, Volume 25, Issue 6, pp 1083–1090 | Cite as

Branching of annual shoots in common walnut (Juglans regia L.) as affected by bud production and indol-3-acetic acid (IAA) content

  • Anita SolarEmail author
  • Gregor Osterc
  • Franci Štampar
  • Damijan Kelc
Original Paper


Relationship of bud production (axillary and terminally) of annual shoot (1Y) and/or the content of bud-derived indol-3-acetic acid (IAA) to branching of the 1Y was studied in common walnut (Juglans regia L.), cvs. Franquette and Lara. Cultivar-related branch architecture was determined. Lara tended to branch more densely than Franquette (53 vs. 42%). Significantly more fruiting off-spring shoots (FO) than vegetative ones (VO) grew-out per 1Y in both cultivars, whereas the ratio FO/VO of Lara exceeded that of Franquette by four times. An acrotonic branching pattern was more strongly expressed in Lara compared to Franquette. Bud-derived IAA was influenced by the cultivar (Franquette had 3.6 times more cumulative IAA along the 1Y than Lara), and by the relative position (terminal, subterminal, medial and basal) of the buds along 1Y. An opposite relationship between branching density and cumulative IAA content was established in both cultivars. At the 1Y relative position level, the opposite ratio between branching density and IAA content was clearly shown only on the basal position of the bud along 1Y in the Lara cultivar. Such an inconsistent linkage between bud production and the IAA spatial distribution along the 1Y illustrated that hormonal factors probably weakly affect the branching of Franquette and Lara. The length of the parent 1Y, the position of the buds along the 1Y-length, and the fate of the buds seemed to have a stronger influence on the bud out-growth and further development of the off-springs. In further analyses, seasonal fluctuations of the IAA, and the following activity of the buds should be investigated in order to improve the understanding of a complex branching phenomenon in walnut.


Juglans regia L. Annual shoots Bud activity Bud-derived IAA Branch architecture Current-year off-springs 



This work was supported by the Slovenian Ministry of Higher Education, Science and Technology, and is a part of the programme Horticulture No P4-0013-0481. The authors thank Dr. Evelyne Costes for her valuable comments and two anonymous reviewers for the greatly appreciated suggestions on the manuscript.


  1. Bangerth F (1994) Response of cytokinin concentration in the xylem exudate of bean plants (Phaseolus vulgaris L.) to decapitation and auxin treatment and relationship of apical dominance. Planta 194:439–442CrossRefGoogle Scholar
  2. Bartel B, LeClere S, Migidin M, Zolman BK (2001) Inputs to active indole-3-acetic acid pool: de novo synthesis, conjugate hydrolysis, and indole-3-butyric acid β-oxidation. J Plant Growth Reg 20:198–216CrossRefGoogle Scholar
  3. Barthélémy D, Sabatier S, Pascal O (1995) Le developpement architectural du noyer commun Juglans regia L. (Juglandaceae). Foret-entreprise 103(3–4):61–68Google Scholar
  4. Bell AD (1998) Plant Form. An illustrated guide to flowering plant morphology. Oxford University Press, Oxford, New York, TokioGoogle Scholar
  5. Bennett T, Sieberer T, Willett B, Booker J, Luschnig C, Leyser O (2006) The Arabidopsis MAX pathway controls shoot branching by regulating auxin transport. Curr Biol 16(6):553–563PubMedCrossRefGoogle Scholar
  6. Caraglio Y, Barthélémy D (1997) Revue critique des termes relatifs à la croissance et à la ramification des tiges des végéteaux vasculaires. In: Bouchon J, de Reffye P, Barthélémy D (eds) Modélisation et Simulation de l’Architecture des Végétaux. Science Update INRA Editions, Paris, pp 11–87Google Scholar
  7. Carraro N, Forestan C, Canova S, Traas J, Varotto S (2006) ZmPIN1a and ZmPIN1b encode two novel putative candidates for polar auxin transport and plant architecture determination of maize. Plant Physiol 142:254–264PubMedCrossRefGoogle Scholar
  8. Cilas C, Costes E, Milet J, Legnaté H, Gnagne M, Clément-Demange A (2004) Characterization of branching in two Hevea brasiliensis clones. J Exp Bot 55(399):1045–1051PubMedCrossRefGoogle Scholar
  9. Cline MG (1997) Concepts and terminology of apical dominance. Am J Bot 84:1064–1069PubMedCrossRefGoogle Scholar
  10. Cline MG (2000) Execution of the auxin replacement apical dominance experiment in temperate woody species. Am J Bot 87(2):182–190PubMedCrossRefGoogle Scholar
  11. Costes E, Guédon Y (1997) Modelling the sylleptic branching on one-year-old trunks of apple cultivars. J Am Soc Hort Sci 122:53–62Google Scholar
  12. Costes E, Belouin A, Brouard L, Lelezec M (2004) Development of young pear trees and occurence of first flowering: a varietal comparison. J Hortic Sci Biotechnol 79:67–74Google Scholar
  13. Costes E, Lauri PE, Régnard JL (2006) Tree architecture and production. Hortic Rev 32:1–60Google Scholar
  14. Crawford S, Shinohara N, Sieberer T, Williamson L, George G, Hepworth J, Muller D, Domagalska MA, Leyser O (2010) Strigolactones enhance competition between shoot branches by dampening auxin transport. Development 137(17):2905–2913PubMedCrossRefGoogle Scholar
  15. Ducousso I, Sabatier S, Barthélémy D, Germain E (1995) Comparison de quelques caracteristiques morphologiques des pousses annuelles et des branches de la cime de sept variétés de Noyer commun, Juglans regia L. (JUGLANDACEAE). In: Les Coloques, No. 74. Colloque Architecture des arbres fruitières et forestières. Montpellier, France, 23–25 Nov 1993:91–108Google Scholar
  16. Dun EA, Brewer PB, Beveridge CA (2009) Strigolactones: discovery of the elusive shoot branching hormone. Trends Plant Sci 14(7):364–372PubMedCrossRefGoogle Scholar
  17. Foo E, Bullier E, Goussot M, Foucher F, Rameau C, Beveridge CA (2005) The branching gene RAMOSUS1 mediates interactions among two novel signals and auxin in pea. Plant Cell 17(2):464–474PubMedCrossRefGoogle Scholar
  18. Frimi J (2003) Auxin transport—shaping the plant. Curr Opin Plant Biol 6:7–12CrossRefGoogle Scholar
  19. Germain E (1990) Inheritance of late leafing and lateral bud fruitfulness in walnut (Juglans regia L.), phenotypic correlations among some traits of the trees. Acta Hortic 284:125–134Google Scholar
  20. Germain E, Prunet JP, Garcin A (1999) Le Noyer. CTIFL, Paris, FranceGoogle Scholar
  21. Gomez-Roldan V, Fermas S, Brewer PB, Puech-Pages V, Dun EA, Pillot JP, Letisse F, Matusova R, Danoun S, Portais JC, Bouwmeester H, Becard G, Beveridge CA, Rameau C, Rochange SZ (2008) Strigolactone inhibition of shoot branching. Nature 455:189–194PubMedCrossRefGoogle Scholar
  22. Hallé F, Oldeman RAA, Tomlinson PB (1978) Tropical Trees and forests. An architectural analysis. Springer, Berlin, Heidelberg, New York, pp 43–45Google Scholar
  23. Ito A, Yaegaki H, Hayama H, Kusaba S, Yamaguchi I, Yoshioka H (1999) Bending shoots stimulates flowering and influences hormone levels in lateral buds of Japanese pear. Hortic Sci 34:1224–1228Google Scholar
  24. Kelc D, Štampar F, Solar A (2007) Fruiting behaviour of walnut tree influences relationships between morphometric traits of parent wood and nut weight. J Hortic Sci Biotechnol 82(3):439–445Google Scholar
  25. Kelc D, Štampar F, Solar A (2010) Distribution of indol-3-acetic acid (IAA) in the annual fruit-bearing shoots of walnut cultivars ‘Franquette’ and ‘Lara’. Acta Hortic 861:273–278Google Scholar
  26. Klee HJ, Lanahan MB (1995) Transgenic plants in plant biology. In: Davies PJ (ed) Plant hormones: physiology, biochemistry, molecular biology, 2nd edn. Kluwer Academic, Dordrecht, The Netherlands, pp 340–353Google Scholar
  27. Kovač M, Piskernik D, Ravnikar M (2003) Jasmonic acid induced morphological changes are reflected in auxin metabolism of beans grown in vitro. Biol Plant 47:273–275Google Scholar
  28. Langrova V, Sladky Z (1971) The role of growth regulators in the differentiation of walnut buds. Biol Plant 13(5/8):361–367CrossRefGoogle Scholar
  29. Lauri PE (2007) Differentiation and growth traits associated with acrotony in the apple tree (Malus x domestica, Rosaceae). Am J Bot 94(8):1273–1281PubMedCrossRefGoogle Scholar
  30. Lauri PE (2009) Does plant architecture only result from growing meristems? Atlan’s principle of life and death as regulated morphogenetic processes. In: Karam WE (ed) Tree growth: influences, layers and types. Nova Science Publishers, Hauppauge, New York, pp 45–55Google Scholar
  31. Lauri PE, Terouanne E, Lespinasse JM, Regnard JL, Kelner JJ (1995) Genotypic differences in the axillary bud growth and fruitting pattern of apple fruiting branches over several years—an approach to regulation of fruit bearing. Sci Hortic 64:265–281CrossRefGoogle Scholar
  32. Lauri PE, Bourdel G, Trottier C, Cochard H (2008) Apple shoot architecture: evidence for strong variability of bud size and composition and hydraulics within a branching zone. New Phytol 178:798–807PubMedCrossRefGoogle Scholar
  33. Lespinasse JM, Delort F (1986) Apple tree management in vertical axis: appraisal after ten years of experiments. Acta Hortic 160:120–155Google Scholar
  34. Leyser O (2009) The control of shoot branching: an example of plant hormone processing. Plant Cell Environ 32(6):694–703PubMedCrossRefGoogle Scholar
  35. McGranahan G, Leslie C (1991) Walnuts (Juglans). In: Moore JN, Ballington JR Jr (eds) Genetic resources of temperate fruit and nut crops, vol 2, pp 907–951Google Scholar
  36. Moore R, Clark WD, Vodopich DS, Stern KR, Lewis R (1998) Botany. Second Edition. McGraw-Hill, Boston, pp 413–441Google Scholar
  37. Muradoglu F, Balta F, Battal P (2009) Endogenous hormone levels in bearing and non-bearing shoots of walnut (Juglans regia L.) and their mutual relationships. Acta Physiol Plantar 32(1):53–57CrossRefGoogle Scholar
  38. Rameau C (2010) Strigolactones, a novel class of plant hormone controlling shoot branching. C.R. Biol 333(4):344–349PubMedCrossRefGoogle Scholar
  39. Renton M, Guedon Y, Godin C, Costes E (2006) Similarities and gradients in growth unit branching patterns during ontogeny in ‘Fuji’ apple trees: a stochastic approach. J Exp Bot 57:3131–3143PubMedCrossRefGoogle Scholar
  40. Sabatier SA, Barthélémy D (2001a) Annual shoot morphology and architecture in Persian walnut, Juglans regia L. (Juglandaceae). Acta Hortic 544:255–264Google Scholar
  41. Sabatier SA, Barthélémy D (2001b) Bud structure in relation to shoot morphology and position on the vegetative annual shoots of Juglans regia L. (Juglandaceae). Ann Bot 87:117–123CrossRefGoogle Scholar
  42. Shimizu-Sato S, Mori H (2001) Control of outgrowth and dormancy in axillary buds. Plant Physiol 124(4):1405–1413CrossRefGoogle Scholar
  43. Shimizu-Sato S, Tanaka M, Mori H (2009) Auxin-cytokinin interactions in the control of shoot branching. Plant Mol Biol 69(4):429–435PubMedCrossRefGoogle Scholar
  44. Solar A, Štampar F (2003) Genotypic differences in branching pattern and fruiting habit in common walnut (Juglans regia L.). Annal Bot 92:317–325CrossRefGoogle Scholar
  45. Solar A, Solar M, Štampar F (2006) Stability of the annual shoot diameter in Persian walnut: a case study of different morphotypes and years. Trees Struct Funct 20:449–459CrossRefGoogle Scholar
  46. Souza BM, Kraus JE, Endres L, Mercier H (2003) Relationships between endogenous hormonal levels and axillary bud development of Ananas comosus nodal segments. Plant Physiol Biochem 41:733–739CrossRefGoogle Scholar
  47. Štefančič M, Štampar F, Osterc G (2006) Influence of endogenous IAA levels and exogenous IBA on rooting and quality of leafy cuttings of Prunus ‘GiSelA 5’. J Hortic Sci Biotechnol 81:508–512Google Scholar
  48. Tamas IA (1995) Hormonal regulation of apical dominance. In: Davies PJ (ed) Plant hormones: physiology, biochemistry, molecular biology, 2nd edn. Kluwer Academic, Dordrechet, The Netherlands, pp 572–597Google Scholar
  49. Tworkoski T, Miller S (2007) Endogenous hormone concentrations and bud-break response to exogenous benzyl adenine in shoots of apple trees with two growth habits grown on three rootstocks. J Hortic Sci Biotechnol 82:960–966Google Scholar
  50. Umehara M, Hanada A, Yoshida S, Akiyama K, Arite T, Takeda-Kamiya N, Magome H, Kamiya Y, Shirasu K, Yoneyama K, Kyozuka J, Yamaguchi S (2008) Inhibition of shoot branching by new terpenoid plant hormones. Nature 455:195–200PubMedCrossRefGoogle Scholar
  51. Xie X, Yoneyama Ka, Yoneyama Ko (2010) The strigolactone story. Annu Rev Phytopathol 48:93–117PubMedCrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2011

Authors and Affiliations

  • Anita Solar
    • 1
    Email author
  • Gregor Osterc
    • 2
  • Franci Štampar
    • 2
  • Damijan Kelc
    • 2
  1. 1.Department of Agronomy, Experimental Field for Nut Crops, Biotechnical FacultyUniversity of LjubljanaMariborSlovenia
  2. 2.Department of Agronomy, Biotechnical FacultyUniversity of LjubljanaLjubljanaSlovenia

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