The effect of soil moisture on the tolerance of Lupinus pilosus genotypes to a calcareous soil

Commercial narrow-leafed lupins (Lupinus angustifolius L.) grown on calcareous soils commonly display chlorotic symptoms resembling Fe deficiency. The severity of chlorosis increases with concurrent increases in soil moisture content. Our research has indicated that the rough-seeded lupin species, Lupinus pilosus Murr., has a range of adaptation to calcareous soils, from tolerant to intolerant. A pot experiment was conducted comparing a tolerant, a moderately tolerant and a moderately intolerant genotype of L. pilosus. Plants were grown for 35 days in a calcareous soil (50% CaCO₃) at three moisture contents (80%, 100% and 120% of field capacity); the growth was compared with that on a fertile black cracking clay control soil at 70% of field capacity. Visual chlorosis score, chlorophyll meter readings, number of leaves and shoot dry weights were recorded at 14, 21, 28 and 35 days after sowing. Concentrations of chlorophyll, active Fe and nutrients in the youngest fully expanded leaves were also measured. Results showed that increased soil moisture increased the severity of chlorotic symptoms (increased chlorosis score) in all genotypes. The tolerant genotype showed significantly less symptoms than other genotypes at all moisture contents. All genotypes were able to recover from chlorosis symptoms at 80% moisture in the calcareous soil. Chlorosis score negatively correlated with chlorophyll meter readings, chlorophyll concentration and foliar active and total Fe, and Mn concentrations. Visual chlorosis score appeared to be a cost effective, accurate and efficient method enabling classification of the tolerance of genotypes. The chlorotic symptoms were likely to be due to HCO₃ ^- induced nutrient deficiencies or a direct effect of HCO₃ ^- on chlorophyll synthesis. This study indicates that the most probable mechanism of tolerance is related to an ability to prevent uptake of HCO₃ ^- or efficiently sequester it once inside the root which prevents increases in internal pH and transport to the shoots.     

Screening rough-seeded lupins (Lupinus pilosus Murr. and Lupinus atlanticus Glads.) for tolerance to calcareous soils

Soil- and solution-based screening methods were used to identify interspecific and intraspecific variation in lupins for tolerance to calcareous soils. Plants were grown for 21 days in a calcareous soil (pH 8.2; 50% CaCO₃; moisture content 90% of field capacity) for soil-based screening and in nutrient solution containing 15 mM KHCO₃ for solution-based screening. Chlorosis as an indicator of tolerance was recorded. Lupinus pilosus Murr. had the most tolerant genotypes and had the greatest range of intraspecific variation. Most genotypes of Lupinus atlanticus Glads. and Lupinus angustifolius L. were moderately intolerant, although two genotypes of L. atlanticus appeared to be tolerant. Lupinus albus L. had moderately tolerant to moderately intolerant genotypes, whilst the single genotypes of Lupinus cosentinii Guss. and Lupinus digitatus Forsk. appeared tolerant. In a field study six genotypes of L. pilosus identified in the soil-based screening as differing in their tolerance to the calcareous soil were grown on comparable calcareous (pH 8.3; topsoil 3% CaCO₃, subsoil 13% CaCO₃) and non-calcareous (pH 7.3) soils within a paddock. Chlorosis and nutrient concentrations in the youngest leaves were measured 53 days after sowing, whilst grain yield was estimated at harvest. Despite the soil containing a much lower CaCO₃ content than used in the screening method, the field study confirmed that moderately intolerant to intolerant genotypes had lower relative grain yields than more tolerant genotypes. Chlorosis rankings of the genotypes were correlated between field and the screening studies. It is suggested that the incorporation of genes conferring tolerance to calcareous soils into high yielding, agronomically suitable genotypes of L. pilosus should be an important objective in a lupin breeding program for calcareous soils.     

Microelement composition of plants grown with low to high levels of sulfur applied to calcareous soil in a glasshouse

Summary PI54619-5-1 soybeans (Glycine max L.), which are very susceptible to Fe deficiency, were grown for 24 days in calcareous (10%) Hacienda loam soil with different levels of S each with and without 2 ppm Fe added as FeEDDHA (ferric ethylenediamine di (o-hydroxyphenylacetic acid). The S application rates ranged from sufficient to neutralize about 15% to more than all of the CaCO₃ present if the S were all oxidized. The soil pH values at harvest time ranged from 7.4 to 6.0. The highest S rate was 10% by weight of soil and it overcame Fe deficiency without FeEDDHA. The S treatments resulted in increased concentrations of Fe and other metals in leaves, but the FeEDDHA treatments increased yields more than did S. At the lower levels of S, the effects of S and FeEDDHA on Fe concentrations in leaves were additive, but not at the highest level of S. The FeEDDHA overcame much of the effect that S had on increasing Mn concentrations in leaves. It had a similar effect, particularly at the low S levels, on Zn, Cu, Al, B, and Ni concentrations in leaves. A level of S sufficient to neutralize only 15% of the CaCO₃ of the soil increased leaf concentrations of Fe, Mn, Zn, Cu, Al, B, Ni, Si, and P. The effect for Zn, Cu, and Al appreared maximum at this level. A combination of the1/2% S and the FeEDDHA resulted in the most favorable micronutrient balance. Bush beans (Phaseolus vulgaris L. var. Improved Tendergreen) grown in calcareous soil with S insufficient to neutralize all the CaCO₃ had increased Mn, Ni, and Mo and decreased Ba levels in leaves. CaSO₄ as a source of S did not have the same effects as elemental S.     

Evaluation of early generations for iron chlorosis in relation to productivity in groundnut (Arachis hypogaea L.)

Five popular but iron-inefficient cultivars were crossed with three efficient genotypes and both parents and F₁s were evaluated for iron-efficiency in potted calcareous and noncalcareous soil. The iron-efficient genotypes were dark green or green in both noncalcareous and calcareous soils whereas inefficient types were light green to yellow in calcareous soil. The chlorophyll and active iron (Fe²⁺) concentration of leaves was less in iron-efficient genotypes compared to efficient types in calcareous soil and reduction of both the parameters from noncalcareous to calcareous soil was considerably high in iron-inefficient lines. There was significant correlation between visual scores, chlorophyll and active iron content. There were no differences among F₁s for iron chlorosis and they were all iron-inefficient. The frequency of iron-inefficient plants was higher than the efficient plants in all F₂ populations. But most of the productive plants came from iron-efficient segregants indicating strong association between iron-efficiency and productivity. Based on the results selection for iron-efficiency in early generations and extensive evaluation for productivity in advanced generations is suggested for developing varieties for cultivation in calcareous soils.     

Foliar fertilization to control iron chlorosis in pear (Pyrus communis L.) trees

The effectiveness of foliar fertilization to re-green chlorotic leaves in iron-deficient pear trees has been studied. Trials were made to assess the influence of (i) the level of Fe deficiency, (ii) the leaf surface treated (adaxial or abaxial), and (iii) two different surfactants, L-77 and Mistol. Treatments were ferrous sulphate alone, ascorbic, citric and sulphuric acids, applied either alone or in combination with ferrous sulphate, Fe-DTPA and water as a control. Solutions were applied with a brush and leaves were treated twice each year. None of the treatments caused a full recovery from Fe deficiency chlorosis. Treatments containing Fe caused the largest re-greening effects, and FeSO₄ had a similar re-greening effect to Fe(III)-DTPA. Increases in leaf Chl were more pronounced with abaxial leaf surface applications and in severely deficient leaves. Using Fe(III)-DTPA in foliar sprays does not seem to be justified, since their effects are not better than those of FeSO₄. The joint use of Fe(III)-DTPA and L-77 and that of FeSO₄ and citric acid do not seem to be suitable. With a single foliar application, FeSO₄ combined with acids gave slightly better results than FeSO₄ alone. Acidic solution applications without Fe may be effective in alleviating chlorosis in some cases, especially in the case of citric acid. In the current state of knowledge, foliar fertilization cannot offer yet a good alternative for full control of Fe chlorosis, although its low environmental impact and cost make this technique a good complementary measure to soil Fe-chelate applications and other chlorosis alleviation management techniques. Abbreviations: Chl – chlorophyll; EDDCHA – ethylenediamine di(5-carboxy-2-hydroxyphenylacetic) acid; EDDHA – ethylenediamine di(o-hydroxyphenylacetic) acid; EDDHMA – ethylenediamine di(o-hydroxy-p-methylphenylacetic) acid; EDDHSA – ethylenediamine di(2-hydroxy-5-sulfophenylacetic) acid     

The effect of drought stress on chlorophyll fluorescence in Lolium-Festuca hybrids

The effects of drought on photochemical efficiency of PSII in leaves of 22 hybrids of Festuca pratensis × Lolium multiflorum and Festuca pratensis × Lolium perenne and of Festuca pratensis cv. Skra were investigated. A significant decrease of electron transport efficiency (about 25%) in PSII (Φ_PSII) was not found before 9 days of seedling growth in hydroponics with water potential (Ψ_w) equal to −0.8 MPa (simulated “soil drought”). The decrease of Φ_PSII was similarly related to that of excitation energy capture by open PSII reaction centre (Fv’/Fm’) and also to the decrease of the proportion of oxidized to reduced Q_A (photochemical fluorescence quenching, q_p). According to the drought prolongation, variation of all parameters of fluorescence between genotypes significantly increased. The seedlings of some genotypes were able to recover electron transport efficiency in PSII after increasing water potential in nutrient solution (removing the “soil drought”). When plants grew in containers with soil and 4 genotypes with the highest sensitivity of electron transport to drought (S) as well as 4 genotypes with the highest tolerance (T) were compared 17 days after watering ceased, Ψ_w in leaves considerably decreased, but the differences between S and T genotypes were often not significant in this respect. The differences between S and T genotypes, as values of Fv/Fm were concerned, also appeared small (about 5%), similarly as that of Fv’/Fm’ (5%), q_p (12%) and Φ_PSII (about 15%). Drought stress increased non-photochemical quenching of chlorophyll fluorescence (NPQ) 15 to 47% and this could protect the PSII reaction centres from damages because of energy excess. The increase of NPQ was not closely connected with drought resistance of plants because it was similar in some genotypes tolerant to dehydration as well as in sensitive ones. The results of the experiments suggest that resources of genetic variability in Festulolium may be sufficient for revealing differences between genotypes on the basis of measurement of chlorophyll a fluorescence, as far as their tolerance to soil drought is concerned. As the tolerance of PSII against drought is high, the determinations of fluorescence should be performed rather under severe stress. Such methods seem to be useful for selection of genotypes with high drought tolerance as well as with the ability to at least partial repairing of PSII after drought.     

Effects of phosphorus and iron fertilizers on the growth of two soybean varieties at two soil temperatures

The influence of FeEDDHA (0, 0.2 and 2 μg Fe g⁻¹ soil) and NaH₂PO₄·H₂O (0 and 120 μg Pg⁻¹ soil) on the growth of two Fe-ineffective soybean (Glycine max L. Merr.) varieties (anoka and T203) on a calcareous soil at two soil temperatures (16 and 24°C) was compared under greenhouse conditions. The two soybean varieties differed in the following respects: (a) T203 accumulated smaller concentrations of Fe in washed tops than Anoka under comparable conditions; (b) T203 was more susceptible to Fe deficiency and its accentuation by high levels of fertilizer P than Anoka; (c) T203 accumulated lower quantities of Mn in tops than Anoka under comparable conditions; (d) T203, but not Anoka, developed Mn deficiency symptoms when treated with P and 2 μg Fe g⁻¹ at 16°C. Fe deficiency was more severe in both varieties at the higher soil temperature due apparently to: (a) greater plant concentration of P in tops at 24°C; and/or (b) an increased rate of plant growth and greater dilution of Fe in young tissue at 24°C. Foliar P concentration was increased much more than foliar Fe concentration by an increase in soil temperature. Severely Fe deficient T203 plants grown without FeEDDHA at 24°C accumulated less foliar Mn than their FeEDDHA counterparts. Comparisons of Fe effectiveness of various soybean cultivars based on relative responses to FeEDDHA can be influenced by differential effects on Mn nutrition.     

Penetration of iron and some organic substances through isolated cuticular membranes

Cuticular membranes were isolated enzymically from tomato fruits and from the dorsal and ventral surfaces of the leaves of Euonymus japonicus. Penetration of Fe from FeSO₄ and FeEDDHA (ferric ethylenediamine di(o-hydroxyphenylacetate) in the absence and presence of urea through the isolated cuticular membranes was studied. Fe from FeSO₄ penetrated more rapidly through the cuticles than Fe from FeEDDHA. Urea reduced the penetration of Fe from FeSO₄ and FeEDDHA. Binding of Fe on the inner surfaces of tomato fruit cuticles was also reduced by EDDHA.     

Rational approaches to control of iron deficiency other than plant breeding and choice of resistant cultivars

Satisfactory progress has been made in recent years in preventing and correcting Fe deficiency in plants, and more can be expected in the future. Important advances include uses of acid- and Fe-fortified organic wastes and use of amended sulfur-pyrite mixes in soil. Three different approaches with organics are as an acidified matrix with Fe, as a means of chelating Fe, and as a carrier of acidifiers. Several procedures can help minimize Fe deficiency. (1) Avoid mis-management of soil physical properties and overirrigation. (2) Avoid nutrient imbalance, such as excess P or excess micronutrients. (3) Use preplant application of mildly acid-organic matter-Fe-sulfur or pyrite mixes worked into zones of soil or banded into seed rows. It is important that small bands or spots in soil be completely neutralized of CaCO₃. (4) Where foliar sprays can or need be used, especially to correct mild chlorosis, use ferrous compounds prepared to be delivered at pH 3± so that the Fe does not easily oxidize or precipitate in the solution. (5) For established trees that have become Fe deficient, inject, via slant drilling of small holes in tree trunks, dilute ferric ammonium citrate sufficient to supply not more than 100 mg kg^-1 Fe to leaves (dry weight basis). Most but not all species will respond. Procedure may be repeated in two or three weeks if necessary. (6) Iron chelates may be used in drip irrigation. If soil is sandy, soil pH not over 7.2, FeDTPA may be used; otherwise, FeEDDHA should be used. If the Fe is supplied with no other nutrients, pH may be at 4 and some FeSO₄ included to recycle the chelating agents. If Fe is used without chelating agents, the pH may be 1.0 or less and other nutrients included. (7) Urea-acid sulfate-Fe sulfate may be irrigated into soil around plants, especially if soil was polymer treated. (8) Efficiency of use of Fe chelates may be increased by making them slow release or by applying with seeds.     

Characterization of the tolerance to iron chlorosis in different peach rootstocks grown in nutrient solution

Iron chlorosis is an important problem in peach trees, but differences exist between peach rootstocks in their tolerance to Fe chlorosis in calcareous soils. The purpose of this investigation was to characterize the tolerance of different rootstocks to Fe chlorosis induced by bicarbonate in nutrient solution. The rootstocks studied included peach (Nemaguard), plums (Brompton, San Julian A and Puebla de Soto 101) and almond × peach hybrids (Adafuel and GF677). Young plants obtained from rooted cuttings or from in vitro culture techniques were grown individually, under controlled conditions, in flasks with 700 mL of aerated nutrient solution low in iron and with or without 10 mM bicarbonate or 10 mM phosphate. Susceptiblity to bicarbonate-induced chlorosis was inversely correlated with both the Fe content in young leaves and the reducing capacity of roots, but not with the phosphorus content in young leaves. The plum Puebla de Soto 101 and the hybrid GF677 showed the lowest degree of chlorosis and the highest reducing capacity. Phosphate did not induce chlorosis.     

Investigations into the exploitation of heterogeneous soils by Lupinus albus L. and L. pilosus Murr. and the effect upon plant growth

In calcareous soils, genotypes of Lupinus albus L. generally grow poorly, resulting in stunted plants that often develop lime-induced chlorosis. In contrast, some genotypes of L. pilosus Murr. occur naturally in calcareous soils without developing any visible symptoms of stress. Some genotypic variation for tolerance to calcareous soil does exist in L. albus and the tolerance mechanisms need to be determined. The adaptation through root system morphological plasticity of L. albus and L. pilosus, to heterogeneous limed soil profiles (pH 7.8) containing either patches of acid (non-limed) soil, or vertically split between acid and limed soil, was investigated. When grown in the presence of patches of acid soil, L. albus had a 52% greater shoot dry weight and visibly greener leaves compared with plants grown in the homogeneous limed soil. Total root dry matter in the acid-soil patches was greater than in the control limed-soil patches. This was due to a four-fold increase in the cluster root mass, accounting for 95% of the root dry matter in the acid-soil patch. Although these cluster roots secreted no more citric acid per unit mass than those in the limed soil did, their greater mass resulted in a higher citrate concentration in the surrounding soil. L. pilosus responded to the patches of acid soil in a manner comparable with L. albus. When grown in the homogeneous limed soil, L. pilosus had a greater maximum net CO₂ assimilation rate (P_max) than L. albus, however, the P_max of both species increased after they had accessed a patch of acid soil. Differences were apparent between the L. albus genotypes grown in soil profiles split vertically into limed and acid soil. A genotype by soil interaction occurred in the partitioning between soils of the cluster roots. The genotype La 674 was comparable with L. pilosus and produced over 11% of its cluster roots in the limed soil, whereas the other genotypes produced only 1–3% of their cluster roots in the limed soil. These results indicate L. pilosus is better adapted to the limed soil than L. albus, but that both species respond to a heterogeneous soil by producing mainly cluster roots in an acid-soil patch. This revised version was published online in June 2006 with corrections to the Cover Date.     

Stress-induced changes in the root architecture of white lupin (Lupinus albus) in response to pH, bicarbonate, and calcium in liquid culture

Current agronomic cultivars of white lupin (Lupinus albus) are intolerant of calcareous or limed soils. In these soils, high pH, bicarbonate (HCO₃^?), and calcium (Ca) concentrations are the major chemical stresses to the root system. To determine the responses of the root system to these factors, evaluate root architecture, and compare genotypes for tolerance, a series of liquid culture experiments was completed using root chambers that allowed the study of the root system in two dimensions. Each stress condition caused changes in different parts of the root system and there was no generalised stress response. HCO₃^? (5 mM) had the greatest effect on cultivars intolerant of calcareous soil; it decreased the dry weight of the shoot and caused the highest percentage of tap root deaths. HCO₃^? also discriminated between short (determinate) and long (indeterminate) roots, as it decreased the number and density of the determinate roots only. Calcium (3 mM) affected all parts of the root system. The tap root was shortened and showed an increased tortuousness in its path compared with 1 mM Ca, although no plants suffered tap root death. The numbers and densities of the two lateral root forms were also decreased, as were the lengths of the indeterminate roots. Stress from alkaline pH (7.5) media caused a lower number and density of determinate lateral roots to be produced than at pH 6.5. The experiments demonstrated that each culture condition elicited a definable stress response. Stress conditions altered the root architecture of genotypes reported to be tolerant of calcareous soil less than in intolerant genotypes. Although soil is more complex than liquid culture, it is possible that in a calcareous or limed soil each stress condition examined may affect the overall stress of the plant, and increased tolerance may result from tolerance to a single stress.     

Nicotianamine and the distribution of iron into the apoplasm and symplasm of tomato (Lycopersicon esculentum Mill.)

The apoplasmic and symplasmic iron pools were determined in roots and leaves of Lycopersicon esculentum Mill. cv. Bonner Beste and its mutant chloronerva. The mutant is auxotrophic for the ubiquitous plant constituent nicotianamine (NA) and exhibits an impaired iron metabolism. Formation of apoplasmic iron pools in roots was dependent on the iron source in the nutrient solution. With Fe-ethylenediaminedi-(2-hydroxyphenylacetate) (FeEDDHA) only a very small apoplasmic iron pool was formed in the roots of both genotypes. Plants grown with FeEDTA increased their apoplasmic iron pool with increasing exogenous iron concentrations in the nutrient solution. The size of the apoplasmic pools in roots did not differ between the wild-type and the mutant (about 85 mol Fe · g^–1 DW). By contrast, the symplasmic iron concentrations in roots and leaves of the mutant were significantly higher when compared to the wild-type. An exogenous NA supply to the leaves of the mutant reduced the high symplasmic iron concentrations to the level of the wild-type. Mutant leaves exhibited a gradient of symplasmic iron concentrations depending on the developmental age of the leaves. The oldest leaves contained considerably more symplasmic iron than the youngest. The results demonstrate that the apparent iron deficiency of the mutant is not the consequence of an impaired iron transport from the apoplasm to the symplasm. Therefore, it is concluded that NA is not required for the transport of Fe(II) through the plasmalemma into the cell.Abbreviations BPDS bathophenanthroline disulfonic acid, Na₂ salt - FeEDDHA ferric N-N-ethylenediaminedi-(2-hydroxy-phenylacetate) - NA nicotianamine Part 40 in the series The normalizing factor for the tomato mutant chloronerva. For part 39 see Pich et al. (1991)The valuable technical assistance of Mrs. Christa Kallas and Mr. Günter Faupel is gratefully acknowledged.     

Effects of pH and bicarbonate on the nutrient status and growth of three Lupinus species

Aims

High pH, and high bicarbonate (HCO₃⁻) and calcium (Ca) availability characterise calcareous soils. High [Ca] only partially explains why some Lupinus species are calcifuge, so we explored high [HCO₃⁻] and high pH.

Methods

We grew six Lupinus genotypes in hydroponics with pH 5, 6.5 and 8^a (adjusted by KOH), and 8^b (adjusted by KHCO₃). Leaf symptoms and areas, root appearance and biomass were recorded; whole leaf and root nutrient concentrations, and leaf cellular phosphorus (P), Ca and potassium (K) concentrations were determined using elemental X-ray microanalysis.

Results

Chlorosis was observed in young leaves at high pH for L. angustifolius and L. cosentinii, and P deficiency at high pH for all genotypes. High pH decreased iron (Fe) and zinc (Zn) uptake in all genotypes. It also decreased lateral root growth, the uptake of P, K, Ca, and manganese (Mn) by all sensitive species; and translocation of P, Fe, Zn, Mn, and Ca to leaves in most sensitive species. However, leaf [Ca], leaf [K], [K] within each measured cell type, and translocation of K and Ca to leaves of L. pilosus and L. cosentinii at pH 8 were greater than at pH 5 and 6.5. Compared with pH 8^a, all L. angustifolius genotypes translocated more P, Fe, Zn, Mn and K from roots to leaves at pH 8^b. High pH did not affect the leaf cell types that accumulated P and Ca, but decreased the leaf cellular [P].

Conclusions

Lupinus angustifolius and L. cosentinii were sensitive to high [HCO₃⁻] and/or high pH; L. pilosus was relatively tolerant. High pH decreased lateral root growth and nutrient uptake, inhibiting growth of sensitive species. High [HCO₃⁻] diminished the negative effect of pH 8 on nutrient translocation to leaves in most L. angustifolius genotypes. This knowledge provides critical insights into the habits of Lupinus species to guide breeding of calcicole plants.

     

Does high bicarbonate supply to roots change availability of iron in the leaf apoplast?

The role of the leaf apoplast in iron (Fe) uptake into the leaf symplast is insufficiently understood, particularly in relation to the supposed inactivation of Fe in leaves caused by elevated bicarbonate in calcareous soils. It has been supposed that high bicarbonate supply to roots increases the pH of the leaf apoplast which decreases the physiological availability of Fe in leaf tissues. The study reported here has been carried out with sunflower plants grown in nutrient solution and with grapevine plants grown on calcareous soil under field conditions. The data obtained clearly show that the pH of the leaf apoplastic fluid was not affected by high bicarbonate supply in the root medium (nutrient solution and field experiments). The concentrations of total, symplastic and apoplastic Fe were decreased in chlorotic leaves of both sunflower (nutrient solution experiment) and grapevine plants in which leaf expansion was slightly inhibited (field experiment). However, in grapevine showing severe inhibition of leaf growth, total Fe concentration in chlorotic leaves was the same or even higher than in green ones, indicative to the so-called `chlorosis paradox'. The findings do not support the hypothesis of Fe inactivation in the leaf apoplast as the cause of Fe deficiency chlorosis since no increase was found in the relative amount of apoplastic Fe (% of total leaf Fe) either in the leaves of sunflower or grapevine plants. It is concluded that high bicarbonate concentration in the soil solution does not decrease Fe availability in the leaf apoplast.     

Iron accumulation in seed of common bean

The effect of soil and genotype on iron concentration [Fe] in common bean (Phaseolus vulgaris L.) seed was studied in the greenhouse. Liming an acid soil increased soil pH from 6.0 to 7.3 but had no effect on seed [Fe] of three bean genotypes (Voyager, T39, UI911) from the Middle American gene pool in North Dakota. However, liming decreased seed-manganese concentration [Mn]. The influence of FeEDDHA on Fe accumulation in seed of the three bean genotypes, grown on acid (pH=6.0) and naturally calcareous (pH=8.2) soils, was also studied in North Dakota. Seed from the acid soil contained 25% higher [Fe] than seed from the calcareous soil. FeEDDHA increased seed [Fe] only on the calcareous soil, but reduced seed [Mn] on both soils. Voyager seed, characterized by a relatively low [Fe] in the seed coat, had a higher seed [Fe] than the other two genotypes. The hypothesis that high seed [Fe] is characterized by a low seed-coat [Fe] was next investigated. Voyager, T39 and 10 diverse Latin American genotypes from the Middle American gene pool were grown on a soil (pH=7.0) with Andic properties in Mexico in the presence and absence of FeEDTA. FeEDTA increased seed [Fe]. Seed of Voyager and a Mexican genotype (Bayo 400) had the highest seed [Fe]. However, Bayo 400, unlike Voyager, contained a high percentage of its seed Fe in the seed coat. Consequently, a high seed [Fe] genotype does not necessarily have a low seed-coat [Fe]. Both soil and genotype affect Fe accumulation in bean seed.     

Mineral composition and growth of Colorado spruce (Picea pungens) seedlings under calcareous soil conditions

Summary Field studies were conducted to assess the mineral nutrition and growth of Colorado spruce (Picea pungens Engln) seedlings (2–4 yr) from provenances selected for superior growth on calcareous prairie soils. Tissue nutrient concentrations and response to nitrogen were determined by use of foliar analysis and growth pattern studies. Soil conditions ranged from 7.6–7.8 for pH, 12–23% for total CaCO₃, and 5–6% for active CaCO₃. Foliage mineral composition showed relatively low phosphorus (0.09–0.15%) and high calcium (0.45–1.52%) assimilation. Seasonal growth and seedling response to added nitrogen was not adversely affected by the calcareous soil condition. Levels of nitrogen in the foliage required for optimum growth ranged from 1.5–2.0% and were similar to that of other conifer species.     

Changes in Chlorophyll Fluorescence in Maize Plants with Imposed Rapid Dehydration at Different Leaf Ages

A comparison of the effects of a rapidly imposed water deficit with different leaf ages on chlorophyll a fluorescence and gas exchange was performed in maize (Zea mays L.) plants. The relationships between photosynthesis and leaf relative turgidity (RT) and ion leakage were further investigated. Leaf dehydration substantially decreased net photosynthetic rate (A) and stomatal conductance (G _s), particularly for older leaves. With dehydration time, F _v /F _m maintained a relatively stable level for youngest leaves but significantly decreased for the older leaves. The electron transport rate (ETR) sharply decreased with intensifying dehydration and remained at lower levels during continuous dehydration. The photochemical quenching of variable chlorophyll fluorescence (q _P) gradually decreased with dehydration intensity for the older leaves but increased for the youngest leaves, whereas dehydration did not affect the nonphotochemical chlorophyll fluorescence quenching (NPQ) for the youngest leaves but remarkably decreased it for the older leaves. The leaf RT was significantly and positively correlated with its F _v /F _m, ETR, and q _P, and the leaf ion leakage was significantly and negatively correlated with F _v /F _m and NPQ. Our results suggest that the photosynthetic systems of young and old leaves decline at different rates when exposed to rapid dehydration.     

Seasonal development of the photosynthetic performance of Norway spruce (Picea abies [L.] Karst.) under magnesium deficiency

In order to investigate the influence of different magnesium nutrition on photosynthesis, one hundred 6-year-old spruce trees derived from one clone were planted in October 1990 into a special out-door experimental construction, where they were cultivated in sand culture with an optimal supply of nutrients, except magnesium, via circulating nutrient solutions. Magnesium was added to the nutrient solutions in three different concentrations, varying from optimal to severe deficient supplies. During the first vegetative period in 1991, photosynthetic performance and carboxylation efficiency were measured under saturating light, controlled CO₂ conditions, optimal temperature and humidity, using a minicuvette system.During summer, the trees under moderate magnesium deficiency developed tip yellowing symptoms on older needles, while the youngest needles remained green with unchanged chlorophyll contents. Trees under severe magnesium deficiency showed yellowing symptoms on all needle age classes combined with decreased chlorophyll contents in the youngest needles as well. In comparison with the controls, the photosynthetic performance of the 1-year-old needles was significantly lower in both deficiency treatments. The same was observed in the youngest needles of the trees under severe deficiency. Trees under moderate deficiency treatment decreased in photosynthetic performance during the summer without reduction of chlorophyll contents. The reduction of photosynthetic rates corresponded to a decrease in carboxylation efficiency, which is taken as a measure of the activity of the enzyme ribulose-1,5-bisphosphate carboxylase. This reduction, together with the observed increase of carbohydrate contents in needles of trees growing under magnesium deficiency, led to the assumption that the photosynthetic carbonfixation is reduced as a consequence of the accumulation of carbohydrates.     

Accumulation of nutrients by eucalyptus seedlings from acidic and calcareous soils of the cerrado region of Central Brazil

Summary The accumulation of nutrients by seedlings ofEucalyptus grandis andE. saligna from a latosol and two other fertile soils was compared. Both species performed well in a calcareous soil and an acidic but fertile gallery forest soil but not in the dystrophic latosol.E. grandis accumulated more nutrients from the calcareous soil thanE. saligna. In the other two soils these two species showed similar growth and nutrient accumulation. In spite of the high acidity, manganese toxicity did not seem to be a limiting factor in the latosol or the gallerty forest soil. Correlations among the shoot concentrations were not always the same as those among the total contents of the nutrients in the aerial biomass.