Journal of Plant Growth Regulation

, Volume 30, Issue 1, pp 100–113 | Cite as

Auxins or Sugars: What Makes the Difference in the Adventitious Rooting of Stored Carnation Cuttings?

  • María Ángeles Agulló-Antón
  • José Sánchez-Bravo
  • Manuel Acosta
  • Uwe DruegeEmail author


Cold storage of cuttings is frequently applied in the vegetative propagation of ornamental plants. Dianthus caryophyllus was used to study the limiting influences of auxin and sugars on adventitious root formation (ARF) in cuttings stored at 5°C. Carbohydrate levels during storage were modulated by exposing cuttings to low light or darkness. The resulting cuttings were treated (or not) with auxin and planted, and then ARF was evaluated. Carbohydrate levels in the cuttings were monitored and the influence of light treatment on indole-3-acetic acid (IAA) and zeatin (Z) in the basal stem was investigated. Dark storage for up to 4 weeks increased the percentage of early rooted cuttings and the final number and length of adventitious roots, despite decreased sugar levels in the stem base. Light during cold storage greatly enhanced sugar levels, particularly in the stem base where the Z/IAA ratio was higher and ARF was lower than observed in the corresponding dark-stored cuttings. Sugar levels in nonstored and dark-stored cuttings increased during the rooting period, and auxin application enhanced the accumulation of sugars in the stem base of nonstored cuttings. Auxin stimulated ARF most strongly in nonstored, less so in light-stored, and only marginally in dark-stored cuttings. A model of auxin-sugar interactions in ARF in carnation is proposed: cold storage brings forward root induction and sink establishment, both of which are promoted by the accumulation of auxin but not of sugars, whereas high levels of sugars and probably also of cytokinins act as inhibitors. Subsequent root differentiation and growth depend on current photosynthesis.


Root development Light Dark exposure Temperature Carbohydrates Source-sink Plant hormones Signaling 



The work was supported by the Ministry of Science and Innovation of Spain (projects MICINN/FEDER AGL-2008-02472 and AGL-2004-07902); the Federal Ministry of Food, Agriculture and Consumer Protection of Germany; the State Ministry of Rural Development, Environment and Consumer Protection of Brandenburg; and the Ministry for Agriculture, Nature and Environment Protection of the Free State of Thüringen. M. A. Agulló thanks the Spanish MICINN for a FPI fellowship and a travel grant. We appreciate the skillful technical assistance of Baerbel Broszies and Sabine Czekalla (IGZ, Erfurt) and the valuable help of Dr. A. Torrecillas (CAID, Universidad de Murcia) with hormonal analyses. The authors are grateful to Drs. G. Garrido and E. Cano (Barberet & Blanc S.A., Murcia, Spain) for the plant material supplies.


  1. Acosta M, Oliveros-Valenzuela MR, Nicolás C, Sánchez-Bravo J (2009) Rooting of carnation cuttings. The auxin signal. Plant Signaling Behav 4:234–236CrossRefGoogle Scholar
  2. Ahkami AH, Lischewski S, Haensch KT, Porfirova S, Hofmann J, Rolletschek H, Melzer M, Franken P, Hause B, Druege U, Hajirezaei MR (2009) Molecular physiology of adventitious root formation in Petunia hybrida cuttings: involvement of wound response and primary metabolism. New Phytol 181:613–625CrossRefPubMedGoogle Scholar
  3. Albacete A, Ghanem ME, Martinez-Andujar C, Acosta M, Sanchez-Bravo J, Martinez V, Lutts S, Dodd IC, Perez-Alfocea F (2008) Hormonal changes in relation to biomass partitioning and shoot growth impairment in salinized tomato (Solanum lycopersicum L.) plants. J Exp Bot 59:4119–4131CrossRefPubMedGoogle Scholar
  4. Altman A, Wareing PF (1975) The effect of IAA on sugar accumulation and basipetal transport of 14C-labelled assimilates in relation to root formation in Phaseolus vulgaris cuttings. Physiol Plant 33:32–38CrossRefGoogle Scholar
  5. Blakesley D (1994) Auxin metabolism and adventitious root initiation. In: Davis TD, Haissig BE (eds) Biology of adventitious root formation. Plenum Press, New York, pp 143–154Google Scholar
  6. Calamar A, de Klerk GJ (2002) Effect of sucrose on adventitious root regeneration in apple. Plant Cell Tissue Organ Cult 70:207–212CrossRefGoogle Scholar
  7. Corrêa LR, Paim DC, Schwambach J, Fett-Neto AG (2005) Carbohydrates as regulatory factors on the rooting of Eucalyptus saligna Smith and Eucalyptus globulus Labill. Plant Growth Regul 45:63–73CrossRefGoogle Scholar
  8. de Klerk GJ, Van der Krieken W, de Jong JC (1999) The formation of adventitious roots: new concepts, new possibilities. In Vitro Cell Dev-Pl 35:189–199CrossRefGoogle Scholar
  9. Dennis DT, Blakeley SD (2001) Carbohydrate metabolism. In: Buchanan BB, Gruissem W, Jones RL (eds) Biochemistry & molecular biology of plants. American Society of Plant Physiology, Rockville, MD, pp 630–675Google Scholar
  10. Druege U, Zerche S, Kadner R, Ernst M (2000) Relation between nitrogen status, carbohydrate distribution and subsequent rooting of Chrysanthemum cuttings as affected by pre-harvest nitrogen supply and cold-storage. Ann Bot 85:687–701CrossRefGoogle Scholar
  11. Druege U, Zerche S, Kadner R (2004) Nitrogen- and storage-affected carbohydrate partitioning in high-light-adapted Pelargonium cuttings in relation to survival and adventitious root formation under low light. Ann Bot 94:831–842CrossRefPubMedGoogle Scholar
  12. Eliasson L (1978) Effects of nutrients and light on growth and root formation in Pisum sativum cuttings. Physiol Plant 43:13–18CrossRefGoogle Scholar
  13. Fett-Neto AG, Fett JP, Goulart LWV, Pasquali G, Termignon RR, Ferreira AG (2001) Distinct effects of auxin and light on adventitious root development in Eucalyptus saligna and Eucalyptus globulus. Tree Physiol 21:457–464PubMedGoogle Scholar
  14. Garrido G, Cano EA, Arnao MB, Acosta M, Sánchez-Bravo J (1996) Influence of cold storage period and auxin treatment on the subsequent rooting of carnation cuttings. Sci Hortic 65:73–84CrossRefGoogle Scholar
  15. Garrido G, Cano EA, Acosta M, Sánchez-Bravo J (1998) Formation and growth of roots in carnation cuttings: influence of cold storage period and auxin treatment. Sci Hortic 74:219–231CrossRefGoogle Scholar
  16. Garrido G, Guerrero JR, Cano EA, Acosta M, Sánchez-Bravo J (2002) Origin and basipetal transport of the IAA responsible for rooting of carnation cuttings. Physiol Plant 114:303–312CrossRefPubMedGoogle Scholar
  17. Garrido G, Arnao MB, Acosta M, Sánchez-Bravo J (2003) Polar transport of indole-3-acetic acid in relation to rooting in carnation cuttings: influence of cold storage duration and cultivar. Biol Plant 47:481–485CrossRefGoogle Scholar
  18. Geiger DR, Koch KE, Shieh WJ (1996) Effect of environmental factors on whole plant assimilate partitioning and associated gene expression. J Exp Bot 47:1229–1238CrossRefPubMedGoogle Scholar
  19. Guo J, Hu X (2008) Noninvasive expressions of ipt in whole plants or roots through pOp/LhG4 indicate a role of plant aerial parts and light in cytokinin synthesis and root inhibition. J Plant Growth Regul 27:251–262CrossRefGoogle Scholar
  20. Haissig BE (1974) Metabolism during adventitious root primordium initiation and development. New Zeal J For Sci 4:324–337Google Scholar
  21. Haissig BE (1984) Carbohydrate accumulation and partitioning in Pinus banksiana seedlings and seedling cuttings. Physiol Plant 61:13–19CrossRefGoogle Scholar
  22. Haissig BE (1986) Metabolic processes in adventitious rooting of cuttings. In: Jackson MB (ed) New root formation in plants and cuttings. Martinus Nijhoff Publishers, Dordrecht, pp 141–189Google Scholar
  23. Hajirezaei MR, Takahata Y, Trethewey RN, Willmitzer L, Sonnewald U (2000) Impact of elevated cytosolic and apoplastic invertase activity on carbon metabolism during potato tuber development. J Exp Bot 51:439–445CrossRefPubMedGoogle Scholar
  24. Hansen J, Eriksen EN (1974) Root formation of pea cuttings in relation to irradiance of stock plants. Physiol Plant 32:170–173CrossRefGoogle Scholar
  25. Hartmann HT, Kester DE, Davies FT, Geneve RL (2002) Plant propagation. Principles and practices. Englewood Cliffs, NJ: Prentice Hall, 880 ppGoogle Scholar
  26. Ichimura K, Kohata K, Koketsu M, Shimamura M, Ito A (1998) Identification of pinitol as a main sugar constituent and changes in its content during flower bud development in carnation (Dianthus caryophyllus L.). J Plant Physiol 152:363–367Google Scholar
  27. Jarvis BC (1986) Endogenous control of adventitious rooting in non-woody cuttings. In: Jackson MB (ed) New root formation in plants and cuttings. Martinus Nijhoff Publishers, Dordrecht, pp 191–222Google Scholar
  28. Karve A, Moore BD (2009) Function of Arabidopsis hexokinase-like 1 as a negative regulator of plant growth. J Exp Bot 60:4137–4149CrossRefPubMedGoogle Scholar
  29. Kubota C, Rajapakse NC, Young RE (1997) Carbohydrate status and transplant quality of micropropagated broccoli plantlets stored under different light environments. Postharvest Biol Technol 12:165–173CrossRefGoogle Scholar
  30. Li MS, Leung DWM (2000) Starch accumulation is associated with adventitious root formation in hypocotyl cuttings of Pinus radiata. J Plant Growth Regul 19:423–428Google Scholar
  31. Loewus FA, Murthy PPN (2000) myo-Inositol metabolism in plants. Plant Sci 150:1–19CrossRefGoogle Scholar
  32. Ludwig-Müller J (2009) Molecular basis for the role of auxins in adventitious rooting. In: Niemi K, Scagel C (eds) Adventitious root formation of forest trees and horticultural plants—from genes to applications. Research Signpost, Kerala, India, pp 1–29Google Scholar
  33. Ludwig-Müller J, Vertocnik A, Town CD (2005) Analysis of indole-3-butyric acid-induced adventitious root formation on Arabidopsis stem segments. J Exp Bot 56:2095–2105CrossRefPubMedGoogle Scholar
  34. Mandadi KK, Misra A, Ren S, McKnight TD (2009) BT2, a BTB protein, mediates multiple responses to nutrients, stresses, and hormones in Arabidopsis. Plant Physiol 150:1930–1939CrossRefPubMedGoogle Scholar
  35. Mishra BS, Singh M, Aggrawal P, Laxmi A (2009) Glucose and auxin signaling interaction in controlling Arabidopsis thaliana seedlings root growth and development. PLoS ONE 4(2):e4502CrossRefPubMedGoogle Scholar
  36. Moubayidin L, Di Mambro R, Sabatini S (2009) Cytokinin–auxin crosstalk. Trends Plant Sci 14:557–562CrossRefPubMedGoogle Scholar
  37. Muller B, Sheen J (2008) Cytokinin and auxin interaction in root stem-cell specification during early embryogenesis. Nature 453:1094–1097CrossRefPubMedGoogle Scholar
  38. Nanda KK, Jain MK (1972) Mode of action of IAA and GA3 on root and shoot growth of epiphyllous buds of Bryophyllum tubiflorum. J Exp Bot 23:980–986CrossRefGoogle Scholar
  39. Naqvi SM, Gordon SA (1967) Auxin transport in Zea mays coleoptiles. II. Influence of light on the transport of indoleacetic acid-2–14C. Plant Physiol 42:138–143CrossRefPubMedGoogle Scholar
  40. Normanly J, Slovin JP, Cohen JD (2004) B1. Auxin biosynthesis and metabolism. In: Davies PJ (ed) Plant hormones. Biosynthesis, signal transduction, action. Kluwer Academic Publishers, Dordrecht, pp 36–62Google Scholar
  41. Okoro OO, Grace J (1976) The physiology of rooting populus cuttings. I. Carbohydrates and photosynthesis. Physiol Plant 36:133–138CrossRefGoogle Scholar
  42. Oliveros-Valenzuela MR, Reyes D, Sánchez-Bravo J, Acosta M, Nicolás C (2008) Isolation and characterization of a cDNA clone encoding an auxin influx carrier in carnation cuttings. Expression in different organs and cultivars and its relationship with cold storage. Plant Physiol Biochem 46:1071–1076CrossRefGoogle Scholar
  43. Osterc G, Štefančič M, Štampar F (2009) Juvenile stockplant material enhances root development through higher endogenous auxin level. Acta Physiol Plant 31:899–903CrossRefGoogle Scholar
  44. Perilli S, Moubayidin L, Sabatini S (2010) The molecular basis of cytokinin function. Curr Opin Plant Biol 13:21–26CrossRefPubMedGoogle Scholar
  45. Ramírez-Carvajal GA, Morse AM, Dervinis C, Davis JM (2009) The cytokinin type-B response regulator PtRR13 is a negative regulator of adventitious root development in Populus. Plant Phys 150:759–771CrossRefGoogle Scholar
  46. Rapaka VK, Bessler B, Schreiner M, Druege U (2005) Interplay between initial carbohydrate availability, current photosynthesis, and adventitious root formation in Pelargonium cuttings. Plant Sci 168:1547–1560CrossRefGoogle Scholar
  47. Roitsch T (1999) Source-sink regulation by sugar and stress. Curr Opin Plant Biol 2:198–206CrossRefPubMedGoogle Scholar
  48. Roitsch T, Balibrea ME, Hofmann M, Proels R, Sinha AK (2003) Extracellular invertase: key metabolic enzyme and PR protein. J Exp Bot 54:513–524CrossRefPubMedGoogle Scholar
  49. Smith AM, Zeeman SC, Smith SM (2005) Starch degradation. Annu Rev Plant Biol 56:73–98CrossRefPubMedGoogle Scholar
  50. Sorce C, Picciarelli P, Calistri G, Lercari B, Ceccarelli N (2008) The involvement of indole-3-acetic acid in the control of stem elongation in dark- and light-grown pea (Pisum sativum) seedlings. J Plant Physiol 165:482–489CrossRefPubMedGoogle Scholar
  51. Takahashi F, Sato-Nara K, Kobayashi K, Suzuki M, Suzuki H (2003) Sugar-induced adventitious roots in Arabidopsis seedlings. J Plant Res 116:83–91PubMedGoogle Scholar
  52. Toledo MEA, Ueda Y, Imahori Y, Ayaki M (2003) L-ascorbic acid metabolism in spinach (Spinacia oleracea L.) during postharvest storage in light and dark. Postharvest Biol Technol 28:47–57CrossRefGoogle Scholar
  53. Trethewey RN, Geigenberger P, Riedel K, Hajirezaei MR, Sonnewald U, Stitt M, Riesmeier JW, Willmitzer L (1998) Combined expression of glucokinase and invertase in potato tubers leads to a dramatic reduction in starch accumulation and a stimulation of glycolysis. Plant J 15:109–118CrossRefPubMedGoogle Scholar
  54. Veierskov B (1988) Relations between carbohydrates and adventitious root formation. In: Davis TD, Haissig BE, Sankhla N (eds) Adventitious root formation in cuttings. Dioscorides Press, Portland, OR, pp 70–101Google Scholar
  55. Veierskov B, Andersen AS (1982) Dynamics of extractable carbohydrates in Pisum sativum. III. The effect of IAA and temperature on content and translocation of carbohydrates in pea cuttings during rooting. Physiol Plant 55:179–182CrossRefGoogle Scholar
  56. Werner T, Schmülling T (2009) Cytokinin action in plant development. Curr Opin Plant Biol 12:527–538CrossRefPubMedGoogle Scholar
  57. Werner T, Holst K, Pors Y, Guivarc’h A, Mustroph A, Chriqui D, Grimm B, Schmülling T (2008) Cytokinin deficiency causes distinct changes of sink and source parameters in tobacco shoots and roots. J Exp Bot 59:2659–2672CrossRefPubMedGoogle Scholar
  58. Zelená E (2000) The effect of light on metabolism of IAA in maize seedlings. Plant Growth Regul 30:23–29CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media, LLC 2010

Authors and Affiliations

  • María Ángeles Agulló-Antón
    • 1
  • José Sánchez-Bravo
    • 1
  • Manuel Acosta
    • 1
  • Uwe Druege
    • 2
    Email author
  1. 1.Departamento de Biología Vegetal (Fisiología Vegetal), Facultad de BiologíaUniversidad de MurciaEspinardoSpain
  2. 2.Department Plant PropagationLeibniz-Institute of Vegetable and Ornamental Crops (IGZ)Erfurt-KuehnhausenGermany

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