Towards Pathogen-Free Coconut Germplasm Exchange
Abstract
:1. Introduction
2. The Disease-Causing Agents of Concern
2.1. Phytoplasma
Agent | Disease | Localization | Symptoms | References |
---|---|---|---|---|
Phytoplasma | Coconut lethal yellowing disease (LY) | Husk, shell, endosperm embryo, plumule, phloem cells and haustorial tissue from diseased coconut palms | Untimely shedding of fruit, discoloration in younger leaves, and darkening of the inflorescence, finally leading to the death of the coconut palm. | [22,38,39] |
Coconut root wilt disease (CRWD) | Leaves, inflorescence, and roots | Flaccid bending of leaflets, accompanied by leaf yellowing, necrosis, compromised stomatal regulation, and damage to the root system. | [24] | |
Weligama coconut leaf wilt disease (WCLW) | Sieve tubes, leaflets | The leaves exhibit flaccidity and marginal necrosis, accompanied by intense yellowing of the fronds. With disease advancement, the crown diminishes in size, and the trunk starts to taper. In addition, the female flower production declines, as does the palm’s productivity. Moderately affected palms show necrosis in the root tips, while severely affected palms lack young roots. Some of the infected palms also exhibit leaf rot disease. However, no studies have reported the death of a coconut palm caused by this disease. | [23,40,41,42] | |
Cape St. Paul wilt disease (CSPW) | Fronds, inflorescence, and leaves | Premature nut drops, yellowing of fronds, and necrosis of immature inflorescences, followed by progressive yellowing of the crown from the older leaves upwards, and death of palms. | [43] | |
Bogia coconut syndrome (BCS) | Fruit husk, shell, endosperm, embryo, trunk phloem | Premature dropping of fruit of all ages. The outer fronds of the crown will droop and develop a pale yellow colour. Fronds then turn brown and hang down the stem, like a skirt. A dry rot develops in the newly expanding spear, progressing downwards to the growing point. Complete necrosis—death of the palm. | [22] | |
Virus | Coconut foliar decay virus disease (CFDV) | Leaflet, frond, phloem, tissue adjacent to and within necrotic zones | Yellowing in certain leaflets below the unfolding spear leaf. Then, a broader yellowing occurs in both the affected fronds and neighboring organs. Synchronously, these fronds undergo a lateral necrosis near the petiole’s base, resulting in their collapse. | [44] |
Viroid | Coconut cadang-cadang viroid disease (CCCVd) | Vascular tissues, nucleolus of mesophyll cells, embryos. | The fruit assumes a rounded shape, displaying distinctive scarifications at its equator, while the initial non-necrotic, translucent, bright yellow leaf spots emerge. Inflorescences undergo necrosis, halting fruit production, slowing down new frond development, and causing larger and more frequent leaf spots. Later, fronds start to appear chlorotic when observed from a distance. Finally, preceding death, leaf spots merge, and the entire crown exhibits a distinct yellow or bronze hue, significantly reduced in size with a diminished number of fronds. | [45,46,47] |
Coconut tinangaja viroid disease (CTiVd) | Unreported | A decrease in frond quantity, along with diminished size and quantity of fruit, leads to the shriveling and deformation of the fruit. This results in the cessation of fruit production, failure to generate inflorescences, tapering of the distal end of the trunk, persistent stipules, stippling of leaflets characterized by fine chlorotic spots, and the development of brittleness in both leaflets and fronds. | [48] |
2.2. Viruses
2.3. Viroids
3. Tissues Used for Coconut Germplasm Exchange
3.1. Zygotic Embryos
3.2. Apical Meristems
3.3. Somatic Embryos/Embryogenic Callus
4. Methods for the Detection of Lethal Pathogens
4.1. Detection Using Artificial Transmission Test Systems
4.2. Visual Detection Methods
4.3. Immunological Detection Methods
4.4. Molecular Detection Methods
4.4.1. Molecular Hybridization Assays
4.4.2. Polymerase Chain Reaction (PCR)-Based Methods
- (a)
- Conventional PCR: The molecular detection of phytoplasmas present in symptomatic tissue is routinely undertaken by the PCR using phytoplasma-specific universal or phytoplasma group-specific primers designed based on the highly conserved 16S ribosomal RNA (rRNA) gene sequences, the ribosomal protein, and elongation factor genes [101]. Phytoplasmas can be easily detected using PCR with the highest sensitivity in immature rather than mature tissues [101], as evidenced by the detection of LY phytoplasma in coconut leaves [102] and the Coconut yellow decline (CYD) phytoplasma in spear leaves, inflorescences, or trunk tissues of affected symptomatic and asymptomatic coconut palm varieties [103]. A high population of viroid-like molecules has been identified using PCR coconut palms infected with the Coconut tapering disease in Sri Lanka [104]. Similarly, a sense-specific single-primer PCR assay has been employed to identify CFDV DNA from coconut palms in Vanuatu [105].
- (b)
- Reverse transcriptase-PCR (RT-PCR): As most plant viruses have RNA genomes (and even DNA viruses produce RNA transcripts), it is most effective to detect virus infections by analyzing RNA sequences from infected plant samples [106]. The RT-PCR technique has been employed for detecting low concentrations of viruses and viroids in infected plants, with the only requirement being that of obtaining good quality RNA [106]. The application of the RT-PCR technique has successfully helped to detect the CCCVd viroid variant in oil palm leaf tissues [99], the Areca palm necrotic ringspot virus (ANRSV) in areca palm (Areca catechu Linn. [86]), and African oil palm ringspot virus (AOPRV) in infected oil palm leaves [107].
- (c)
- Nested PCR: Nested PCR methods, where four primers are used in two consecutive rounds of DNA amplification, have greatly enhanced specificity and sensitivity, and therefore aid in the detection of a low titer of pathogens [108,109]. Most of the nested PCR protocols developed have been focusing on phytoplasma detection and are used to increase the sensitivity of the available protocol [98]. Nesting of universal group primers with group-specific primers has helped in the diagnosis of phytoplasma infection in well-defined taxonomic groups [110]. For example, the WCLW-causing phytoplasma has been successfully detected using an optimized nested PCR in symptomatic coconut plants in Sri Lanka [111]. Similarly, CRWD- and CSPW-causing phytoplasma have also been detected in coconut palms using nested PCR [15,24,112].
- (d)
- Quantitative PCR (qPCR): In recent years, qPCR has proven to be an indispensable tool for the molecular diagnosis of pathogens. It is about 10 times more sensitive than standard PCR and does not require gel electrophoresis for target confirmation. The qPCR can simultaneously detect and quantify the pathogen, and this method is less affected by cross-contamination and is less work-intensive as compared to RT-PCR and nested PCR. For example, phytoplasma belonging to the 16SrIVgroup causing CRWD have been easily and effectively detected by qPCR [113], especially when compared to conventional and nested PCR [114]. The qPCR approach using the TaqMan probe gives higher sensitivity than nested-PCR when detecting the 16SrIV subgroups of the LY phytoplasma in the coconut [83,115,116]. The CCCVd variants in oil palm have been detected by using the qPCR technique [117]. This technique could be used in the coconut as well.
- (e)
- Digital PCR: Digital PCR (dPCR) is a breakthrough next-generation PCR technology, and it works by partitioning the sample into thousands of separate reaction compartments, conducting single, parallel PCR reactions [118]. It is used to detect phytoplasma and viruses, with several advantages over qPCR diagnostic assays, including higher sensitivity, precision, accuracy in detecting extremely rare target sequences, absolute quantification without standard curve and reference samples, greater tolerance to PCR inhibitors, and suitability for the preparation of in-house reference materials [119]. Reverse transcription (RT) dPCR has been used to detect viruses and viroids in apple and citrus plants [120,121], and this technique can be used for the coconut.
4.4.3. Isothermal Amplification Techniques
4.4.4. Next-Generation Sequencing
4.5. Limitations of Current Detection Methods
5. Methods for the Elimination of Lethal Pathogens
5.1. Chemotherapy
5.2. Thermotherapy and Cryotherapy
5.3. Limitations of Elimination Methods
6. Research Gaps and Research Priorities towards Safe Exchange of Coconut Germplasm
6.1. Lack of Knowledge on Vertical Transmission of Lethal Coconut Pathogens, and Their Detection in Various Tissues
6.2. Lack of Robust Disease Elimination Methods
6.3. Lack of Plant Health and Exchange Regulations among Coconut-Growing Countries
7. Conclusions
Author Contributions
Funding
Data Availability Statement
Acknowledgments
Conflicts of Interest
References
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---|---|---|---|
Field transmission | Artificial transmission test systems | Phytoplasma | [43,81] |
In vitro transmission | Artificial transmission test systems | Phytoplasma | [20,83] |
Transmission electron microscope (TEM) | Visual | Viroids, virus, phytoplasma | [86] |
Fluorescent stain 4′,6-diamidino-2-phenylindole (DAPI). | Visual | Phytoplasma | [88] |
ELISA assay | Immunological | Phytoplasma | [93,94,95] |
Antibiotic response | Immunological | Phytoplasma | [97] |
Hybridization assay | Molecular | Viroids, phytoplasma | [100] |
Polymerase chain reaction (PCR)-based methods | Molecular | Virus, viroids, phytoplasma | [15,24,83,99,102,103,104,105,111,112,113,114,115,116] |
Isothermal amplification techniques (IAT) | Molecular | Virus, viroids, phytoplasma | [126,127,128,129] |
Next-generation sequencing (NGS) | Molecular | Not yet studied in coconut | [106,130] |
Methods | Targeted Pathogens | Targeted Coconut Pathogens | Source |
---|---|---|---|
Chemotherapy | Phytoplasma | Phytoplasma | [136,137] |
Thermotherapy | Virus | Not yet studied in coconut | [142,143] |
Cryotherapy | Virus | Not yet studied in coconut | [144,145,146,147] |
Name of Guideline | Source | Year Published |
---|---|---|
FAO/IBPGR Technical Guidelines for the Safe Movement of Coconut Germplasm | [154] | 1993 |
Germplasm Health Management for COGENT’s Multi-Site International Coconut Genebank | [156] | 2004 |
COGENT Global Conservation Strategy for Cocos Nucifera: A Framework for Promoting the Effective Conservation and Use of Coconut Genetic Resources Developed in Consultation with COGENT Members and Partners | [157] | 2008 |
Technical guidelines for the safe movement and duplication of coconut (Cocos nucifera L.) germplasm using embryo culture transfer protocols | [158] | 2012 |
A Global Strategy for the Conservation and Use of Coconut Genetic Resources, 2018–2028: Summary Brochure | [159] | 2018 |
Coconut risk management and mitigation manual for the Pacific Region | [160] | 2021 |
Year | Coconut Germplasm Guidelines | Advances/Gaps in the Understanding of Phytoplasma, Virus and Viroid Diseases |
---|---|---|
Before 2003 | Coconut fruit (dehusked and surface-sterilized), some zygotic embryos and pollen were the main internationally exchanged materials. The guidelines stated that pollen had to be tested for the presence of phytoplasma, and de-husked fruit and in vitro cultured embryos needed to be tested for viruses, viroids, or insect pests [154]. The embryos should first be cultured in vitro in the country of origin rather than the receiving country. If the country of origin or the recipient lacks the necessary tissue culture facilities, embryos should be sent to a different, third country for their culture [154]. | Phytoplasma diseases were still largely unexplored. Coconut husk, shell, and endosperm tissues were all shown to contain the Coconut cadang-cadang viroid [45]. DNA of the Coconut foliar decay virus was detected on the coconut fruit husk and in zygotic embryos but was not shown to be transferred through zygotic embryo cultures [161]. |
2003–2005 | Germplasm should be distributed in an in vitro form. No strict management guidelines were developed for the donor or recipient countries for zygotic embryos and other in vitro transferred materials. Tissues should not be sourced from disease-stricken areas, and pathogen testing should be undertaken. Pest risk analysis approaches were introduced to assist in pest control [156]. Food and Agriculture Organization/The International Board for Plant Genetic Resources (FAO/IBPGR) released technical guidelines for the safe movement of coconut germplasm, which recommended the transfer of isolated zygotic embryos instead of whole or de-husked fruit [137,156]. | Lethal yellowing phytoplasma DNA was detected in embryos extracted from diseased fruit by in situ PCR [38]. |
2005–2012 | The International Coconut Genetic Resources Network (COGENT) adopted the FAO/IBPGR guidelines and incorporated them into its global strategy for 2005–2015 and designated the embryo culture as a key technique to be further refined for the international exchange of coconut germplasm [157]. Pollen transfer was identified as having fewer quarantine complications than other forms of tissue transfer [157]. A general recommendation was made for the transfer of in vitro coconut materials, including a full procedure for the transfer of zygotic embryos. The recommendation was that in vitro germination should be undertaken in the donor country and the ex vitro growth take place in the recipient country [153,158] | Studies show that some embryos taken from phytoplasma-diseased coconut palms do not contain phytoplasma DNA [15,112]. Pollen is reported to transmit several lethal palm virus and viroid diseases [162]. |
2012–2020 | Guidelines were developed that required International genebanks to duplicate their collections in other geographical locations due to the threat from phytoplasma diseases [159]. | Phytoplasma DNA was detected in the plumules of in vitro germinated coconut embryos coming from infected fruit [20]. Bogia coconut syndrome and Banana wilt-associated phytoplasma DNA were shown to be introduced to coconut palms when caged with phytoplasma-carrying insects [22]. |
2021-present | Pollen transfer was still allowed; however, suspicion was raised about embryo culture; current embryo exchange activities became restricted. Further transmission confirmation research is now considered necessary for zygotic embryo transfer [160]. | An in vitro assay was developed to confirm the transmission of Lethal yellowing phytoplasma by planthoppers (Halaxius crudus) onto in vitro growing coconut seedlings, 16SrIV-group phytoplasmas were detected by nested PCR [83]. |
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Yang, C.; Nguyen, V.A.; Nulu, N.P.C.; Kalaipandian, S.; Beveridge, F.C.; Biddle, J.; Young, A.; Adkins, S.W. Towards Pathogen-Free Coconut Germplasm Exchange. Plants 2024, 13, 1809. https://doi.org/10.3390/plants13131809
Yang C, Nguyen VA, Nulu NPC, Kalaipandian S, Beveridge FC, Biddle J, Young A, Adkins SW. Towards Pathogen-Free Coconut Germplasm Exchange. Plants. 2024; 13(13):1809. https://doi.org/10.3390/plants13131809
Chicago/Turabian StyleYang, Chongxi, Van Anh Nguyen, Naga Prafulla Chandrika Nulu, Sundaravelpandian Kalaipandian, Fernanda Caro Beveridge, Julianne Biddle, Anthony Young, and Steve W. Adkins. 2024. "Towards Pathogen-Free Coconut Germplasm Exchange" Plants 13, no. 13: 1809. https://doi.org/10.3390/plants13131809
APA StyleYang, C., Nguyen, V. A., Nulu, N. P. C., Kalaipandian, S., Beveridge, F. C., Biddle, J., Young, A., & Adkins, S. W. (2024). Towards Pathogen-Free Coconut Germplasm Exchange. Plants, 13(13), 1809. https://doi.org/10.3390/plants13131809