll cycle in photoprotection.     

Iron deficiency in peach trees: effects on leaf chlorophyll and nutrient concentrations in flowers and leaves

The effects of iron deficiency on the leaf chlorophyll concentrations and on the macro- (N, P, K, Ca and Mg) and micro-nutrient (Fe, Mn, Zn and Cu) composition of flowers (at full bloom) and leaves (60 and 120 days after full bloom) of field-grown peach (Prunus persica L. Batsch) trees were investigated. Flowers and leaves were taken and analysed from fifty individual trees. Our data indicate that large decreases in leaf chlorophyll concentration were found at the beginning of the season in control trees, possibly associated to a dilution effect by leaf growth, that were later followed by leaf chlorophyll concentration increases. Leaf Fe chlorosis apparently results from two different processes, the dilution of leaf Chl caused by growth and the subsequent inability to produce and/or stabilize new Chl molecules in the thylakoid membrane. Iron chlorosis did not change the seasonal change patterns of any of the nutrients studied. In Fe-deficient trees the K concentration and the K/Ca ratio were high not only in leaves but also in flowers, indicating that this is a characteristic of Fe-deficient plant tissue in the whole fruit tree growing season. Flower Fe concentrations were well correlated with the degree of chlorosis developed later in the season by the trees, suggesting that flower analysis could be used for the prognosis of Fe deficiency in peach.     

Influence of aluminum and phosphorus on growth and xylem sap composition in Melastoma malabathricum L.

Melastoma (Melastoma malabathricum L.) is an aluminum-accumulating woody plant that accumulates more than 10 000 mg kg^–1 of aluminum (Al) in mature leaves. The influence of Al and phosphorus (P) applications on plant growth and xylem sap was examined in the present study in order to elucidate the interaction between Al-induced growth enhancement and P nutrition, and to determine the form of Al for translocation from roots to shoots. Although the Al application significantly increased the growth of Melastomaseedlings with the high P pre-treatment, and P concentrations in the leaves and Pi concentrations in the xylem sap regardless of the P pre-treatment, we could not come to the conclusion that a primary cause of the Al-induced growth enhancement in Melastoma is the stimulation of P uptake. The degree of Al-induced growth enhancement corresponded not with the P concentrations but with the Al concentrations in the plant tissue, suggesting that the Al-induced growth enhancement in Melastoma is primarily caused by Al itself in the plant tissue rather than by the stimulation of P uptake. Through the analysis of organic acids and Al in the xylem sap and plant tissue, the form of Al for translocation from roots to shoots was shown to be an Al-citrate complex that was transformed into Al-oxalate complex for Al storage in the leaves. In addition, the xylem sap of Melastoma seedlings grown in the absence of Al contained higher concentrations of malate. In the presence of Al, however, higher concentrations of citrate were found, indicating that Melastoma changes its organic acid metabolism in the presence or absence of Al; more specifically, it increases the synthesis of citrate.     

Effect of Fe 3+, Zn 2+ and Mn 2+ on ferric reducing capacity and regreening process of the peach rootstock Nemaguard (Prunus persica (L.) Batsch)

The peach rootstock Nemaguard is susceptible to lime-induced iron deficiency chlorosis. Under field conditions, application of ferric chelates to the soil is effective in correcting the Fe-deficiency symptoms. The objectives of this work were to study the induction of the root ferric reducing capacity and the relationship between chlorosis and leaf Fe concentration of plants grown hydroponically under different treatments. Results showed that bicarbonate-treated plants grown with a low Fe concentration increased their reducing capacity if they received additional Fe or Zn for a short period (15 h). However, the addition of Mn had no effect. When these Mn-treated plants were changed to nutrient solution with no bicarbonate and sufficient Fe, regreening was retarded several days in relation to the other treatments. In plants grown without bicarbonate, the reducing capacity was higher in plants grown with a low amount of Fe than in plants grown with either 0 Fe or sufficient Fe. In plants grown with bicarbonate and low Fe, the leaves became chlorotic and had low Fe concentration. When these plants received higher Fe supply, the regreening of the old leaves was not complete, though they had even higher Fe concentrations that the new developing leaves which were completely green. Results are discussed in relation to the Fe or Zn requirements of plants to induce reducing capacity and the incapacity of the cells from chlorotic leaves to absorb Fe and repair metabolic and structural damages.     

Stomatal response to drying soil in relation to changes in the xylem sap composition of Helianthus annuus. II. Stomatal sensitivity to abscisic acid imported from the xylem sap

Sunflower plants [Helianthus annuus L.) were subjected to soil drought. Leaf conductance declined with soil water content even when the shoot was kept turgid throughout the drying period. The concentration of abscisic acid in the xylem sap increased with decreasing soil water content. No general relation could be established between abscisic acid concentration in the xylem sap and leaf conductance due to marked differences in the sensitivity of leaf conductance of individual plants to abscisic acid from the xylem sap. The combination of these results with data from Gollan, Schurr & Schulze (1992, see pp. 551–559, this issue) reveals close connection of the effectiveness of abscisic acid as a root to shoot signal to the nutritional status of the plant.     

Collection and composition of xylem sap and root structure in two halophytic species

Leptochloa fusca (L.) Kunth and Atriplex hortensis (L.) were grown on quartz sand or in liquid culture in the presence of varied concentrations of NaCl. Xylem sap was collected as (a) root pressure exudate, in L. fusca even at 100 mM NaCl, (b) by applying pressure to excised roots of L. fusca and (c) from leaves of the whole plant growing in quartz sand by pressurizing the root system. The latter procedure failed in L. fusca due to the passage of air and soil solution into the leaves. This was caused by an extensive aerenchyma in root cortex. In Atriplex hortensis remarkably high pressures were required to induce a flow of sap. The mineral cation and anion and the amino acid composition of the xylem sap obtained by the different methods was measured and is examined in view of using it for determining the flows of minerals in the whole plant and in relation to the effects of salinity. The spacious aerenchyma in roots of L. fusca has been found to persist also after prolonged exposure to dry air.Presented at the Fourth International Symposium on Structure and Function of Roots, Starà lesna, Slovak Republic. See also PLSO 167/1 (1994).     

Effect of nitrogen nutrition on amino acid composition of xylem sap and stem wood in Alnus glutinosa

Citrulline was the major amino acid in root pressure sap, stem sap and stem wood from Alnus glutinosa L. Gaertn. plants relying on fixed nitrogen or, partly or wholly, on mineral nitrogen for growth. Glutamine increased in prominence in plants assimilating mineral nitrogen but asparagine remained a relatively insignificant component. Differences in the relative amounts of the free amino compounds of stem sap from nitrogen-fixing and mineral nitrogen-fed plants were usually small compared to differences between plants fed different sources of mineral nitrogen. In contrast, relatively high values for the ratios of citrulline/total free amino nitrogen compounds and particularly of citrulline/amides in root pressure sap distinguished nitrogen-fixing plants from those receiving mineral nitrogen. Although the amino acid ratios of stem wood extracts showed closer similarity to those for root pressure sap than stem sap, the seasonal accumulation of citrulline, possibly as a storage amino acid, in stem wood from field-grown plants negated the possibility of utilising stem wood analyses as an indicator of the form of nitrogen assimilation. Comparative data on the levels of citrulline or other free amino acids in Alnus glutinosa are unlikely to be useful as an index of nitrogen fixation, under most experimental conditions.     

Nitrate and reduced-N concentrations in the xylem sap of Stellaria media, Xanthium strumarium and six legume species

Abstract At an applied nitrate concentration of 1 mol m^?3, the proportion of xylem sap nitrogen as nitrate was < 15% for Cajanus cajan, Lupinus albus and Trifolium repens, 33% for Pisum sativum and within the range 57–62% for Glycine max, Phaseolus vulgar is, Stellaria media and Xanthium strumarium. At an applied nitrate concentration of 10 mol m～³ the value had increased to 66% for T. repens while at 20 mol m^?3 nitrate values had increased to 46, 51 and 49% for C. cajan, L. albus and Pisum sativum, respectively, and 89% and 85% for 5. media and X. strumarium, respectively. Glycine max and Phaseolus vulgaris differed from the other species in that the proportion of their xylem sap nitrogen as nitrate remained constant (～ 60%) as applied nitrate concentration increased from 1 to 20 mol m^?3. The proportion of total plant nitrate reductase activity in the shoot of C. cajan, S. media and X. strumarium increased as applied nitrate concentration increased from 1 to 20 mol m^?3. Values at the lower and upper concentrations were, respectively, 26 and 72% for C. cajan. 48 and 80% for X. strumarium and 68 and 87% for S. media. The partitioning of nitrate assimilation between root and shoot in these species is discussed.     

Photosystem II efficiency and mechanisms of energy dissipation in iron-deficient,field-grown pear trees (Pyrus communis L.)

The dark-adapted Photosystem II efficiency of field-grown pear leaves, estimated by the variable to maximum chlorophyll fluorescence ratio, was little affected by moderate and severe iron deficiency. Only extremely iron-deficient leaves showed a decreased Photosystem II efficiency after dark adaptation. Midday depressions in Photosystem II efficiency were still found after short-term dark-adaptation in iron-deficient leaves, indicating that Photosystem II down-regulation occurred when the leaves were illuminated by excessive irradiance. The actual Photosystem II efficiency at steady-state photosynthesis was decreased by iron deficiency both early in the morning and at midday, due to closure of Photosystem II reaction centers and decreases of the intrinsic Photosystem II efficiency. Iron deficiency decreased the amount of light in excess of that which can be used in photosynthesis not only by decreasing absorptance, but also by increasing the relative amount of light dissipated thermally by the Photosystem II antenna. When compared to the controls, iron-deficient pear leaves dissipated thermally up to 20% more of the light absorbed by the Photosystem II, both early in the morning and at midday. At low light iron-deficient leaves with high violaxanthin cycle pigments to chlorophyll ratios had increases in pigment de-epoxidation, non-photochemical quenching and thermal dissipation. Our data suggest that pH could be the major factor controlling thermal energy dissipation, and that large (more than 10-fold) changes in the zeaxanthin plus antheraxanthin to chlorophyll molar ratio caused by iron deficiency were associated only to moderate increases in the extent of photoprotection.This revised version was published online in October 2005 with corrections to the Cover Date.     

The concentration of xylem sap constituents in root exudate, and in sap from intact, transpiring castor bean plants (Ricinus communis L.)

Root exudates were sampled from detopped root systems of castor bean (Ricinus communis). Different volume flux rates were imposed by changing the pneumatic pressure around the root system using a Passioura-type pressure chamber. The concentrations of cations, anions, amino acids, organic acids and abscisic acid decreased hyperbolically when flux rates increased from pure root exudation up to values typical for transpiring plants. Concentrations at low and high fluxes differed by up to 40 times (phosphate) and the ratio of substances changed by factors of up to 10. During the subsequent reduction of flux produced by lowering the pneumatic pressure in the root pressure chamber, the concentrations and ratios of substances deviated (at a given flux rate) from those found when flux was increased. The flux dependence of exudate composition cannot therefore be explained by a simple dilution mechanism. Xylem sap samples from intact, transpiring plants were collected using a Passioura-type root pressure chamber. The concentrations of the xylem sap changed diurnally. Substances could be separated into three groups: (1) calcium, magnesium and amino acid concentrations correlated well with the values expected from their concentration-flux relationships, whereas (2) the concentrations of sulphate and phosphate deviated from the expected relationships during the light phase, and (3) nitrate and potassium concentrations in intact plants varied in completely the opposite manner from those in isolated root systems. Abscisic acid concentrations in the root exudate were dependent on the extent of water use and showed strong diurnal variations in the xylem sap of intact plants even in droughtstressed plants. Calculations using root exudates overestimated export from the root system in intact plants, with the largest deviation found for proton flux (a factor of 10). We conclude that root exudate studies cannot be used as the sole basis for estimating fluxes of substances in the xylem of intact plants. Consequences for studying and modelling xylem transport in whole plants are discussed.     

福建桃、梨果蛀虫及梨小食心虫初步研究

本文报道福建省桃、梨果蛀虫概况及梨小食心虫的初步研究和防治意见。     

Effects of drought on the yellowing status and the dynamics of mineral elements in the xylem sap of declining spruce (Picea abies L.)

In order to test the effects of drought on the occurrence of spruce yellowing, in the Vosges Mountains (northeastern France), the soil of a plot of a 30-year-old spruce stand was protected from throughfall input by a roof for two months during the summer. The degree of yellowing of the trees from the dry and control plots was measured before and after the experiment. Sap flow and sap concentrations were measured in the control plot during the whole of the growing season, and in the dry plot during the drought period. Drought brought about an increase of needle yellowing linked to a reduction in Mg uptake. A decrease in sap concentration of concentration of Ca, Mg and K occurred in the dry plot as compared with the control plot at the end of the drought period. It is concluded that climatic stress could have been the triggering factor of spruce yellowing in the Vosges during the eighties.     

Seasonal dynamics of major cations in xylem sap and needles of Norway spruce (Picea abies L. Karst.) in stands with different soil solution chemistry

Soil solution, xylem sap and needles of mature trees were sampled in three spruce stands over one vegetation period and analysed for major cations. Investigations of nutrient distribution between these three pools and evaluations of seasonal dynamics give the following results: The highest nutrient concentrations in the xylem sap occur at the time of bud break and become gradually lower during the vegetation period. The trees show similar trends of xylem sap concentrations with time for potassium, calcium and magnesium regardless of the nutritional status of the plots. No coupling of xylem sap to soil solution composition can be observed in spite of a high variability of soil solution chemistry in time. The major cations in the current needles exhibit a significantly different trend with time. No time-based correlations for nutrient contents could be found for the needles from the previous year.Despite mobilisation of storage pools in the deficient stand, trees are not able to increase the Ca and Mg contents in the needles up to the level of the other stands. Potassium could be retranslocated in sufficient extent for nutrition of current needles. Due to seasonal variability and dependence upon internal processes, such as retranslocation and mobilisation of nutrients, xylem sap does not seem to be a good tool for the estimation of the nutritional status of forest sites.It was concluded that only minor transport into new foliage via xylem sap will proceed after nutrient flush during the bud break and the nutrient content in the new biomass will be governed by dilution due to biomass growth or by nutrient transport by other means than xylem sap.     

Amino acid content in xylem sap of regrowing alfalfa (Medicago sativa L.): Relations with N uptake,N2 fixation and N remobilization

During vegetative regrowth of Medicago sativa L., soil N, symbiotically fixed N₂ and N reserves meet the nitrogen requirements for shoot regrowth. Experiments with nodulated or non-nodulated plants were carried out to investigate the changes in N flows originating from the different N sources and in xylem transport of amino acids during regrowth. Exogenous N uptake, N₂ fixation and endogenous N remobilization were estimated by ¹⁵N labelling and amino acids in xylem sap were analysed. Removal of shoots resulted in great declines of exogenous N flows derived either from N₂ or from NH₄NO₃ during the first week of regrowth, thereafter recovery increased linearly. Mineral N uptake as well as N₂ fixation occurred mainly between the 10th and 18th day after removal of shoots while exogenous N assimilation in intact plants remained at a steady level. Nitrogen remobilization rates in defoliated plants increased by at least three to five-fold, especially during the first 10 days following shoot removal. Compared to control plants, contents of amino acids in xylem sap, during the first 10 days of regrowth, were reduced by about 72% and 82% in NH₄NO₃ grown and in N₂ fixing plants, respectively. Asparagine was the main amino acid transported in xylem sap of both treated plants. Its relative contents during this period significantly decreased from 75% to 59% and from 67% to 36% respectively in non-nodulated plants and in nodulated ones. This decline was accompanied by compensatory increase in the relative contents of aspartate and glutamine.     

Effects of combined nitrogen on anapleurotic carbon assimilation and bleeding sap composition in Phaseolus vulgaris L.

Dwarf french beans, Phaseolus vulgaris L., were grown with or without inoculation with rhizobia (strain 3644), and with or without a combined nitrogen source (nitrate or ammonium ions). The distribution of radioactivity into products of dark ¹⁴CO₂ assimilation was studied in roots or nodules from these plants. A detailed study was also made of the distribution and rates of excretion of nitrogen in xylem bleeding sap in 28 day old plants grown on the various sources of nitrogen. Whereas detached nodules accumulated radioactive glycine, serine and glutamate when incubated with ¹⁴CO₂, bleeding sap from plants root fed ¹⁴CO₂ contained low levels of radioactivity in these compounds but higher levels in allantoin. Chemical analysis showed allantoin to be the major compound transported in the xylem of nodulated plants, whether or not they were fed on combined nitrogen. In contrast uninoculated plants accumulated mainly amino acids in the bleeding sap, the amount and chemical composition of which depended on the combined nitrogen source.Abbreviations PEP phosphoenol pyruvate - OAA oxaloacetate     

Effects of water stress and rootstocks on fruit phenolic composition and physical/chemical quality in Suncrest peach

The aim of the present work was to evaluate the effects of water stress and different rootstocks on the quality indices and nutritional characteristics of peach fruit, cv. Suncrest. The results show how pre‐harvest factors, such as the type of rootstock and water stress, can influence the quality indices and nutritional properties of peach fruits. Rootstocks type influenced both quality indices such as flesh firmness (FF) and soluble solid content (SSC), but it is unclear how rootstock exerted its influence. Also water stress induced changes in quality indices of peach fruit grafted on GF 677 and Montclar whereas minor effects were recorded when water stress was applied on Penta. The content of hydroxycinnamic acids was similar in control fruits of GF677, Montclar and Penta and it increased significantly only in fruit of trees grafted on GF 677 following water stress. Anthocyanins content in fruit of control plants was higher in Montclar in comparison with the fruit grafted on the other two rootstocks while procyanidin content was higher in fruit of control plants grafted on Penta. Following the water stress an increase in total anthocyanins was observed in GF 677 and Penta whereas no significant differences were found for Montclar. Water stress induced a significant decrease in procyanidin content in Penta. In conclusion in fruits of trees grafted on Penta following water stress quality indices did not show changes but significant variations in some phytochemicals, i.e. anthocyanins and procyanindins were observed. The results suggest that the regulation of water management is an important factor to maintain a good quality of the fruits. In fact, if irrigation stress can induce a higher biosynthesis of phytochemical compounds and an increase of SSC, at the same time, it can change fruit quality traits, such as a lower FW in Montclar, a higher titratable acidity (TA) in GF 677 and Penta and a lower FF in Gf 677 and Montclar.     

桃园生草对土壤有机碳及活性碳库组分的影响

果园生草是解决传统清耕引起的水土流失、土壤有机质减少、土壤肥力下降和果品品质变劣的果园地面管理措施之一。阐明不同牧草种类对土壤有机碳及其活性组分的影响是实现果园土壤生态管理、促进有机果品生产技术体系完善和果园土壤质量提高过程中亟待解决的问题。通过研究桃园人工种植黑麦草和毛苕子对土壤总有机碳(TOC)及其活性组分(微生物量碳(MBC)、水溶性有机碳(WSOC)、轻组有机碳(LFOC)、重组有机碳(HFOC)以及团聚体有机碳)的影响,探明豆科牧草与禾本科牧草种植对桃园土壤有机碳及其组分的影响及其差异。结果表明:与清耕对照(CK)相比,种植黑麦草后土壤TOC、MBC和WSOC分别提高了5.13%、76.4%和18.1%,种植毛苕子土壤LFOC提高了11.3%,土壤HFOC降低了13.2%,但对土壤TOC、MBC和WSOC未产生显著影响。此外,黑麦草显著提高土壤较大粒级(74μm)团聚体有机碳含量,而毛苕子则显著降低土壤较小粒级(2000μm)团聚体有机碳含量。对提高TOC而言,在桃园种植禾本科牧草黑麦草优于豆科牧草毛苕子。土壤MBC、WSOC、LFOC以及HFOC可以作为指示桃园土壤质量提高与否、表征土壤有机碳变化的敏感指标。