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DIVISION S-4-SOIL FERTILITY & PLANT NUTRITION

Seed-Zinc Concentration and the Zinc-Efficiency Trait in Navy Bean

^a Dep. of Soil Science, North Dakota State Univ., Fargo, ND 58105 USA
^b Dep. of Plant Sciences, North Dakota State Univ., Fargo, ND 58105 USA

moraghan{at}prairie.nodak.edu

	ABSTRACT

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Some navy bean (Phaseolus vulgaris L.) cultivars are susceptible to Zn deficiency. Norstar and Voyager are examples of Zn-efficient cultivars; Avanti and Albion are examples of Zn-inefficient cultivars. The objective of this study was to compare growth and Zn accumulation, especially in seed, of these four cultivars. In field and greenhouse experiments, except under extremely low soil-Zn levels, the two Zn-efficient cultivars contained >25% higher seed-Zn concentrations than did the two Zn-inefficient cultivars. Both Albion and Avanti, but not Norstar and Voyager, were Zn deficient at a field site with a relatively high level of Zn (1.7 mg kg^-1) extractable by diethylenetrinitrilopentaacetic acid (DTPA). The Zn-efficiency trait was associated with higher levels of Zn in plant tops and was not associated with differences in P or Fe nutrition. Zinc deficiency delayed pod maturity most in the two Zn-inefficient cultivars. Analysis for Zn in seed from routine breeder-yield trials, in which a Zn-efficient cultivar is not Zn deficient, is proposed as a suitable technique for selecting navy bean genotypes that are both agronomically Zn efficient and better sources of Zn for human nutrition.

Abbreviations: DTPA, diethylenetrinitrilopentaacetic acid • EDDHA, ethylenediiminobis(2-hydroxyphenyl)acetic acid

	INTRODUCTION

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ACCORDING TO Graham (1984), a Zn-efficient genotype has the ability to produce a high yield in a soil that is limiting in Zn for a standard genotype. Most studies concerned with diversity in the bean germplasm pool with regard to Zn nutrition have involved the navy bean cultivars Sanilac and Saginaw. Sanilac is a Zn-inefficient cultivar, while Saginaw is a Zn-efficient cultivar (Judy et al., 1965). In various studies, Saginaw contained Zn concentrations similar to (Judy et al., 1965; Polson and Adams, 1970; Edwards and Mohamed, 1973) or greater than (Ambler and Brown, 1969; Mugwira and Knezek, 1971; Jolley and Brown, 1991) Sanilac in vegetative tissue. Differences in absorption of Fe and P (Ambler and Brown, 1969) and of Fe (Jolley and Brown, 1991), rather than differences in Zn absorption, were considered to be the cause of the divergent Saginaw–Sanilac response patterns. Restricted translocation of Zn from roots to tops of Sanilac may be another factor for this cultivar difference (Polson, 1968). Saginaw was also more resistant to Zn toxicity than Sanilac, but this superiority was not associated with variable Zn accumulation in different vegetative tissues (Polson and Adams, 1970). Neither of these cultivars is now widely grown in the United States.

Zinc–phosphorus relationships of a Great Northern, Zn-efficient genotype (cultivar not designated) and Sanilac were compared in a controlled environmental study (Boawn and Brown, 1968). The authors concluded that the two cultivars had similar P–Zn nutritional requirements and that other factors were responsible for their differential behavior under field conditions. No information pertaining to seed-Zn concentration was given in any of the above studies.

Responses of five Australian navy bean cultivars to foliar Zn sprays were compared in a field study (Brouwer et al., 1981). Three of the five cultivars were Zn-inefficient. A major but largely overlooked finding from this study was that seed of the two Zn-efficient cultivars contained higher Zn concentrations than did the seeds of their Zn-inefficient counterparts. Apart from agronomic considerations, an understanding of environmental and genetic factors that influence Zn accumulation in bean seed is also needed, since this commodity is an important food source. Some groups of humans, particularly those on low-meat diets, appear to be at risk in regard to Zn nutrition (Wise, 1995).

We have frequently observed Zn deficiency symptoms in certain navy bean cultivars in our region: Norstar and Voyager (two Zn-efficient cultivars); and Avanti and Albion (two Zn-inefficient cultivars). The objective of our research was to compare the growth and Zn accumulation, especially by seed, of these four cultivars. An associated objective was to determine whether seed-Zn concentration could be used to select Zn-efficient genotypes in a plant breeding program.

	Materials and methods

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This study consisted of two greenhouse and two field experiments. Selected properties of the experimental soils and their descriptions are given in Table 1 . The Zn and Fe concentrations and contents of the seed used in the various experiments are given in Table 2 .

Eight bean seeds were planted in all pots, and seedlings were thinned to three plants per pot 7 d after emergence. All pots were watered, as needed, with distilled, deionized water to the approximate field capacity to avoid wilting. Watering weights were adjusted periodically to account for plant growth. Greenhouse temperatures were maintained between 18 and 22°C. Natural light was supplemented with fluorescent light, 34 µmol m^-2 s^-1 photosynthetically active radiation, to give a 12-h photoperiod.

Greenhouse Experiment 1
This experiment was conducted between October 1996 and February 1997 to determine the effect of Zn (0 and 8 mg ZnSO₄–Zn kg^-1 of air-dried soil) on growth, seed yield, and seed composition of four navy bean cultivars (Albion, Avanti, Norstar, and Voyager). The eight treatments were arranged in a randomized complete block design with four replications.

Mature pods were harvested whenever pedicel tissue died, and a record was kept of cumulative counts. The number of days between emergence and harvest of 75% of the ultimate number of harvested pods was used to measure the effect of added Zn on pod maturity. Senescent leaves were collected daily, and these were combined with any living leaves left on the plants after the final pod harvest. Stem tissue above the cotyledonary node was harvested after the completion of the pod harvest. Seed was separated from pods and counted. Leaves, stems, seed, and pod walls were dried at 70°C for 48 h, weighed, ground to pass a sieve with 0.25-mm openings, redried at 70°C, and analyzed for Zn, Fe, and P. All plant tissue except seed was ground in a stainless steel mill. Seed was ground in an agate mortar with an agate pestle.

Greenhouse Experiment 2
The purpose of this follow-up experiment, conducted during November and December 1997, was to determine the early growth responses of Albion, Avanti, and Norstar to five rates of ZnSO₄–Zn (0, 0.5, 1.0, 2.0, and 4.0 mg kg^-1 Zn of air-dried soil). The 15 treatments were arranged in a randomized complete block design with four replications. The plant tops were cut 1 cm below the cotyledonary node 31 d after emergence. No flowers or pods were present at harvest. The plant material was processed as described for Experiment 1.

Field Experiments
These two 1997 field experiments were established to determine if the differential seed-Zn accumulation patterns of Albion, Avanti, Norstar, and Voyager obtained in Greenhouse Experiment 1 could be reproduced at diverse acid and calcareous soil sites with relatively high levels of DTPA-extractable Zn (Table 1). The dryland field sites were located at Erie and Johnstown, ND. Adequate stored soil moisture and rain prevented plant–water stress at the two experimental sites. The four cultivars were arranged in a randomized complete block design with four replications. Each plot consisted of four rows, 6.1 m in length, with an interrow spacing of 0.76 m.

Four-meter lengths of the two center rows of each plot were harvested mechanically for seed yield. Prior to harvest, 30 pods were randomly selected from the interior rows for chemical analysis. Seed was removed by hand, counted, washed with tapwater containing detergent, rinsed with tapwater, washed in five successive baths with deionized water, dried with cellulose tissue, and processed as described under Greenhouse Experiment 1.

Chemical and Statistical Analysis
Subsamples of the ground plant materials were digested on an aluminum block with 4 mL HNO₃ and 2 mL HClO₄. The acid digests were analyzed for Fe and Zn by atomic absorption spectroscopy and for P by a molybdenum-blue procedure. Standard Reference Material 1572 from the National Institute of Standards and Technology, Gaithersburg, MD, was digested and analyzed concurrently with the bean samples to provide an indication of the accuracy of the analytical procedures. DTPA-extractable Zn and NaHCO₃-extractable P in soil were determined as described by Lindsay and Norvell (1978) and Olsen et al. (1954), respectively. Statistical analyses for the experiments were performed with SAS procedures (SAS Institute, 1985). Differences among mean values were compared by use of Tukey's test (Snedecor, 1956) when the F test from the analysis of variance was significant at the P = 0.05 level.

	Results

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Greenhouse Experiment 1
Norstar and Voyager without added Zn displayed mild leaf chlorosis, typical of Zn-deficiency in beans, prior to flowering. In contrast, Albion and Avanti had severe leaf chlorosis, less vining, more leaf crinkling, and a greater reduction in leaf size. Absence of additional Zn delayed flowering and extended the flowering period for all cultivars, especially for Avanti and Albion. As a result, pod maturity was delayed to the greatest extent in Avanti and Albion (Table 3) . Flower abortion was most pronounced for Albion without added Zn.

Seed-zinc concentration differed little among the four cultivars in the absence of added Zn (Table 3). However, Norstar and Voyager seed had higher Zn concentrations than Albion and Avanti seed in the presence of added Zn. Consequently, the Zn x cultivar interaction was highly significant. Addition of Zn reduced seed P concentration in the four cultivars (Table 3). The effect of Zn on the P content (mg per 3 plants) of plant tops was complex. Although this Zn effect was not significant, the Zn x cultivar effect was highly significant. This resulted from added Zn increasing the P content in Albion tops and decreasing it in Avanti, Norstar, and Voyager. Differential resistance of the four cultivars to Zn deficiency was not related to the Fe accumulation patterns (Table 3). The harvested Voyager seed had the highest Fe concentration in this experiment. In addition, the planted Voyager seed also had the highest Fe concentration among the four cultivars (Table 2).

Greenhouse Experiment 2
By the third trifoliate leaf stage Avanti and Albion had developed severe leaf chlorosis. Addition of 0.5 and 1.0 mg kg^-1 Zn decreased the intensity but did not eliminate this chlorosis. Norstar showed little leaf chlorosis in the absence of added Zn, but leaf size and stem elongation were restricted. Except for slight leaf chlorosis on older leaves of Albion and to a lesser extent on Avanti, plants of the three cultivars treated with 2 mg kg^-1 Zn were healthy and vigorous at harvest.

Dry matter yields for Avanti and Albion tops were relatively similar for given Zn treatments (Table 4) . However, Norstar was less responsive to added Zn. For instance, increasing the Zn-fertilizer rate from 0 to 4 mg kg^-1 Zn resulted in relative dry matter responses of 217, 185, and 77%, for Albion, Avanti, and Norstar, respectively. Response functions showing the different relationships between dry matter yields and Zn-fertilizer rate for Avanti and Albion, as contrasted to Norstar, are given in Table 5 . Polynomial regression functions indicate that higher rates of added Zn, in the range 0 to 4 mg kg^-1 Zn, had progressively smaller effects on Zn concentration in the three cultivars (Table 5). Zinc concentration in plant tops was in the order: Norstar > Avanti > Albion (Table 4). Added Zn decreased P concentration in the tops of all three cultivars. Albion accumulated more Fe in plant tops treated with 0 and 0.5 mg kg^-1 Zn than did Avanti and Norstar.

	Discussion

TOP ABSTRACT INTRODUCTION Materials and methods Results Discussion REFERENCES

Our research with four cultivars, and the results of Brouwer et al. (1981) for five other cultivars, show that analysis of seed Zn in routine plant breeding experiments can be used for selecting Zn-efficient navy bean genotypes. Results from the Erie and greenhouse experiments indicated that plants do not have to be Zn deficient in order for efficient and inefficient genotypes to be separated by this technique. Our experience is that genotypes with seed-Zn concentrations 15 to 20% less than those of Zn-efficient cultivars such as Norstar will likely to be Zn-inefficient. The data from Greenhouse Experiment 1 indicated, however, that in cases of very low soil-Zn availability, where Zn-efficient genotypes are Zn deficient, seed-Zn concentration cannot be used to identify the Zn-efficiency trait. In such cases, the severity of Zn-deficiency symptoms can be used as a selection criterion. Alternatively, selection for Zn-efficient genotypes can be conducted only in experiments where known Zn-efficient genotypes show no symptoms of Zn deficiency. The relationship between agronomic Zn efficiency and seed-Zn concentration found in navy bean does not occur in all crops. For instance, a Zn-efficient wheat (Triticum aestivum L.) cultivar had a lower grain-Zn concentration than a Zn-inefficient cultivar (Graham et al., 1992; Rengel and Graham, 1995).

Analysis of seed has several advantages over analysis of leaf material. Seed can be stored immediately after harvest, while leaf material has to be quickly dried. Mature seed is a defined growth stage and is easy to identify. In contrast, obtaining young recently mature leaflets is subject to error, particularly when genotypes with determinate and indeterminate growth habits are included in the same experiment. A final advantage of using seed as a diagnostic tool is that routine yield-trials of plant breeders do not have to be disturbed during the growth period. Mechanically harvested seed that comes in contact with rubber, galvanized, or brass machinery parts should be washed prior to analysis for Zn (Moraghan, 1985).

Albion showed severe Zn-deficiency symptoms and Avanti showed mild Zn-deficiency symptoms at the Johnstown field site even though DTPA-extractable Zn was 1.7 mg kg^-1 of soil. Likewise, Brouwer et al. (1981) found severe Zn-deficiency symptoms in three Zn-inefficient cultivars at sites with DTPA-extractable Zn ranging from 0.9 to 2.1 mg kg^-1 of soil. The average critical level of DTPA-extractable Zn for most crops is only 0.8 mg kg^-1 of soil (Sims and Johnson, 1991). Commonly used critical levels of 0.5 to 0.8 mg kg^-1 of DTPA-extractable Zn do not work with Zn-inefficient navy bean genotypes.

The responsiveness of seed-Zn concentration to increased availability of soil Zn appears to be common in many higher plants (Rashid and Fox, 1992). However, the potential for increasing seed-Zn concentration in P. vulgaris, compared to that in some other crops, may be limited. Application of 0, 0.6, 4.0, and 8.0 mg ZnSO₄–Zn kg^-1 of soil to the Zn-efficient cultivar Norstar gave seed-Zn levels of 22, 24, 35, and 38 mg kg^-1 (Moraghan, 1996). In contrast, application of 0 and 3.2 mg Zn kg^-1 of soil to wheat resulted in grain with 12 and 155 mg kg^-1 Zn, respectively (Rengel and Graham, 1995). Seed of Norstar and Voyager from the Erie field site, with 5.8 mg kg^-1 of DTPA-extractable soil Zn, had only 32 and 33 mg kg^-1 Zn, respectively. The standard reference value for Zn concentration of navy bean seed consumed in the USA, corrected to a dry-weight basis, is 29 mg kg^-1 (USDA, 1997).

The underlying mechanisms for the Zn-inefficiency trait in navy bean is not obvious from our study. However, differences in Fe and P nutrition were not major causal factors. Results from Greenhouse Experiment 2 indicated that Zn-inefficient genotypes absorb less Zn, but the possibility remains that these genotypes translocate less Zn from the root system to the tops. The three planted seeds used in this experiment contained 15 (Albion), 15 (Avanti), and 18 (Norstar) µg Zn. This contrasted with 23 (Albion), 28 (Avanti), and 57 (Norstar) mg of Zn in the 31-d-old harvested tops without added Zn. Consequently, differential Zn input from seed explains only a small part of the Zn-efficiency trait.

Enhanced release of phytosidephores under Zn-deficiency stress may be causally involved in Zn efficiency in genotypes of graminaceous species (Cakmak et al., 1994). Navy bean is a nongraminaceous species and would not be expected to produce phytosidephores. The possibility that genotypic Zn-efficiency traits may be related to the ability of some genotypes to support mycorrhizal activity has been suggested (Graham and Rengel, 1993). However, high soil P availability reduces vesicular–arbuscular mycorrhizal infection (Tinker, 1980). The Zn-efficiency trait in regard to seed-Zn concentration was exhibited strongly at the Erie site where soil NaHCO₃–P was 47 mg kg^-1. These results do not support the concept that differential mycorrhizal activity is a major cause of the observed genotypic differences.

Zinc deficiency had a greater detrimental effect on Albion than on Avanti, despite the fact that both Zn-inefficient cultivars have similar seed-Zn concentrations when grown under comparable conditions. Seed yield was decreased more, flower abortion was greater, and pod maturity was delayed longer in Albion than in Avanti. Zinc deficiency is known to delay pod maturity in P. vulgaris (Boawn et al., 1969; Blaylock, 1995). This delay can occur even when early-season symptoms of Zn deficiency do not result in seed-yield decreases (Hamilton et al., 1993).

	ACKNOWLEDGMENTS

 
Appreciation is extended to Mr. Kevin Horsager for assistance with this study.

Received for publication April 6, 1998.

	REFERENCES

TOP ABSTRACT INTRODUCTION Materials and methods Results Discussion REFERENCES

 

• Ambler J.E., Brown J.C. Cause of differential susceptibility to zinc deficiency in two varieties of navy beans (Phaseolus vulgaris L.). Agron. J. 1969;61:41-43.
• Blaylock A.D. Navy bean yield and maturity response to nitrogen and zinc. J. Plant Nutr. 1995;18:163-178.
• Boawn L.C., Brown J.C. Further evidence for a P–Zn imbalance in plants. Soil Sci. Soc. Am. Proc. 1968;32:94-97.
• Boawn L.C., Rasmussen P.E., Brown J.W. Relationship between tissue zinc levels and maturity period of field beans. Agron. J. 1969;61:49-51.[Abstract/Free Full Text]
• Brouwer H.M., Stevens G.R., Fletcher J.G. Differential varietal response to zinc foliar sprays in navy beans. Queensl. J. Agric. Anim. Sci. 1981;38:179-185.
• Cakmak S., Gülüt K.Y., Marschner H., Graham R.D. Effect of zinc and iron deficiency on phytosiderophore release in wheat genotypes differing in zinc deficiency. J. Plant Nutr. 1994;17:1-17.
• Edwards G.E., Mohamed A.K. Reduction in carbonic anhydrase activity in zinc deficient leaves of Phaseolus vulgaris L. Crop Sci. 1973;13:351-354.[Abstract/Free Full Text]
• Graham R.D. Breeding for nutritional characteristics. In: Tinker P.B., Läuchli A., eds. Advances in plant nutrition, Vol. 1. New York: Praeger Publ, 1984:57-102.
• Graham R.D., Ascher J.S., Hynes S.C. Selecting zinc-efficient cereal genotypes for soil of low Zn status. Plant Soil 1992;146:241-250.
• Graham R.D., Rengel Z. Genotypic variation in zinc uptake and utilization by plants. In: Robson A.D., ed. Zinc in soils and plants. Dordrecht, the Netherlands: Kluwer Academic Publ, 1993:107-118.
• Hamilton M.A., Westermann D.T., James D.W. Factors affecting zinc uptake in cropping systems. Soil Sci. Soc. Am. J. 1993;57:1310-1315.[Abstract/Free Full Text]
• Jolley V.D., Brown J.C. Factors in iron-stress response mechanism enhanced by Zn deficiency in Sanilac, but not Saginaw navy bean. J. Plant Nutr. 1991;14:257-265.
• Judy, W., J. Melon, G. Lessman, B. Ellis, and J. Davis. 1965. Zinc fertilization of pea beans, corn and sugarbeets in 1964. Mich. Agric. Exp. Stn. Res. Rep. 33., East Lansing, MI.
• Lindsay W.L., Norvell W.A. Development of a DTPA soil test for zinc, iron, manganese, and copper. Soil Sci. Soc. Am. J. 1978;42:421-428.
• Moraghan J.T. Plant tissue testing for micronutrient deficiencies and toxicities. In: Vlek P.L.G., ed. Micronutrients in tropical food production. Dordrecht, the Netherlands: M. Nijhoff Publ, 1985:201-219.
• Moraghan J.T. Zinc concentration of navy bean seed as affected by rate and placement of three zinc sources. J. Plant Nutr. 1996;19:1413-1422.
• Mugwira L.M., Knezek B.D. Growth and plant zinc distribution in two navy bean varieties fertilized with zinc. Commun. Soil Sci. Plant Anal. 1971;2:407-413.
• Olsen S.R., Cole C.V., Watanabe F.S., Dean L.A. Estimation of available phosphorus in soil by extraction with sodium bicarbonate. USDA Circ. 939. Washington, DC: U.S. Gov. Print. Office, 1954.
• Polson D.E. A physiologic-genetic study of the differential response of navy beans (Phaseolus vulgaris L.) to zinc. Diss. Abstr. 1968;29:450B-451B.
• Polson D.E., Adams M.W. Differential response of navy beans (Phaseolus vulgaris L.) to zinc. I. Differential growth and elemental composition at excessive Zn levels. Agron. J. 1970;62:557-560.[Abstract/Free Full Text]
• Rashid A., Fox R.L. Evaluating internal zinc requirement of grain crops by seed analysis. Agron. J. 1992;84:469-474.[Abstract/Free Full Text]
• Rengel Z., Graham R.D. Importance of seed Zn content for wheat growth on Zn-deficient soil. II. Grain yield. Plant Soil 1995;173:267-274.
• SAS Institute. SAS/STAT guide for personal computers, 6th ed Cary, NC: SAS Inst, 1985.
• Sims J.T., Johnson G.V. Micronutrient soil tests. In: Mortvedt J.J., ed. Micronutrients in agriculture. Madison, WI: SSSA, 1991:427-476.
• Snedecor G.W. Statistical methods. Ames, IA: Iowa State College Press, 1956.
• Tinker, P.B. 1980. Role of rhizosphere microorganisms in phosphorus uptake by plants. p. 617–654. In F.E. Khasawneh et al. (ed.) The role of phosphorus in agriculture. ASA, CSSA, and SSSA, Madison, WI.
• U.S. Department of Agriculture. 1997. USDA nutrient database for standard reference. [Online]. Release 11-1. Nutrient data laboratory home page. Available at http://www.nal.usda.gov/fnic/foodcomp (verified 28 April 1999; this web site has replaced USDA-SCS Agric. Handb. 8).
• Wise A. Phytate and zinc bioavailability. Int. J. Food Sci. Nutr. 1995;46:53-63.[Medline]

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