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Advances in research and development of mango industry/Avancos na pesquisa e desenvolvimento da industria de manga.

WORLD MANGO STATISTICS

Globally the mango industry is the 5th largest tropical fruit industry with production of over 34.3 million tons (FAO 2008 Statistics-check 2009 stats). Production is based across approximately 100 countries in tropical and subtropical regions of the world, with Asia the largest production region with 25,95 5 x [10.sup.3] tons followed by Africa with 3,389 x [10.sup.3] tons, North and Central America with 2,929 x [10.sup.3] tons and South America 2,008 x [10.sup.3] tons (GALAN SAUCO, 2010). Production has been increasing with a doubling of production over the past 18 years, a trend that is expected to continue with current expansion in Brazil, Ecuador, Peru, China West Africa and Spain. Although about 100 countries grow mangoes, about 80% of production comes from the top nine countries in order of production, India, China, Indonesia, Mexico, Thailand, Pakistan Brazil, Philippines and Nigeria (GALAN SAUCO, 2010). Approximately 1% to 2% of the world production is traded internationally the rest is traded and consumed within the countries of production.

The main exporting producing nations include Mexico (23% of production), Brazil (14.3%), Pakistan (3.2%), Peru (10.3%) and India (9.71%). The largest importing destinations are the European community (34%), USA (20%), Arabian Peninsula (14%) and Asia (27%) (GERBAUD, 2009). A key factor in international mango trade is fruit quality at the consumer end of the chain. Factors such as a fruit shelf life, ripening and controlled temperature storage facilities available at the export destination and good agricultural practice certification are significant limitations to this trade.

BREEDING AND GENETIC IMPROVEMENT

Some of the most significant advances in mango research and development that have the capacity to improve the industry and consumer acceptance of mangoes are occurring in the discipline of breeding and genetic improvement. Many of the mango industries in the larger producing nations are based on traditional mango cultivars that have been grown for hundreds of years. Often these cultivars are not up to the challenges of modern international trade, supply chain and consumer demands. As a result, most of the repeat international trade is confined to a few cultivars, often those developed in Florida in the first half of the 20th centaury such as 'Tommy Atkins', 'Kent', 'Keitt'. Although widely adapted these cultivars have a narrow genetic base (SCHNELL; BROWN et al., 2006) and fruit lower fruit quality than the traditional cultivars. With these limitations recognised by many growing and exporting nations several breeding programs have been established to develop new cultivars to improve their export fruit.

Breeding tropical tree fruits such as mango is a long term endeavour taking up to 20 years from crossing to commercial uptake of a new cultivar. Some of the new cultivars coming out of these breeding programs that are beginning to gain market share include: 'Calypso' (TM), 'Honey Gold' A, R2R2, from Australia, 'Maha Chanok' from Thailand, and 'Alpha" among others from Brazil. These and others are listed in Table 1.

It will be these new genetics that drive world trade in mango in the future due to their superior adaption to production conditions, improved consumer appeal and legal proprietary protection through plant variety rights and associated patents and trademarks. One example of this success is the 'Calypso' ([TM]) where the value of the benefit derived from the investment in breeding the cultivar has a benefit cost ratio of 9.5:1, ten years after its release and marketing on the Australian domestic market (StrahAN; PRATT, 2011).

MARKER BASED DIVERSITY AND SELECTION

Molecular genomics technologies are being used to assist our understanding of mango genetics and the diversity of the world mango genepoole. In recent years many studies using molecular marker to discover the diversity and phylogenetic relationships between mango varieties and related Mangifera species have been undertaken. The genetic relationships and origins of individual mangoes and groups of varieties is providing a good understanding of how mangoes have spread around the tropical and subtropical regions of the world. Some of these early analyses used protein or isozyme markers that are influenced by environment and have low polymorphism (SMITH; SMITH, 1992). More recently the more reliable DNA markers generated by PCR-based methods that are reproducible and fast (Varshney, Graner et al., 2005) have been used. Markers such as restriction fragment length polymorphisms (RFLPs), random amplification of polymorphic DNAs (RAPDs), amplified fragment length polymorphisms (AFLPs), inter simple sequence repeat (ISSR) markers and microsatellites or SSRs have been used in mango. SSRs have become the preferred marker for plant analysis because of their reproducibility, multiallelic nature, co-dominant inheritance, relative abundance and good genome coverage (Phillips e Vasil, 2001). In mango SSR marker analyses has revealed genetic relationship over a wide cross-section of the mango and Mangifera genepoole through a series of studies undertaken by (EIADTHONG; YONEMORI et al., 1999; DUVAL; BUNEL et al., 2005; HONSHO; NISHIYAMA et al., 2005; SCHNELL; OLANO et al., 2005; VIRUEL; ESCRIBANO et al., 2005; UKOSKIT, 2007; GALVEZ-LOPEZ; HERNANDEZ-DELGADO et al., 2009). Studies on the Phylogenetic relationships among Mangifera L. species (EIADTHONG; YONEMORI et al., 2000; YONEMORI; HONSHO et al., 2002; HIDAYAT; PANCORO et al., 2011) are revealing some large diversity within species and new and unrecognised relationships between species previously not revealed in traditional taxonomic models of the genus.

Molecular markers have also been used to identify of fingerprint individual mango varieties by unique patterns of marker alleles. Fingerprinting data has been used in support of plant breeders rights and patients (IP AUSTRALIA, 2008) and in maker assisted selection (MAS). MAS is being used routinely in the Australian and Japanese mango breeding programs to identify unknown pollen parents in hybrid progeny lines (Dillon, N., Bally, I. S. E. et al., 2010) and to determine the self- and cross pollination origins of progeny (Kanzaki, Honsho et al., 2005). Fingerprinting of some of the major mango collections around the world has identified duplicated, misnamed and otherwise mislabelled individuals within the collections (SCHNELL, 2006; DILLON, N. L., BALLY, I. S. E. et al., 2010).

The improved understanding of mango genetics and adoption of molecular genetics in breeding programs is enabling breeders to greatly improve their breeding efficiency through improved selection of parents and progeny. This along with research in mango genomics and gene discovery will identify genes and markers associated with traits of interest and advance the quality of the next generation of cultivars released from breeding programs.

ROOTSTOCKS

In mango both polyembryonic and monoembryonic rootstocks are used with only a few rootstock cultivars identified as conferring beneficial effects on scions. In recent times more of an effort has been put in to discovering and evaluation rootstocks for mango that deliver environmental adaptability, uniformity, vigour control or yield enhancement to scions. Some of the specific rootstock cultivars that have been shown to advantage certain scions include, the three root-rot resistant rootstocks IAC 101, IAC104, and IAC106 (ROSSETTO, BORTOLETTO et al., 2006). Rootstocks 'Brodie' and 'MYP' that increased the yield efficiency of 'Kensington Pride' Smith et al (2003). Rootstock evaluation and selection is ongoing in many countries and is likely to provide specific rootstock/ scion combination that will enhance the production and quality performance of specific scions and help to even out annual fluctuations in production and quality.

MANGO DISEASES

Mangoes are susceptible to a range of pest and diseases that can significantly reduce commercial production and fruit quality. One of the most significant diseases of mango is anthracnose caused by the fungal pathogen Colletotrichum gloeosporioides. Anthracnose affects the flowers, leaves and fruit at various stages of growth and is a major cause of fruit rots in the postharvest supply chain. Despite the development of chemical and cultural control measures for this disease, significant losses to production during harvest are still a reality, especially in years where wet wether is experienced. Over recent years several areas of pathogen/host interactions have been studied that may lead to new management options for the disease. These include: investigations of natural defence mechanisms with in the mango and identification genotypes with high levels of natural resistance to the disease. In many plants pre-formed natural anti-fungal compounds act as protective chemical barriers against a range of pathogen infections when found in high concentrations.

In mango high concentrations of the antifungal compounds (alk(en)ylresorcinols) are found the in the peel and in resin ducts of developing fruit. The decline in concentration of alk(en) ylresorcinols during fruit ripening have been significantly correlated with the development of anthracnose symptoms (Hassan, Dann et al., 2007; Hassan, Irving et al., 2009). Alk(en)ylresorcinols concentrations have been shown to vary between mango cultivars (HASSAN; DANN et al., 2007) and can be influenced by tree nitrogen nutrition (BALLY, 2006; BALLY; HOFMAN et al., 2009). Reduced incidence and severity of postharvest anthracnose has been demonstrated in the cultivar 'Kensington Pride' when anti-fungal compounds are retained in the fruit skin during ripening (HASSAN; DANN et al., 2011). This was achieved by leaving long (2-3 cm) peduncles attached to the fruit during harvest which prevents sap loss and maintains anti-fungal concentrations in the peel and in resin ducts.

In Australia a screening program to identify mango cultivars with natural resistance to anthracnose has identified several accessions with superior disease resistance. The strongest being in one accession of the wild mango species Mangifera laurina (Akem, Grice et al., 2007; Grice e Bally, 2007). This is ongoing work with ongoing screening of germplasm and research in to the mechanisms of resistance. Breeders in the Australian Mango Breeding Program have also begun to develop hybrid breeding populations segregating for disease resistance and to integrate natural resistance in to commercially acceptable cultivars (BALLY, AKEM et al., 2010).

HARVESTING AND HANDLING MANGO SAP

Harvesting fruit at optimum maturity to maximise the eating quality when ripe has traditionally been done using visual signals from fruit shape internal flesh colour and skin colour. In some cultivars these visual signals are week making maturity determination in unripe fruit difficult. Over recent years several instruments have been developed to non-destructively measure fruit maturity. Near infrared (SWNIR) (400-1100 nm) spectroscopy to non-destructively determine fruit maturity at harvest and link it to eating quality in ripe fruit (SUBEDI; WALSH et al., 2007; SUBEDI; WALSH, 2009). Leuchaudel et al (2010) have also used chlorophyll fluorescence to measure fruit maturity in developing fruits. A near infrared hand held instrument (Nirvana) supplied by Integrated Spectronics Pty. Ltd. Sydney (www.intspec.com/products) is being routinely used to determine fruit maturity of Calypso[TM] mangoes by larger growers to schedule harvesting to maximise fruit eating quality for consumers.

Lactiferous sap from the resin canals in the mango peel contains several caustic compounds that will burn the skin of people harvesting the fruit and the skin of the fruit being harvested if not managed appropriately. The causticity of the sap varies between cultivars and with fruit maturity and environmental conditions (HESSE; BOWDEN, 1995; HASSAN; IRVING et al., 2009). Sap is responsible for several fruit blemishes that detract from the fruit aesthetically and reduce the market value of affected fruit. The blemishes commonly referred to as 'sap-burn' and 'skin-browning' have lactiferous sap as their primary causal agent. In the cultivar 'Kensington Pride', the sap is particularly caustic and the Australian industry has developed methods of managing the harvesting process to minimise sap contamination of fruit. Fruit are essentially harvested in one of two ways, with long stems and de-sapped in the packing shed or with stem removal in the field with the use of harvest aids. Shed de-sapping to remove the caustic sap generally involves harvesting fruit with the peduncle attached and pre-dipping fruit in a detergent solution to protect the fruit before hand de-stemming in a manner that directs the sap flow away from the fruit. Field harvest-aids have been developed to de-sap fruit as it is picked in the field and avoid the de-sapping bottlenecks in the packing shed. Harvest-aid design has progressed over the past decade with a wide range of designs being used to day. Two popular designs are elevated platform cherry picker styles where fruit is hand picked into an elevated bin that is constantly being sprayed with a de-sapping agent and mobile ground models that use a detergent covered tarpaulin to catch fruit picked from the tree by hand or by harvest hocks with peduncles removed (figure 1 and 2).

The examples above of some of the advanced in mango production and quality derived through research and development, demonstrate the health of world mango industries and its adaptability for current and future challenges.

[FIGURE 1 OMITTED]

[FIGURE 2 OMITTED]

REFERENCES

AKEM, C. et al Preliminary evaluation of mango varieties for postharvest disease resistance. In: THE AUSTRALIAN MANGO CONFERENCE, 6., 2007. Proceedings ... Melbourne: Australian Mango Industry Association, 2007. p.3-4.

BALLY, I. S. E. The effect of preharvest nutrition and crop load on fruit quality and postharvest disease in mango (Mangifera indica L). 2006. 196 f. Thesis (Doctor of Philosophy)--School of Agronomy and Horticulture, University of Queensland, Brisbane, 2006.

BALLY, I. S. E. et al Mango breeding for disease resistance, current activities and future prospects. In: INTERNATIONAL MANGO SYMPOSIUM, 9., 2010. Sanya. Proceedings. Sanya: International Society for Horticultural Science, 2010.

BALLY, I. S. E. et al. The effects of nitrogen on postharvest disease in mango (Mangifera indica L. 'Keitt'). Acta Horticulturae, The Hague, v. 820, p. 365-370, 2009.

DILLON, N. et al. Implementation of SSR markers in mango breeding in Australia. In: INTERNATIONAL MANGO SYMPOSIUM, 9., 2010, Sanya. Proceedings. Sanya: International Society for Horticultural Science, 2010.

DILLON, N. L. et al Genetic diversity of Australian Mango (Mangifera spp.) genebank Tree Genetics & Genomes, 2010. In press.

DUVAL, M. F. et al. Development of microsatellite markers for mango (Mangifera indica L.). Molecular Ecology Notes, Oxford, v. 5, n. 4, p. 824-826, 2005.

EIADTHONG, W. et al. Amplified fragment length polymorphism analysis for studying genetic relationships among Mangifera species in Thailand. Journal of the American Society for Horticultural Science, Alexandria, v. 125, n. 2, p. 160-164, 2000.

EIADTHONG, W. et al.. Identification of mango cultivars of Thailand and evaluation of their genetic variation using the amplified fragments by simple sequence repeat-(SSR-) anchored primers. Scientia Horticulturae, Amsterdam, v. 82, n. 1-2, p. 57-66, 1999.

GALAN SAUCO, V Worldwide mango production and market. Current situation and future prospects. In: INTERNATIONAL MANGO SYMPOSIUM, 9., 2010. Sanya. Proceedings. Sanya: International Society for Horticultural Science, 2010.

GALVEZ-LOPEZ, D. et al. Genetic analysis of mango landraces from Mexico based on molecular markers. Plant Genetic Resources, Cambrigde, v. 7, n. 03, p. 244-251, 2009.

GERBAUD, P. Close-Up. Mango. FruiTROP, Paris, v. 164, p. 7-39, 2009.

GRICE, K.; BALLY, I. S. E. Preliminary screening for anthracnose resistance in mango. THE AMISTAR AUSTRALIAN MANGO CONFERENCE, 6., 2007. Queensland. Proceedings ... Queensland: Australian Interactive Media Industry Associaion, 2007.

HASSAN, M. K. et al. Retention of the fruit peduncle at harvest retains sap and contributes to resistance against post-harvest anthracnose disease in 'Kensington Pride', but not in 'R2E2' mango. Journal of Horticultural Science & Biotechnology, Asford, v. 86, n. 3, p. 261-266, 2011.

HASSAN, M. K. et al. Concentrations of constitutive alk(en)ylresorcinols in peel of commercial mango varieties and resistance to postharvest anthracnose. Physiological and Molecular Plant Pathology, London, v. 71, p. 158-165, 2007.

HASSAN, M. K. et al. Sap properties and alk(en) ylresorcinol concentrations in Australian-grown mangoes. Annals of Applied Biology, Warwick, v. 154, n. 3, p. 419-427, 2009.

HESSE, A.; BOWDEN, B. Chemical investigation of mango sap components. In: MANGO 2000 MARKETING SEMINAR AND PRODUCTION WORKSHOP, 2000, Townsville. p.265-270.

HIDAYAT, T. et al. Molecular Diversification and Phylogeny of Mangifera (Anacardiaceae) in Indonesia and Thailand. In: INTERNATIONAL CONFERENCE ON ADVANCED SCIENCE, ENGINEERING AND INFORMATION TECHNOLOGY, 14 15 january 2011, Malaysia. Proceedings. p.88-91.

HONSHO, C. et al. Isolation and characterization of new microsatellite markers in mango (Mangifera indica). Molecular Ecology Notes, Oxford, v. 5, n. 1, p. 152-154, 2005.

IP AUSTRALIA. Mango (Mangifera indica), Variety: NMBP1243. Plant Varieties Journal, Cambridge, v. 21, n. 3, p. 84-85; 275-283, 2008.

KANZAKI, S. et al. Use of microsatellite markers to test self- and cross-pollinated seedlings of some mango cultivars. In: INTERNATIONAL SYMPOSIUM ON BIOTECHNOLOGY OF TEMPERATE FRUIT CROPS AND TROPICAL SPECIES, 2005. Daytona Beach. Proceedings.

LECHAUDEL, M. et al. Chlorophyll fluorescence, a nondestructive method to assess maturity of mango fruits (Cv. a[euro]~Cogshalla[euro] [TM]) without Growth Conditions Bias. Journal of Agricultural and Food Chemistry, Easton, v. 58, n. 13, p. 7532-7538, 2010.

PHILLIPS, R. L.; VASIL, I. K. (Ed.). DNA-based markers in plants. 2nd ed. Dordrecht: Kluwer Academic Publishers, 2001.

ROSSETTO, C. J. et al. Seca-da-mangueira. avaliacao de porta-enxertos resistentes para produtividade. Planaltina: EMBRAPA Cerrados, 2006.

SCHNELL, R. J. Integrating genomics into applied tropical fruit breeding programs. In: IFAS. Tropical fruit production and handling. Miami; University of Florida, 2006. p.27-29. Disponivel em: <http:// tfphw.ifas.ufl.edu>.

SCHNELL, R. J. et al. Mango genetic diversity analysis and pedigree inferences for Florida cultivars using microsatellite markers. Journal of the American Society for Horticultural Science, Alexandria, v. 131, n. 2, p. 214-224, 2006.

SCHNELL, R. J. et al. Isolation and characterization of 15 microsatellite loci from mango (Mangifera indica L.) and cross-species amplification in closely related taxa. Molecular Ecology Notes, Osford, v. 5, p. 625-627, 2005.

SMITH, J. S. C.; SMITH, O. S. Fingerprinting Crop Varieties. Advances in Agronomy, Madison, v. 47, p. 86-140, 1992.

SMITH, M. W. et al. Rootstock affects yield, yield efficiency, and harvest rate of 'Kensington Pride' mango. Hortscience, Alexandria, v. 38, n. 2, p. 273-276, 2003.

STRAHAN, R.; PRATT, R. investment analysis of the development of the calypso mango variety. Brisbane: DEEDI, 2011. p.5.

SUBEDI, P. P.; WALSH, K. B. Non-invasive techniques for measurement of fresh fruit firmness. Postharvest Biology and Technology, Amsterdam, v. 51, n. 3, p. 297-304, 2009.

SUBEDI, P. P. et al. Prediction of mango eating quality at harvest using short-wave near infrared spectrometry. Postharvest Biology and Technology, Amsterdam, v. 43, n. 3, p. 326-334, 2007.

UKOSKIT, K. Developmento f Microsatellite Markers in Mango (Mangifera indica L.) using 5' anchored PCR. Thammasat International Journal of Science and Technology, New York, v. 12, n. 3, p. 1-7, 2007.

VARSHNEY, R. K. et al. Genic microsatellite markers in plants: features and applications. Trends in Biotechnology, Amsterdam, v. 23, n. 1, p. 48-55, 2005.

VIRUEL, M. A. et al. Fingerprinting, embryo type and geographic differentiation in mango (Mangifera indica L., Anacardiaceae) with microsatellites. Molecular Breeding, Dordrecht, v. 15, p. 383-393, 2005.

YONEMORI, K. et al. Phylogenetic relationships of Mangifera species revealed by ITS sequences of nuclear ribosomal DNA and a possibility of their hybrid origin. Plant Systematics and Evolution, New York, v. 231, n. 1-4, p. 59-75, 2002.

IAN S. E. BALLY (2)

(1) Palestra Sinfruit 093--Simposio Internacional de Fruticultura--Avancos na Fruticultura (17 a 21 Outubro)

(2) Professional qualifications, B. app. Sc. (Biol), Central Queensland University, PhD, Agricultural Science, University of Queensland.

E-mail: ian.bally@gmail.com
TABLE 1--Some of the new recently breed mango cultivars that are being
adopted by growers and traded nationally and internationally.

Cultivar name     Country     Plant Variety       Comments
                  of origin   Protection

Delta R2E2        Australia   Public Domain       Large fruit popular
                                                    in Asia
                                                    as a gift fruit
B74A              Australia   PBR (A) Australia   High blush,
  (Calypso[TM])                 &                   fibreless pulp
                                Internationally     with excellent
                                                    shipping
                                                    qualities
Honey Gold (A)    Australia   PBR (A) Australia   Rich and sweet
                                                    'Kensington'
                                                    style flavour
                                                    with a firm,
                                                    fibre-free flesh
NMBP1201 (A)      Australia   PBR (A) Australia   Early production
                                                    with sweet
                                                    'Kensington'
                                                    style flavour
NMBP1243 (A)      Australia   PBR (A) Australia   Firm high yielding
                                                    fruit with firm
                                                    flesh and
                                                    strong red blush
NMBP4069 (A)      Australia   PBR (A) Australia   Firm fruit with
                                                    sweet juicy
                                                    flesh and pink
                                                    blush
Joa               South       PBR (A) South       Mid season
                    Africa      Africa              cropping,
                                                    tolerant to
                                                    bacterial black
                                                    spot disease
Chene             South       PBR (A) South       Resistant to sun
                    Africa      Africa              scorch, tolerant
                                                    to bacterial
                                                    black spot
                                                    disease and
                                                    mid season
                                                    cropping
Alfa              Brazil                          Semi Dwarf, high
                                                    yielding with
                                                    some resistance
                                                    to fruit flies,
                                                    post-harvest
                                                    rots and powdery
                                                    mildew.
Roxa              Brazil                          High yield,
                                                    fibreless very
                                                    sweet
Lita              Brazil                          Late season,
                                                    regular cropping,
                                                    excellent fruit
                                                    quality.
Maha Chanok       Thailand    Public Domain       Long shape fruit
                                                    with orange-red
                                                    blush, thick
                                                    blemish resistant
                                                    skin
Summer Snow       Taiwan                          'Irwin' style
                                                    fragrance with,
                                                    low fibre, red
                                                    blush
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Author:Bally, Ian S.E.
Publication:Revista Brasileira de Fruticultura
Date:Oct 1, 2011
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