were less than in plants grown in ProMix in the media study. Fertilization regimes were slightly different. Plants in the substrate study received a weekly application of 100 mg·L-1N, whereas plants in the bottom heat study received twice weekly applications of 50 mg·L-1N fertilizer. Also, warmer ambient greenhouse temperatures, 29.1 ± 0.7 °C day and 21.2 ± 0.1 °C night, during the media experiment likely resulted in the greater growth of pawpaw seedling compared to those, 27 ± 0.8 °C day and 21 ± 0.2 °C night in the bottom heat experiment. Plants in the substrate experiment were subjected to warmer daytime temperatures, but lower temperatures at night. It is possible that there were higher losses of carbon during the night in bottom-heated plants compared to plants in the media experiment, due to increased nighttime respiration with bottom heat.

Hurewitz and Janes (1983) noted that tomato leaf carbon exchange rate and leaf area increased as soil temperature increased to 30 °C. They hypothesized that increased plant growth at warmer soil temperatures may have been due to the effect of increased root sink activity on leaf photosynthetic rate. Other factors that may have contributed to the growth enhancement noted for bottom-heated seedlings included changes in water use efficiency and production or availability of root-derived cytokines. Also, with such heating there are potential decreases in both the time to produce a saleable plant and the cost of heating greenhouses (Vogelezang, 1991). Bottom-heated seedlings also produced greater lateral and total root dry weight than ambient seedlings, which could result in a higher rate of establishment of bottom-heated seedlings in the field.

In conclusion, ProMix with a once weekly application of 100 mg·L-1 N as Peters water-soluble fertilizer served as an excellent substrate for the production of robust pawpaw seedlings for transplanting into the field or into larger containers. Pawpaw seedlings responded very favorably to root-zone warming to 32 °C, by speeding seedling emergence, and increasing shoot, root, and total dry matter production in developing seedlings.

Literature Cited

Aiello, A.S. and W.R. Graves. 1997. Container medium and nitrogen form affect production of Amur Maackia (Maackia amurensis Rupr. &

Maxim.). HortScience 32:1200–1203.

Arnold, M.A. and D.K. Struve. 1989. Growing green ash and red oak in CuCO3-treated containers increase root regeneration and shoot growth following transplant. J. Amer. Soc. Hort. Sci. 114:402–406.

Bode-Stoltzfus, R.M., H. G. Taber, and A.S. Aiello. 1998. Effect of increasing root-zone temperature on growth and nutrient uptake by ‘Gold Star’ muskmelon plants. J. Plant Nutr. 21:321–328.

Cruz, C., S.H. Lips, and M.A. Martins-Loucao. 1993. Growth and nutrition of carob plants as affected by nitrogen sources. J. Plant Nutr. 16:1–15.

Darrow, G.M. 1975. Minor temperate fruits, p. 276–277. In: J. Janick and J.N. Moore (eds.) Advances in fruit breeding. Purdue Univ. Press, West Lafayette, Ind.

Davidson, H., R. Mecklenberg, C. Peterson. 2000. Nursery Management, Administration and Culture. Prentice Hall. Upper Saddle River, N.J.

Finneseth, C.L.H., D.R. Layne, and R.L. Geneve. 1998a. Morphological development of pawpaw [Asimina triloba (L.) Dunal] during seed germination and seedling emergence. HortScience 33:802–805.

Finneseth, C.L.H., D.R. Layne, and R.L. Geneve. 1998b. Requirements for seed germination in North American pawpaw [Asimina triloba (L.) Dunal]. Seed Sci. Technol. 26:471–480.

Finneseth, C.L.H., D.R. Layne, and R.L. Geneve. 2000. Establishment of North American paw-paw [Asimina triloba (L.) Dunal] shoots in vitro from mature and juvenile explants. Acta Hort. 520:97–102.

Graves, W.R., R.J. Joly, and M.N. Dana. 1989. Root-zone temperature affects water status and growth of red maple. J. Amer. Soc. Hort. Sci. 114:406–410.

Graves, W.R., R.J. Joly, and M.N. Dana. 1991. Water use and growth of honey locust and tree-of-heaven at high root-zone temperature. HortScience 26:1309–1312.

Graves, W.R. and A.S. Aiello. 1997. High root-zone temperature causes similar changes in water relations and growth of silver maples from 33 degrees and 44 degrees N latitude. J. Amer. Soc. Hort. Sci. 122:195–199.

Hartmann, H.T., D.E. Kester, F.T. Davies, Jr., and R.L. Geneve. 1997. Plant propagation, principles and practices. Prentice Hall. Upper Saddle River, N.J.

Huang, H., J.D. Norton, G.E. Boyan, and B.R. Abrahams. 1994. Graft compatibility among Chestnut (Castanea) species. J. Amer. Soc. Hort. Sci. 119:1127–1132.

Hurewitz, J. and H.W. Janes. 1983. Effect of altering the root-zone temperature on growth, translocation, carbon exchange rate, and leaf starch accumulation in the tomato. Plant Physiol. 73:46–50.

Klock, K.A., W.R. Graves, and H.G. Taber. 1996. Growth and phosphorus, zinc, and manganese content of tomato, muskmelon, and honey locust at high root-zone temperatures. J. Plant Nutr. 19:795–806.

Layne, D.R. 1996. The pawpaw [Asimina triloba (L.) Dunal]: A new fruit crop for Kentucky and the United States. HortScience 31:777–784.

Lopez-Cantarero, I., F.A. Lorente, and L. Romero. 1994. Are chlorophylls good indicators of nitrogen and phosphorus levels? J. Plant Nutr. 17:979–990.

Mandujano, M. and D.W. Fulbright. 1998. Meristem dieback of greenhouse-grown pawpaw is associated with soil pH. Annu. Rpt. Northern Nut Growers 89:103–107.

Menzel, D.M., D.W. Turner, V.J. Doogan, and D.R. Simpson. 1994. Root shoot interactions in passionfruit (Passiflora sp.) under the influence of changing root volumes and soil temperatures. J. Hort. Sci. 69:552–564.

Moran, R. 1982. Formulae for the determination of chlorophyllous pigments extracted with N,N-dimethylformamide. Plant Physiol. 69:1376–1381.

Moss, G.I. 1984. The effects of root-zone warming on the yield and quality of roses grown in a hydroponic system. J. Hort. Sci. 59:549–558.

Nash, L.J. and W.R. Graves. 1993. Drought and flood stress effects on plant development and leaf water relations of five taxa of trees native to bottomland habitats. J. Amer. Soc. Hort. Sci. 118:845–850.

Pomper, K.W., D.R. Lane, and S.C. Jones. 2002. Incident irradiance and cupric hydroxide container treatment effects on early growth and development of container-grown pawpaw seedlings. J. Amr. Soc. Hort. Sci. 127:13–19.

Ruter, J.M. and D.L. Ingram. 1992. High root-zone temperatures influence RuBisCO activity and pigment accumulation in leaves of ‘Rotundifolia’ holly. J. Amer. Soc. Hort. Sci. 117:154–157.

Sartain, J.B. and D.L. Ingram. 1984. Influence of container medium, lime, and nitrogen source on growth of woody ornamentals. J. Amer. Soc. Hort. Sci. 109:882–886.

Toselli, M., J.A. Flore, B. Marangoni, and A. Masia. 1999. Effects of root-zone temperature on nitrogen accumulation by non-bearing apple trees. J. Hort. Sci. Biotechnol. 74:118–124.

Vogelezang, J.V.M. 1991. Effect of root-zone and air temperature on growth, ornamental value and keepability of Ficus benjamina and Scheffler arboricola 'Compacta'. Sci. Hort. 46:301–313.

Wright, A.N., A.X. Niemiera, J.R. Harris, and R.D. Wright. 1999. Preplant lime and micronutrient amendments to pine bark affect growth of seedlings of nine container-grown tree species. HortScience 34:669–673.

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