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Brandon R. Smith and Lailiang Cheng

The objective of this study was to quantify how photoprotective mechanisms in the leaves of `Concord' grapevines (Vitis labruscana Bailey) respond to a range of iron (Fe) supply. Own-rooted, 1-year-old container-grown vines were fertigated twice weekly for 11 weeks with a complete nutrient solution containing 1, 10, 20, 50, or 100 μm Fe from ferric ethylenediamine di (o-hydroxyphenylacetic) acid (Fe-EDDHA). Leaf total Fe content did not increase in response to Fe supply; however, “active” Fe (extracted with 2,2′-dipyridyl) and chlorophyll (Chl) increased on a leaf area basis as applied Fe increased. At the lowest active Fe level, leaf absorptance and the efficiency of excitation transfer (Fv′/Fm′) was lower, and nonphotochemical quenching (NPQ) was significantly greater. Photosystem II (PSII) quantum efficiency decreased curvilinearly, and the proportion of PSII reaction centers in the open state (qP) decreased linearly as active Fe content decreased. On a Chl basis, the xanthophyll cycle pool size [violaxanthin (V) + antheraxanthin (A) + zeaxanthin (Z)], lutein, and β-carotene increased curvilinearly as active Fe decreased, and neoxanthin (Neo) increased at the lowest Fe level. On a leaf area basis, as active Fe decreased, V+A+Z and β-carotene decreased curvilinearly, and lutein and Neo decreased linearly. At noon, conversion of V to A and Z increased as active Fe decreased. On a Chl basis, activities of antioxidant enzymes superoxide dismutase (SOD), monodehydroascorbate reductase (MDAR), and dehydroascorbate reductase (DHAR) increased curvilinearly, and glutathione reductase (GR) activity increased linearly as active Fe levels declined. Ascorbate peroxidase (APX) and catalase (CAT), on a Chl basis, were relatively constant. On a leaf area basis, a decrease in active Fe increased SOD and MDAR activity, whereas APX, CAT, DHAR and GR activity decreased. Antioxidant metabolites ascorbate (AsA), dehydroascorbate (DAsA), reduced glutathione (GSH) and oxidized glutathione (GSSG) also increased in response to Fe limitation when expressed on a Chl basis, whereas on a leaf area basis AsA and DAsA decreased and GSH increased curvilinearly. The GSH:GSSG ratio increased as active Fe declined, whereas the AsA:DAsA ratio did not change. In conclusion, both photoprotective mechanisms, xanthophyll cycle-dependent thermal dissipation and the ascorbate-glutathione antioxidant system, are enhanced in response to Fe deficiency to cope with excess absorbed light. In a low soil pH tolerant species such as V. labruscana, the foliar antioxidant system was upregulated in response to excess absorbed light from Fe deficiency-induced chlorosis, and there was no evidence of an increase in oxidative stress from high rates of applied Fe-EDDHA.

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Brandon R. Smith and Lailiang Cheng

‘Concord’ grapevines (Vitis labruscana Bailey) are susceptible to lime-induced chlorosis, which decreases growth and productivity. In two separate experiments, we grew own-rooted vines in a peat–perlite medium adjusted to different pHs with CaCO3 to characterize how lime-induced Fe deficiency affects root and leaf ferric chelate reductase (FCR) and key enzymes and metabolites involved with glycolysis and the tricarboxylic acid (TCA) cycle in leaves. In addition, we measured the pH of the xylem sap as well as Fe, citrate, and malate concentrations. For both experiments, foliar levels of total Fe, active Fe (extracted in 0.1N HCl), and chlorophyll decreased as lime rate increased. An increase in root-medium pH from 5.8 to 7.5 resulted in a 10-fold increase in root FCR activity, whereas leaf FCR activity decreased 10-fold. An increase in root-medium pH did not raise xylem sap pH but decreased Fe and citrate to some extent. Xylem malate was highest at pH 6.6 and decreased both above and below this pH. Foliar data were evaluated in relation to active Fe content, because it is a better indicator of Fe nutritional status. Lower active Fe decreased midday CO2 assimilation and PSII quantum efficiency as well as night respiration. As active Fe decreased, aconitase activity decreased linearly, whereas the activity of glucose-6-phosphate dehydrogenase, NAD(P)-isocitrate dehydrogenase, NAD(P)-malic enzyme, malate dehydrogenase, phosphoenolpyruvate (PEP) carboxylase, PEP phosphatase, and pyruvate kinase increased curvilinearly. Glucose-6-phosphate, fructose-6-phosphate, and 3-phosphoglycerate content decreased curvilinearly as active Fe decreased. Malate content increased as active Fe increased to 1.0 mg·m−2 and then decreased above this level. Citrate increased linearly as active Fe decreased and was an order of magnitude lower than malate content. Our results suggest that leaf FCR activity may limit Fe assimilation to a greater extent than root FCR activity. The decreased leaf aconitase activity under Fe deficiency is the most likely cause of the increase in citrate levels. Greater activity of the other glycolytic and TCA enzymes under Fe deficiency may help to funnel carbon into the mitochondria and enhance NAD(P) reduction. Citrate levels (and the citrate:malate ratios) in the xylem exudate and leaf were much lower when compared with other species and may be linked to Fe inefficiency of ‘Concord’.

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Brandon R. Smith and Lailiang Cheng

`Concord' grapevines (Vitis labruscana Bailey) can readily develop iron deficiency-induced leaf chlorosis when grown on calcareous or high pH soils. Iron (Fe) chelates are often applied to the soil to remedy chlorosis but can vary in their stability and effectiveness at high pH. We transplanted own-rooted 1-year-old `Concord' grapevines into a peat-based medium adjusted to pH 7.5 and fertigated them with 0, 0.5, 1.0, 2.0, or 4mg·L–1 Fe from Fe-EDDHA [ferric ethylenediamine di (o-hydroxyphenylacetic) acid] to determine the effectiveness of this Fe chelate for alleviating Fe deficiency-induced chlorosis at high pH. Vines were sampled midseason for iron, chlorophyll, CO2 assimilation, and photosystem II quantum efficiency (PSII) and at the end of the season for leaf area, dry weight, and cane length. We found that leaf total Fe concentration was similar across all treatments, but active Fe (extracted with 0.1 n HCl) concentration increased as the rate of Fe-EDDHA increased. Chlorophyll concentration increased curvilinearly as applied Fe increased and was highly correlated with active Fe concentration. CO2 assimilation, stomatal conductance, and PSII were very low without any supplemental Fe and increased rapidly in response to Fe application. Total leaf area, foliar dry weight, and cane length all increased as Fe application increased to 1 mg·L–1 Fe, but above this rate, a further increase in Fe did not significantly increase growth. Our results demonstrate that Fe-EDDHA is very effective in alleviating Fe deficiency-induced leaf chlorosis in `Concord' grapevines grown at high pH, which provides a foundation for continuing research related to the optimum rate and timing of application of Fe-EDDHA in `Concord' vineyards on calcareous soils. Compared with total Fe, leaf “active Fe” better indicates the actual Fe status of `Concord' vines.

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Brandon R. Smith and Lailiang Cheng

Plants grown on calcareous soils often exhibit symptoms of Fe-deficiency induced chlorosis despite a high content of total Fe in the leaf tissue. Iron is transported in the xylem primarily as the ferric citrate (Fe-Citr) chelate, and changes in pH, HCO - 3, and Citr can lead to the formation of different Fe-Citr species. Understanding how Fe dissociates from these chelates may help explain why Fe is immobilized in the leaves. The goal was to quantify Fe mobilization (Fe-Mob) from Fe-Citr in an assay system buffered at pH 5, 6, or 7 when: 1) the molar ratio of HCO - 3 to Fe increased in a 1 Fe: 1 Citr system; 2) the molar ratio of Citr increased in a 1 Fe: 3 HCO - 3 system; and 3) solutions were photoreduced (PR) or left in the dark. For non-PR solutions, Fe-Mob from Fe-Citr using 500 μmol NADH was the greatest at the 1 Fe: 0 HCO - 3-level, and decreased as HCO - 3 increased. Fe-Mob also decreased as buffer pH increased from 5 to 7. Increasing the Citr ratio was effective in increasing Fe-Mob, but the effect decreased as buffer pH increased from 5 to 7. PR solutions behaved quite differently. In the 1 Fe: 1 Citr system, little to no Fe-Mob was detected at any buffer pH. However, there were already large pools of Fe2+ in solution, which decreased as HCO - 3 increased, irrespective of buffer pH. Increasing the Citr ratio greatly increased Fe-Mob in the 1 Fe: 3 HCO - 3 system, and mobilization decreased as buffer pH increased. Increasing Citr did not increase the amount of Fe2+ in solution. This work illustrates that increasing the HCO - 3: Fe ratio can lead to an immobilization of Fe, and that increasing the Citr ratio can aid in Fe-Mob from Fe-Citr when the HCO - 3: Fe ratio is high. Increasing the Citr ratio, however, does not increase the amount of PR Fe2+.

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Brandon R. Smith, Paul. R. Fisher and William R. Argo

The objective was to quantify the effect of substrate pH and micronutrient concentration on growth and pigment content for two floricultural crop species, Petunia ×hybrida `Priscilla' and Impatiens wallerana `Rosebud Purple Magic'. A 70% peat: 30% perlite medium was amended with dolomitic hydrated lime to achieve five substrate pH's ranging from pH 4.4 to 7.0. Plants were grown in 10-cm-diameter pots in a greenhouse for 4 weeks, and irrigated with a fertilizer containing (in mg·L-1) 210N-31P-235K-200Ca-49Mg. Micronutrients were applied using an EDTA (ethylenedinitrilotetraacetic acid) chelated micronutrient blend (C111), at 1×, 2×, and 4× concentrations (in mg·L-1) of 0.50Fe-0.25Mn-0.025Zn-0.04Cu-0.075B-0.01Mo. Petunia shoot dry mass and stem caliper decreased as substrate pH increased, whereas leaf length and width remained unchanged. The highest level of C111 resulted in higher dry mass and smaller leaf area compared with other C111 levels. Overall, substrate pH and C111 had little effect on plant size or mass for impatiens. For both species, increasing substrate pH above 5.3 resulted in a decline in chlorophyll, carotenoids, and the SPAD chlorophyll index (measured with a Minolta-502 SPAD meter) compared with the lowest three pH levels. Chlorosis was observed at pH 7 after 2 weeks of growth. Increasing C111 concentration had no effect on pigment content below pH 5.3, but increased pigment content at higher pH levels. The SPAD index was highly correlated with chlorophyll content. This research emphasizes that an acceptable range in substrate pH can vary depending on fertilizer practices, with higher micronutrient concentration compensating for lower solubility at high substrate pH.

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Brandon R. Smith, Paul. R. Fisher and William R. Argo

The objective was to quantify the effect of substrate pH and micronutrient concentration on tissue nutrient levels in Petunia ×hybrida Hort. Vilm.-Andr. and Impatiens wallerana Hook. F. Plants were grown in 10-cm-diameter pots for 4 weeks in a 70% peat: 30% perlite medium amended with five lime rates to achieve substrate pH values ranging from pH 4.4 to 7.0. Plants were irrigated with (in mg·L-1) 210N-31P-235K-200Ca-49Mg. Micronutrients were applied as an EDTA (ethylenedinitrilotetraacetic acid) chelated micronutrient blend (C111), at 1×, 2×, and 4× concentrations of 0.50Fe-0.25Mn-0.025Zn-0.04Cu-0.075B-0.01Mo. Patterns of tissue concentrations across substrate pH differed from nutrient solubility in the medium, particularly with regard to Mn. Foliar N content decreased slightly as substrate pH increased, whereas foliar Ca, Mg, and S increased. Although foliar P and K varied with pH, there was no consistent trend between species. Foliar total Fe, ferrous Fe, and Cu decreased as substrate pH increased, whereas foliar Zn increased. Foliar Mn content decreased for both species as pH rose to 6.0, and then increased from pH 6.0 to 7.0. In contrast, Mn level in the substrate, measured in a saturated medium extract using deionized water as the extractant, decreased as pH increased from pH 4.4 to 7.0. Chlorophyll content decreased when the ratio of tissue Fe to Mn was <0.57 (impatiens) or <0.71 (petunia), or Fe was <106 (impatiens) or 112 (petunia) μg·g-1. SPAD chlorophyll index also declined in petunia with foliar Mn >42 μg·g-1. Increasing C111 increased foliar Cu, total Fe and ferrous Fe in both species, and B for impatiens, and partly compensated for reduced nutrient solubility at high pH.

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Brandon R. Smith*, Li-Song Chen and Lailiang Cheng

Own-rooted one-year-old `Concord' grapevines were fertigated twice weekly for 11 weeks with 1, 10, 20, 50, OR 100 μmol iron (Fe) from ferric ethylenediamine di (o-hydroxyphenylacetic) acid in a complete nutrient solution. As Fe supply increased, leaf total Fe content did not change, whereas active Fe (extracted by 2, 2'-dipyridyl) and total chlorophyll content increased curvilinearly. CO2 assimilation and stomatal conductance increased curvilinearly with increasing active Fe, whereas intercellular CO2 concentrations decreased linearly. Activities of key Calvin cycle enzymes, Rubisco, NADP-glyceraldehyde-3-phosphate dehydrogenase, phosphoribulokinase, stromal fructose-1,6-bisphosphatase (FBPase), and a key enzyme in sucrose synthesis, cytosolic FBPase, all increased linearly with increasing active Fe. No difference was found in the activities of ADP-glucose pyrophosphorylase and sucrose phosphate synthase of leaves between the lowest and the highest treatments, whereas slightly lower activities were observed in the middle Fe treatments. Content of 3-phosphoglycerate increased curvilinearly with increased active Fe, whereas glucose-6-phosphate and fructose-6-phosphate did not change. Glucose, fructose, sucrose, starch, and total non-structural carbohydrates at both dusk and pre-dawn increased with increasing active Fe. Carbon export from starch breakdown during the night, calculated as the difference between dusk and predawn levels, increased as active Fe increased. In conclusion, Fe limitation reduces the activities of Rubisco and other photosynthetic enzymes, and hence CO2 assimilation capacity. Fe-deficient grapevines have lower concentrations of non-structural carbohydrates in source leaves, and therefore, are source limited.

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Li-Song Chen, Brandon R. Smith and Lailiang Cheng

Own-rooted 1-year-old `Concord' grapevines (Vitis labruscana Bailey) were fertigated twice weekly for 11 weeks with 1, 10, 20, 50, or 100 μm iron (Fe) from ferric ethylenediamine di (o-hydroxyphenylacetic) acid (Fe-EDDHA) in a complete nutrient solution. As Fe supply increased, leaf total Fe content did not show a significant change, whereas active Fe (extracted by 2,2′-dipyridyl) content increased curvilinearly. Chlorophyll (Chl) content increased as Fe supply increased, with a greater response at the lower Fe rates. Chl a: b ratio remained relatively constant over the range of Fe supply, except for a slight increase at the lowest Fe treatment. Both CO2 assimilation and stomatal conductance increased curvilinearly with increasing leaf active Fe, whereas intercellular CO2 concentrations decreased linearly. Activities of key enzymes in the Calvin cycle, ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), NADP-glyceraldehyde-3-phosphate dehydrogenase (GAPDH), phosphoribulokinase (PRK), stromal fructose-1,6-bisphosphatase (FBPase), and a key enzyme in sucrose synthesis, cytosolic FBPase, all increased linearly with increasing leaf active Fe. No significant difference was found in the activities of ADP-glucose pyrophosphorylase (AGPase) and sucrose phosphate synthase (SPS) of leaves between the lowest and the highest Fe treatments, whereas slightly lower activities of AGPase and SPS were observed in the other three Fe treatments. Content of 3-phosphoglycerate (PGA) increased curvilinearly with increasing leaf active Fe, whereas glucose-6-phosphate (G6P), fructose-6-phosphate (F6P), and the ratio of G6P: F6P remained unchanged over the range of Fe supply. Concentrations of glucose, fructose, sucrose, starch, and total nonstructural carbohydrates (TNC) at both dusk and predawn increased with increasing leaf active Fe. Concentrations of starch and TNC at any given leaf active Fe content were higher at dusk than at predawn, but both glucose and fructose showed the opposite trend. No difference in sucrose concentration was found at dusk or predawn. The export of carbon from starch breakdown during the night, calculated as the difference between dusk and predawn measurements, increased as leaf active Fe content increased. The ratio of starch to sucrose at both dusk and predawn also increased with increasing leaf active Fe. In conclusion, Fe limitation reduces the activities of Rubisco and other photosynthetic enzymes, and hence CO2 assimilation capacity. Fe-deficient grapevines have lower concentrations of nonstructural carbohydrates in source leaves and, therefore, are source limited.

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Paul R. Fisher, Ron M. Wik, Brandon R. Smith, Claudio C. Pasian, Monica Kmetz-González and William R. Argo

The objective was to evaluate and compare foliar spray and soil drench application methods of iron (Fe) for correcting Fe deficiency in hybrid calibrachoa (Calibrachoa × hybrida) grown in a container medium at pH 6.9 to 7.4. Untreated plants showed severe chlorosis and necrosis, stunting, and lack of flowering. An organosilicone surfactant applied at 1.25 mL·L-1 (0.160 fl oz/gal) increased uptake of Fe from foliar applications of both ferrous sulfate (FeSO4) and ferric ethylenediamine tetraacetic acid (Fe-EDTA). Foliar sprays at 60 mg·L-1 (ppm) Fe were more effective when Fe was applied as Fe-EDTA than FeSO4. Increasing Fe concentration of foliar sprays up to 240 mg·L-1 Fe from Fe-EDTA or 368 mg·L-1 Fe (the highest concentrations tested) from ferric diethylenetriamine pentaacetic acid (Fe-DTPA) increased chlorophyll content compared with lower spray concentrations, but leaf necrosis at the highest concentrations may have been caused by phytotoxicity. Drenches with ferric ethylenediaminedi(o-hydroxyphenylacetic) acid (Fe-EDDHA) at 20 to 80 mg·L-1 Fe were highly effective at correcting Fe-deficiency symptoms, and had superior effects on plant growth compared with drenches of Fe-DTPA at 80 mg·L-1 Fe or foliar sprays. Efficacy of Fe-DTPA drenches increased as concentration increased from 20 to 80 mg·L-1 Fe. An Fe-EDDHA drench at 20 to 80 mg·L-1 Fe was a cost-effective option for correcting severe Fe deficiency at high medium pH.