Dynamics of graft formation in fruit trees: a review

Headnote

Abstract

This review paper comprehensively discussed the dynamics of graft union formation in fruit trees. Histological studies confirmed graft union is a continuous process and reliable indicators of graft-compatibility are not visible. Although initiation of scion bud development is signs of graft formation processes but early callus formation determines subsequent survival of scion prior to cambial and vascular connectivity. Process of graft union formation is similar in most plants with few variations in days to completion. Factors attributed to graftincompatibility or otherwise have been mentioned whiles research should also target specific tropical plants.

Key words: Graft incompatibility, Graft union formation, Callus formation, Cambium, Vascular connectivity

1. Introduction

Grafting is defined as the natural or deliberate fusion of plant parts so that vascular continuity is established and functions as a single plant [22]. However, to achieve the beneficial effects of grafting, the plant has to undergo external and internal processes to establish communication between rootstock and scion [25]. Graft success or failure determination may be difficult at one time because graft-failure could increase gradually with ageing of grafted plants. Therefore, in-depth information on graft-union behavior supports better handling of plants and plant environment for success. This paper comprehensively discussed the dynamics of graft union formation in fruit trees which take rather a longer time to examine and also influenced by abiotic conditions. The paper also highlights some areas of future research advances. In this presentation the following terms will be used interchangeably; graftsuccess or compatibility and graft-failure or incompatibility.

2. Graft union formation

Graft union formation is a process that establishes the connection between rootstock and scion that may eventually determine their graftcompatibility or incompatibility [12]. Complete union formation is the final and most reliable determinant of graft-compatibility particularly in tree crop that are difficult to 'take'. With respect to this presumption, there is high degree of similarity in the pattern of the union process in different fruit tree species including Mangifera indica [1, 7, 4, 24] and Malus domestica [28]. The stages of union formation include the death of layers of cells at the graft interface, cohesion of scion and rootstock, generation of callus cells, differentiation of callus and establishment of vascular continuity [1]. Copes [5] also indicated the order of union formation; contact or isolation layer formation, cell enlargement, callus formation, phellogen formation and vascular cambium formation. Soule [27] categorized these stages into four namely precallus, callus, cambial bridge and the healed union, whilst [17] suggested three stages of compatible graft union as cohesion of the rootstock and scion, proliferation of callus and vascular connection across the interface. Essentially, entire union process is the same for all species [7] but interaction between the various stages of the process and subsequent time to completion is plant specific. However, this paper discussed below, three main stages of graft formation; callus formation, cambial formation and vascular connectivity to practically enhance understanding of the entire process.

3. Callus formation

Callus is a mass of soft parenchymatous tissue that is quickly formed on or below the injured surface and the closing of the wound by the formation of a callus is the first stage in the healing after the operation [7]. Celik [3] suggests that the degree of callus formation at the graft union shortly after grafting operation is the main determinant of graftcompatibility. Mendel [13] also indicated that the onset of cell division marks the beginning of callus formation but the exact time of the first division depends on the activity of the tree and on external factors including temperature. According to [11] contact between the cambial regions of the rootstock and scion is capable of producing parenchymatic cells and callus tissues cement the two graft components. Other authors also indicated that the high degree of compatible grafts is due to the high rate of callus proliferation [31]; abundant root development [21]; cohesion of the stock and scion [18]; and vascular connection across the graft union [17]. According to [9] the basis of callus formation depends on protein released from the plasmalemma forming a complex with catalytic activity resulting in the formation of a successful graft.

Observations by [18] noted the presence of callus on the third day in Sedum whilst [30] indicated satisfactory callus by 28 days in Nectarine/Almond. Turkoglu [33] also reported adequate callus production in 14 days samples of Rosa canina I R. centi/olia combination and [2] observed an increased callus production early as three weeks after grafting in nut graft combinations. Unal and Ozcagiran [34] notified large and stable callus formation in 30 days samples of Pyrus communis I Eriobotrya japónica combination. However, [7] reported satisfactory but delayed callus production in 45 days. The presence of the callus initiates rootstock-scion interaction that create bridge or conducting tissues for water and nutrients transport to the scion [1, 14] ; a short fall of moisture in the scion leads to delay or failure of cambial formation [37]. In view of this, early callus formation is vital but influenced by plant type, physiological condition, environmental (temperature and relative humidity) and craftsmanship [11].

4. Cambial formation and vascular connectivity

According to [7] cambial continuity was observed 60 days after grafting with newly formed cambium producing vascular tissues (xylem and phloem). Tekintas and Dolgun [30] confirmed cambial continuity and vascular transformation between 45 and 60 days in nectarin/almond combination. Tekintas [32] also observed cambial continuity in Citrus grafts in 45 days after grafting but 40 days earlier after grafting in loquat/quince-c combination [23]. Copes [5] indicated that initiation of shoot growth by the scion is a good indication that cambium is present and tracheids had differentiated in the union. Cambium maintains vascular connection in the callus bridge and for that reason proper matching of graft partners is important [11]. On the contrary, poorly aligned graft components result in slow cambial formation [5] but severe misalignment may result in complete failure of cambia union [1].

Vascular tissue formation is considered the last stage of the successful grafting beginning after the establishment of cambial continuity and a strong connection may occur in a short time in compatible grafts [35]. Failure to achieve vascular continuity in the union within a period of 4-8 weeks might result in desiccation of the scion and graft failure [36]. Singh [26] gave 2-3 months for complete union formation whilst, [4] suggested healing of the graft joint by 4 months after grafting. The entire anatomical changes discussed above are entirely a continuous process with no definite time limit to the completion of each stage [16]

5. Graft Compatibility and Incompatibility in union formation

Graft-success is defined as the ability of some plant components to form a sufficient graft union that could also be described as "graft-take" or "graftcompatibility". Whiles graft-take could also mean the sprouting of scion few days after grafting operation, and graft-compatibility perhaps is the long-term assessment of grafts into a complete single plant beyond the initial sprouting of scion [11]. There are several external symptoms to detect graftincompatibility; graft union uniformity, lack of lignification, yellowing of foliage, decline in vegetative growth and vigor and anatomical abnormalities [11]. Whiles the appearance of these symptoms could take several years [10]; graftincompatibility is one of the greatest obstacles in breeding rootstocks of fruit trees [6].

Factors affecting graft-incompatibility are multiple but categorized into adverse physiological response between scion and rootstock and/or anatomical abnormalities of vascular system with the later being the major cause [20]. According to [8] graft-compatibility is achieved between homogenetic rather than heterogenetic combination. Güclü and Koyuncu [10] found these two compatible but lignifications finished in homogenetic combination earlier than achieved between heterogenetic combinations. This means that heterogenetic combination is often not absolutely incompatible; hence the causes of this graft-incompatibility remain uncertain. Errea [8] and Mosse [19] suggested that biochemical causes, rather than anatomical ones, are responsible for the graft-incompatibility by altering the cambial continuity. Moore [17] also attributed biochemical causes to toxins but could not ascertain the specific toxins. Rather, [29] identified high indoles content to facilitate callus formation whiles least phenols content decreased cell division and suppressed graft-compatibility. Mng'omba [15] also confirmed the important role of phenol compounds in plants especially in scion-rootstock associations. Güclü and Koyuncu [10] also confirmed peroxidase activity to predicting graft-incompatibility in sweet cherries. This prediction is considered relevant to detect combinations of graft components that might show delayed incompatibility, especially before grafting.

6. Conclusion

In conclusion, an in-depth knowledge of graft union process helps to evaluate performance of graft success or failure. Studies on graft-incompatibility effect in tropical woody plants still limited. Biochemical and molecular mechanisms in incompatibility of tropical fruit trees in particular are lacking. Again, methods for graft-incompatibility prediction are inadequate. Finally, research should explain the statement the fact that diseases' attacking the scion of grafted plant is not strictly graft-incompatibility unless the graft union is affected.

References

7. References

1. Asante AK, Barnett JR: Graft union formation in mango (Mangiferaindica L. ) Journal of horticultural Sciences 1997, 72: 781-90

2. Balta F (1993): Nut Propagation by Grafting and Anatomical and Histological Investigation on Graft Formation. PhD Thesis. 100. Yil Univ. Institute of Natural Science and Technology. Van, Turkey. 1993.

3. Celik H: "The effects of different grafting methods applied by manual grafting units on grafting success in grapevines". Turkish Journal of Agriculture and Forestry 2000, 24: 499-504. 71(5): 752-758.

4. Charkrabarty U, Sadhu MK: Anatomy of graft union in epicotyl grafting of mango. ActaHorticulturael9$$, 231: 182-185

5. Copes D: Graft union formation in Douglas-fir. American Journal of Botany 1969, 56 (3): 285289.

6. Davarynejad GH, Hassanpour H, Aziz M, Shahriaree F: Investigation on the possibility of reducing graft incompatibility in some Iranian pear cultivais on Quince A by interstocks. Agriculture Sciences and Technology Journal. 2007, 21 (1): 45-55

7. Dolgun O, Tekintas FE, Ertan E: A Histological Investigation on Graft Formation of Some Nectarine Cultivars Grafted on Pixy Rootstock. World Journal of Agricultural Sciences 2008, 4 (5): 565-568

8. Errea P: "Implications of phenolic compounds in graft incompatibility in fruit tree species". ScientiaHorticulturael998, 74 (3) 195-205.

9. Gökbayrak ?, Söylemezoglu G, Akkurt M, Çelik H: Determination of grafting compatibility of grapevine with electrophoretic methods. ScientiaHorticulturae 2007, 113: 343-352.

10. Güclü SF, Koyuncu F: A Method for Prediction of Graft Incompatibility in Sweet Cherry / Not Bot HortiAgrobo, 2012, 40(1): 243-246

11. Hartmann HT, Kester DE, Davies FT, Geneve RL: Plant Propagation: principles and practices: (6th edition). Prentice- Hall, Inc. Upper Saddle River, New Jersey, USA. Hort. Rev; 1997, 33: 267-349.

12. Mahunu GK, Adjei PY, Asante AK: Anatomical studies of graft formation in cashew (Ânacardium Occidentale L. ). Agricultural and Biological Journal of North America 2012, 3(4): 150-153.

13. Mendel K: The anatomy and Histology of the bud-union in citrus (contribution from the Division of Horticultural physiology and Genetics, Jewish Agency for Palestine, Agricultural Exp. Station, Rehovot). Palestine Journal of Botany. 1936, 1 (2), 13-46

14. Miller H: The effects of Heat on graft union formation in Dormantsitka spruce Picea sitchensis (Bong, Carr. ) PhD. Thesis, University of Reading. 1991.

15. Mng'omba SA, du Toit ES, Akinnifesi FK: The relationship between graft incompatibility and phenols in UapacakirkianaMüell Arg. Scientiahorticulturae200S, 117, 212-218

16. Moghadam ARL, Ardebili ZO, Rezaie L: Effect of indole butyric acid on micrografting of cactus. African Journal of Biotechnology 2012, 11(24): 6484-6493.

17. Moore R: A model for graft compatibilityincompatibility in higher plants. Amer. J. Bot. 1984, 71(5): 752-758.

18. MooreR, Walker DB: Graft compatibility in plants. Biosciencel9Sl, 31 (5): 389-91.

19. Mosse ?, (1962): Graft incompatibility in fruit trees". Tech. Comm. Com. Bur. Hort. Plant Crops. 1962, 28 and 36.

20. Mudge K, Janick J, Scofield S, Goldschmidt EE: A History of Grafting Horticultural Reviews, 2009, 35 (9) 437-493. Edited by Jules Janick

21. Neves C, Cludia S, Amâncio S: Histochemical detection of H202 by tissue printing as precocious marker of rhizogenesis in grapevine. Plant Physiol. Biochem. 1998, 36: 817-824.

22. Pina ? and Errea P: A review of new advances in mechanism of graft compatibilityincompatibility. Scientia Hort. 2005, 106: 1-11.

23. Polat AA, Kaska N: Determination of budding success in loquats budded on Quince-C rootstock, Bahce, 1992, 21 (1/2): 9-11.

24. Ram S: Propagation of mango in India. Acta Hort. 1997, 11:363 400

25. Shehata SAM, El-Shraiy, Amai M: Regulating Cucumber Grafting by Interactions of Cytokinins in Xylem Exudates of Rootstock and Basipetol Polar Auxin Transport of Scion at Graft Union. Australian Journal of Basic and Applied Sciences, 2010, 4(12): 6179-6184.

26. Singh LB: The Mango botany, cultivation and utilization. World crops books. Leonard Hill, London. 1960.

27. Soulc J: Anatomy of the bud union in mango (Mangiferaindica L. ) Journal American Society of Horticultural Science 1971, 96 (3): 380-383.

28. Soumelidou K, Battey NH, John P, Barnett JR: The anatomy of the developing bud union and its relationship to dwarfing in apple. Ann. Bot. 1994,74:605-611.

29. Stino RG, Ghoneim EI, Marwad ID, Fadl TR: Performance of Summer grafted Superior Seedless Grapes Grafts on Different Rootstocks. Journal of Horticultural Science and Plant Ornamental Plants (2011) 3 (1); 8690

30. Tekintas FE, Dolgun O: An Investigation on Compability in Some Peach and Nectarin Cultivars Grafted on Almond Seedling, 100. Y21 Univ. J. Fac. Agrie. , 1996, 6(1): 51-54.

31. Todic, S. , Beslic, Z, Kuljancic I: Varying degree of grafting compatibility between cv. Chardonnay, Merlot, and different grapevine rootstocks. Journal of Central European Agriculture. 2005, 6(2): 115-120.

32. Tekintas FE: An Anatomical and Histological Investigation on Graft Formation in Citrus Species and Varieties Grafted on Different Rootstocks, 100. Yil Univ. J. Fac. Agrie. , Van. Turkey, 1991, 1(2): 68-81.

33. Turkoglu N: An Investigation on Different Grafting Techniques and Effects of Hormones on Rooting in Rosa Canina in van Ecological Conditions. Master Thesis, 100. Yil Univ. Institute of Science and Technology. Van, Turkey 1990.

34. Unal A, Ozcagiran R: Graft Formation in Budding, Doga. Turkey J. Agrie. Forest. , 1986, 10(3): 399-407

35. Unal A: An Investigation on Anatomical Characteristics of Grafting in Almond, Plum and Apricot Grafted on Apricot Rootstocks. 1. National Horticultural Congress, Izmir, Turkey 1992,41-45.

36. Weatherhead I, Barnett JR: Development and structure of unusual xylem elements during graft union formation in Picea sitchensis. Ann. Bot. 1986,57:593-598.

37. Weatherhead I: Causes of Graft failure in Sitka spruce, Piceasitchensis(Bong). Car Ph. D. Thesis. The University of Reading 1986.