HORTSCIENCE 26(7):853-855. 1991. Response of americana to Nitrogen and Inoculation D.H. Putnam1, G.H. Heichel2, and L.A. Field3 Department of Agronomy and Genetics, 411 Borlaug Hall, University of Minnesota, St. Paul, MN 55108 Additional index words groundnut, , new crops, Bradyrhizobium japonicum, yield, legume symbiosis, nitrogen fertilizers Abstract. Apios americana Medikus (apios) is a wild tuberous legume with potential as a crop plant. Five apios accessions were grown in sand culture in two greenhouse experiments to examine the effect of N fertilization and inoculation with Bradyrhizobium japonicum on yield and plant characteristics. A common B. japonicum strain (USDA76) was applied to watered with 0, 50, 100, 200, or 300 ppm N solutions

(NH4NO3), plus a complete nutrient solution. At 0 N, total dry matter yield of nonin- oculated plants was only » 30% of inoculated plants. However, total dry-matter yields of inoculated plants at 0 N were only »77% of plants supplemented with 50 or 100 ppm, indicating that inoculation alone was insufficient to meet the N needs of the plant. Tuber weight was increased by both N and inoculation, but tuber weight decreased at N concentrations >100 ppm. Differences among plant accessions with regard to tuber fresh weight, harvest index, and modulation were found. These studies indicated that N fertilization maybe required to maximize tuber yields of apios.

Apios americana (groundnut or apios) is its unique role as an N2-fixing and tuber- a wild legume native to the eastern half of producing plant (Putnam et al., 1990). Most North America that produces edible , tuber or root crops, such as potato, fleshy roots, and seeds. According to several (Dioscorea sp.), or sweetpotato (Ipomoea records, the plant was gathered by various batatas L. Lam.), require large inputs of N Native American tribes (Yarnell, 1964). It fertilizers for maximum economic yields. The was consumed by the Pilgrims in their first introduction of a tuberous crop that fixes at-

years in America (Fernald and Kinsey, 1943) mospheric N2 might reduce or eliminate this and probably brought to England as a New requirement. However, to our knowledge, World potato by Sir Walter Raleigh (Schery, there is no published information as to whether 1972). This history, in conjunction with its Rhizobium symbiosis is sufficient to supply N-fixing capabilities and the high protein and the N needs of apios plants. Greenhouse available content of the tuber studies were initiated to assess the impor- (Walter et al., 1986; Wilson et al., 1987), tance of inoculation and additions of mineral has led to interest in apios as a possible new N to plant growth, tuber yield, and nodula- crop (Blackmon and Reynolds, 1986; Duke, tion of A. americana. 1987; Reed and Blackmon, 1985; Reynolds, Greenhouse experiments were conducted et al., 1990; Vietmeyer, 1986). Several wild- using seed harvested from wild plants grown plant food enthusiasts have indicated a taste in Louisiana and North Carolina. The acces- preference for apios over domesticated po- sion numbers and location of origin are pro- tato (Solanum tuberosum L.) (Gibbons, 1974; vided in Table 2. Accessions chosen in the Harris, 1968; Medsger, 1939). 2nd year differed slightly because of their One of the reasons for interest in apios is potentially better adaptation to Minnesota. Accession 5 was added in 1988, and acces- sions 2 and 3 had insufficient seed for further Received for publication 9 Apr. 1990. Joint con- experimentation in 1988. Sand was sterilized tribution of the Minnesota Agr. Expt. Sta. (Sci. and placed in 5.8-liter pots, and surface-ster- J. Series no. 18077), and the USDA-ARS. We ilized seed was planted in a randomized thank Keith Henjum, Univ. of Minnesota, for complete-block design with four replica- helping with the greenhouse work, Peter Graham tions. The trials were initiated on 3 Mar. of the Microbiology Group, Soil Science Dept., Univ. of Minnesota, for producing the inoculum, 1987 (1987 experiment) and 12 Oct. 1987 (1988 experiment). The inoculated treat- H. Keyser, formerly of USDA-ARS of Beltsville, 7 Md., for providing the B. japonicum strain, and ments received 10 cells per plant of a Bra- W.J. Blackmon, Louisiana State Univ., for pro- dyrhizobium japonicum soybean strain, USDA viding apios seed and tubers. The cost of publish- 76, delivered on a peat base. After emer- ing this paper was defrayed in part by the payment gence, the sand was covered with 2 cm of of page charges. Under postal regulations, this pa- perlite to reduce the opportunity for rhizobial per therefore must be hereby marked advertise- cross-contamination and pots were separated ment solely to indicate this fact. by 30 cm. Nitrogen (in the form of NH NO ) 1Assistant Professor. 4 3 2Formerly Plant Physiologist, Plant Science Re- at 0, 100, and 300 ppm N in 1987 and 0, search Unit, USDA-ARS, St. Paul, Minn. Cur- 50, 100, and 200 ppm N in 1988 was applied rent address: Head, Dept. of Agronomy, Univ. twice weekly through plastic access tubes. of Illinois, 1102 S. Goodwin, Urbana, IL 61801. A Hoagland’s complete nutrient solution 3Scientist. (without N), balanced to a pH of 6.8, was

HORTSCIENCE, VOL. 26(7), JULY 1991 853 ditions of 50 (1987) or 100 (1988) ppm N to the inoculated plants (Table 1). Data for 1987 were similar, but are not shown. Noninoculated plants allocated »50% more dry matter to tubers than to shoots than in- oculated plants as indicated by a higher har- vest index at zero N (Table 1). Additions of N did not affect the harvest index or change the difference between inoculated or nonin- oculated plants. These results indicate a dif- ferential allocation of carbon between the inoculated and noninoculated plants that was independent of exogenous N supply. The lower allocation to tubers in the inoculated plants may represent reallocation of carbo-

Fig. 1. Dry matter yield of A. americana in response to inoculation and application of NH4NO3 in hydrate towards N2-fixation in the nodule, 1987 (mean of four accessions). I = inoculated, NI = noninoculated control. (Significant effects at but more detailed studies would have to be P < 0.05: Nitrogen, Inoculation, N ´ I.) conducted to verify this supposition. Combined tuber and root dry-matter yield also applied twice weekly through access tected) least significant difference was used of inoculated treatments was higher than that tubes. Pots were leached to prevent salt ac- to separate accession means (Steel and Tor- of noninoculated treatments at zero N (Figs. cumulation. Supplemental fluorescent light- rie, 1980). 1 and 2). Fresh weight tuber yields of in- ning was provided at a quantum flux density Plant growth and total yield of noninoc- oculated plants at zero N were 86% greater of »500 µmol·s-1·m-2 during an 18/6 h light/ ulated A. americana plants were »30% of than those of noninoculated plants (Table 1). dark cycle. The were trained on bam- those of inoculated treatments at zero N (Figs. However, this difference between inocula- boo stakes. Plants were harvested on 5 Aug. 1 and 2). However, inoculation alone was tion treatments was not consistent in the N- 1987 and 7 June 1988, for the 1987 and 1988 insufficient to maximize total plant weight treated pots. The difference between inocu- experiments, respectively. or tuber yields in either study. Through symb- lated and noninoculated tuber yields, appar- Data collected 2 weeks before harvest in- iosis with B. japonicum, apios produced to- ent at zero N in both years, was reversed at cluded plant height, vigor of shoots, color tal dry matter yields »77% of the yield from 50 or 100 ppm N in the 1988 study. The data of plants, and number of shoots. At harvest, the optimum N plots (Figs. 1 and 2). Total indicated that for production of tubers, non- number of tubers, weight of tubers, root plant and tuber dry matter yield increased inoculated plants were more responsive to weight, viable nodule weight and number, with the addition of N. However, high additions of N than the inoculated plants. and dry weight of herbage were recorded. amounts of N, particularly 200 ppm N (Fig. The increase in tuber yield with 50 or 100 Occasionally, fleshy roots were produced by 2), decreased yield and plants showed symp- ppm added N was accompanied by increases plants, and these were included as tubers. toms of ammonium toxicity. in tuber count only for noninoculated plants Analysis of variance was applied to the data. Tuber production was affected more than (Table 1). Tuber count declined at high con- The sums of squares for N were partitioned total dry matter yields by the addition of N. centrations of N in the noninoculated treat- into orthogonal contrasts of linear, qua- Fresh weight of tubers increased 21% and ments. dratic, and cubic effects. A Fisher’s (pro- 47% (1987 and 1988, respectively) with ad- Plant height, node count, leaf length, color,

854 HORTSCIENCE, VOL. 26(7), JULY 1991 Table 2. Yield and characteristics of Apios americana accessions grown in 1987 and 1988.

zMean of N treatments. yTuber dry weight as a percentage of total dry weight.

Fernald, M.L. and A.C. Kinsey. 1943. Edible wild plants of eastern North America. Eldewild Press, New York. p. 252-255. Gibbons, E. 1974. Stalking the wild asparagus. D. McKay Co., New York. Harris, B.C.. 1968. Eat the weeds. Barre Publi- cations, Barre, Mass. p. 125-127. Medsger, O.P. 1939. Edible wild plants. Mac- millan, New York. p. 187-188. Putnam, D.H., L. Field, and G.H. Heichel. 1990. Inoculation, nitrogen, and cultivar effects on modulation and tuber yield of Apios americana p. 444. In: J. Janick and J.E. Simon (eds.). Advances in new crops. Timber Press, Port- land, Ore. Reed, M.J. and W.J. Blackmon. 1985. Observa- Fig. 2. Dry matter yield of A. americana in response to inoculation and application of NH4NO3 in tions on the potential of Apios americana as a 1988 (mean of three accessions). I = inoculated, NI = noninoculated control. (Significant effects food crop. HortScience 20:557. at P < 0.05; Nitrogen, Inoculation, variety, N ´ I) Reynolds, B.D., W.J. Blackmon, E. Wickreme- sinhe, M.H, Wells, and R.J. Constantin. 1990. and vigor rating (Table 1) were increased by seed-planted apios. However, the fact that Domestication of Apios americana, p. 436442. inoculation and by N in a pattern similar to apios was well nodulated with a common In: J. Janick and J. Simon (eds.). Advances in total dry matter production (Fig. 2). soybean strain in this study and that others new crops. Timber Press, Portland, Ore. There were differences among accessions have reported modulation with rhizobia ob- Schery, R.W. 1972. Plants for man. 2nd ed. Pren- for tuber fresh weight, total dry weight, har- tained from cowpeas (Allen and Allen, 1981) tice Hall, Englewood Cliffs, N.J. vest index, and nodule fresh weight (Table suggests that under agronomic conditions, Steel, R.G.D. and J.H. Torrie. 1980. Principles 2). These apios accessions, obtained from modulation should not be a severe problem. and procedures of statistics, a biometrical ap- the wild, generally exhibited a high degree We have observed nodules on plants grow- proach. 2nd ed. McGraw-Hill, New York. of variability (Blackmon and Reynolds, 1986). ing without any controlled inoculation in ex- Vietmeyer, N. 1986. Lesser-known plants of po- However, the accession XXXVI E3 from perimental plots and on plants collected from tential use in agriculture and forestry. Science Louisiana was more vigorous and produced the wild in Minnesota. 232:1379-1384. a significantly greater fresh weight of tubers Walter, W. M., E.M. Croom, G.L. Catignani, and W.C. Thresher. 1986. Compositional study of in both years than other accessions. Harvest Literature Cited index also tended to be high in this line. This Apios princeana tubers. J. Agr. Food Chem. Allen, D.N. and E.K. Allen. 1981. Apios. The 34:39-41. line also performed well in field trials (Put- Leguminosae-A source book of characteris- nam et al., 1990). Wilson, P.W., F.J. Pichardo, J.A. Livzzo, W.J. tics, uses and modulation. Univ. of Wisconsin Blackmon, and B.D. Reynolds. 1987. Amino If apios is grown in an agronomic or a Press, Madison, Wis. p. 56-57. acids in the American groundnut (Apios amer- horticultural setting it maybe planted by seed Blackmon, W.J. and B.D. Reynolds. 1986. The icana). J. Food Sci. 52(1):224-225. or by tuber. If tubers are used, rhizobial crop potential of Apios americana —preliminary Yarnell, R.A. 1964. Aboriginal relationships be- symbiants likely would accompany the clones, evaluations. HortScience 21:1334-1336. tween culture and plant life in the Upper Great and if the tubers are stored at low tempera- Duke, J.A. 1987. Help rediscover an American Lakes region. Anthropological Papers. no. 23, tures, these Rhizobium may remain viable. vegetable: Apios. Organic Gardening. Jan p. 98- Museum of Anthropology, Univ. of Mich., Ann Inoculation is likely to be more important in 101. Arbor.

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