1977) LAM: REGENERATION OF PLANTLETS 575

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1 1977) LAM: REGENERATION OF PLANTLETS 575 REGENERATION OF PLANTLETS FROM SINGLE POTATOES Shue-Lock Lam CELLS IN Abstract Regeneration of plantlets from isolated single cells of potato tubers can be achieved through several culturing procedures, each maintained on defined medium and under specific environmental conditions. High auxin and high temperature were favorable for cell proliferation while high cytokinin and alternating low temperature promoted embryoid formation and plantlet regeneration. Resumen La regeneraci6n de plfintulas a partir de c61ulas aisladas de tub6rculos de papa puede conseguirse a trav6s de varios procedimientos de cultivo, cada uno en un medio definido y bajo condiciones ambientales espec/ficas. Alta auxina y altas temperaturas fueron favorables para la proliferaci6n celular, mientras que alta citokinina y bajas temperaturas alternadas promovieron la formaci6n de embrioydes y la regeneraci6n de plantas. Introduction In recent years, considerable progress has been made with tobacco and other plant species in single cell culture technology involving isolation of single cells or protoplasts, fusion of protoplasts between different plant species, and the regeneration of hybrid plants (2). In 1975, Upadhya reported on successful induction of callus from isolated potato protoplasts but he failed to regenerate plantlet from such callus (10). The objective of the present study was to determine the environmental conditions for the isolation of single cells of potato and the nutrient requirements for the subsequent regeneration of plants. Materials and Methods Two Solanum species, S. tuberosum L. var. Superior and S. demissum, L., were used in the present studies. In general, preparation of nutrient media and manipulation of experimental materials were the same as previously reported (3) except in the case of special procedures which will be described in detail below. Received for publication September 24, 1977, Journal Paper Number 6895 of the Purdue University Agricultural Experiment Station, West Lafayette, Indiana 47907, Department of Horticulture.

2 576 AMERICAN POTATO JOURNAL (Vol. 54 Suspension Culture - Cell suspension was obtained from friable callus originated from tuber discs grown on the basal medium (3) containing 3 ppm of 2,4-D at 27~ with light intensity of 10,800 lux. About I gm of friable callus was inoculated into 30 ml of liquid medium (Table I under "suspension culture") in 125 ml flask. The liquid suspension was agitated continuously at 100 rpm on a gyratory shaker under continuous dim light at a room temperature of 27~ To promote active cell division, the cell suspension was subcultured using the same nutrient medium renewed at every 2-3 week interval. Isolation of Single Cells - Isolation of single cells from cell suspension was usually made about 7-10 days after cell suspension reculture. The suspended cells were first filtered through one layer of cheesecloth in order to remove large clumps of cells and then through 2 layers of cheesecloth. The cell suspension was then poured into a graduate cylinder and the cells allowed to settle. The cell aggregates settled to the bottom and isolated single cells floated in the upper portion of the cylinder. To determine the time required for separation of single cells and cell aggregates at a certain point in the cylinder, a drop of suspended cells was taken out with a micropipette and examined under the microscope. Generally, minutes after settling, about 90% of the suspended cells above the midpoint of the cylinder were single cells. The number of cells counted in 0.01 ml multiplied by 100 was equal to the number of single cells per ml. A hemacytometer was also used for counting cells. Culture of single cells - About 7 ml of.6% of agar nutrient medium (Table 1 under "cell plating") were poured into 5 cm plastic petri dishes After solidification, 1 ml of isolated single cells (cell density was about 105 cells per ml) was pipetted onto the surface of the agar medium in each petri dish. The petri dishes were then sealed with parafilm and placed in an incubator with low light intensity of about 200 lux. Temperature was held at 27~ and 16-hr. photoperiod. About 3-4 weeks later, small calli reached the size of 2-3mm in diameter. The calli were then transferred singly to another petri dish containing the same medium except that the concentration of NAA was reduced from 2 ppm to 0.2 ppm and the percentage of agar was increased from.6% to 1% and 800 mg of NH4NO3 was also added. Shoot regeneration - Calli reaching the size of mm in diameter were transferred to a 250 ml culture bottle containing 50 ml of the agar medium (Table I under "shoot regeneration"). The culture bottles were then placed in a growth chamber with high light intensity of about 10,800 lux; at the alternate temperature of 21~ during the day and 16~ at night. Photoperiod was 16 hr. Under these environmental conditions, small shoots could be observed approximately 5-7 weeks after transferring.

3 1977) LAM: REGENERATION OF PLANTLETS 577 TABLE Composition of nutrient media (rag~l). Suspension Cell Shoot Culture Plating Regeneration Inorganic Salts Major Elements Minor Elements M.S. (without M.S. (without M.S. NH4NO3) NH4NO3) % of M.S. V2 of M.S. M.S. Organic Constituents Organic Addenda N & N N & N N & N Casein Hydrolysate Sucrose 1% I% 1% Glucose 1% 1% 1% D-Manitot 4% 4% 4% Agar 0 0.6% 1% Growth Regulators NAA IAA BAP Zeatin Kinetin GA 2,4-D M.S. (Murashige and Skoog's Medium) N & N (Nitsch and Nitsch Medium) NAA (l-naphthaleneacetic Acid) IAA (Indole -3-Acetic Acid) BAP (6-Benzylaminopurine) GA (Gibberellic Acid) Results and Discussion Ammonium and full strength of minor elements of Murashige and Skoog's medium have been reported to be toxic in cell culture in some cases (1, 6, 12), but not in others (11). In the present study, the revised formulation (Table I under" suspension culture") appeared to be beneficial for potato cell culture. The basal medium containing 2 ppm of 2,4-D produced more single cells than that containing 3 ppm of NAA and I ppm of BAP. It appeared that 2,4-D increased cell separation and cell division while NAA increased cell aggregation. Light intensity appeared to have a significant influence on growth of plated cells in petri dishes. Many more isolated single cells developed into small colonies when cultures were placed under 200 lux than did those under 800 lux of light. Under 5000 lux of light, the growth of potato cells was totally inhibited. Cell division was usually observed 2-3 days after plating (Fig. I). Figure 2 shows small clumps of potato cells derived from isolated single cells 15 days after plating.

4 578 AMERICAN POTATO JOURNAL (Vol. 54 FIG. 1. Cell division occurred 2 days after plating. FIG. 2. Small clump of potato cells 15 days after plating.

5 1977) LAM: REGENERATION OF PLANTLETS 579 It is generally believed that shoot and root initiation are controlled by the auxin and kinetin ratio in the medium (7, 10). However, other factors may be involved (5, 8, 9). Nishi, et al indicated that callus induction (dedifferentiation) and organ redifferentiation from callus were simply controlled by the presence or absence of auxin in the medium and cytokinin was not necessary for organ redifferentiation (9). Previous work indicated that shoot initiation from callused tuber discs was easily achieved by the proper adjustment of the auxin and kinetin ratio. It was also pointed out that in the adjustment of the auxin level in induction ofplantlet formation, the exogenous auxin level in the medium as well as the endogenous auxin of the plant materials must be considered (4). The results of the present study indicate that the frequency of shoot initiation varied from 10% ins. tuberosum to 70% ins. demissum. The low frequency of shoot initiation in S. tuberosum is probably due to the loss of morphogenic ability; the callus used was 12 months old and had been maintained by subcultures. Figure 3 shows plantlet from callus derived from isolated single cells 7 weeks after transferring to the regeneration medium. Successful regeneration of plantlets from isolated single cells in the present study, would make it possible to proceed to a study of somatic hybridization between the potato species for use in potato improvement programs. FIG. 3. Plantlet from callus derived from isolated single cells 7 weeks after transferring to the regeneration medium.

6 580 AMERICAN POTATO JOURNAL (Vol. 54 Literature Cited 1. Engvild, K.C Growth and plating of cell suspension cultures ofdatura innoxia. Physiol Plant 32: Gamborg, O.L. et al Protoplast and cell culture methods in somatic hybridization in higher plants. Can J Genet Cytol 16: Lain, Shue-Lock Shoot formation in potato tuber discs in tissue culture. Am Potato J 52: Lam, Shue-Lock Plantlet formation from potato tuber discs in vitro. Am Potato J 54: McWilliam, A.A., S.M. Smith and H.E. Street The origin and development of embryoids in suspension cultures of carrot (Daucus carota). Ann Bot 38: Meyer, Y Isolation and culture of tobacco mesophyll protoplasts using a saline medium. Protoplasma 81: Murashige, T. and F. Skoog A revised medium for rapid growth and bioassays with tobacco tissue cultures. Physiol Plant 15: Murashige, T. and R.T. Nakano Morphogenetic behaviour of tobacco tissue cultures and implications of plant senescence. Am J Bot 52: Nishi, T., Y. Yamada and E. Takahashi The role of auxins in differentiation office tissue cultured in vitro. Bot Mag (Tokyo) 86: Skoog, F. and C.O. Miller Chemical regulation of growth and organ formation in plant tissues cultured in vitro. Symp Soc Exp Biol 11: Uchimiya, H. and T. Murashige Evaluation of parameter in the isolation of viable protoplasts from cultured tobacco cells. Plant Physiol 54: Upadhya, M Isolation and culture of mesophyll protoplasts of potatoes (Solanum tuberosurn L.) Potato Res 18: Author's Note Just prior to forwarding this manuscript for publication, I discovered that Profs. Shepard and Totten have reported on successful regeneration of plantiets from mesophyh protoplasts in potato in the August issue of Plant Physiology. 60:

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