Impact of low-cost management techniques on population dynamics of plant-parasitic nematodes in sweet potato
Introduction
Sweet potato (Ipomoea batatas) is a preferred food security crop in sub-Saharan Africa because it is drought tolerant, performs well under unfavorable conditions and requires little or no management inputs compared to other crops (Woolfe, 1992; Kaguongo et al., 2012; Stathers et al., 2013). In Kenya, sweet potato is mainly cultivated by smallholder farmers in Central, Rift Valley, Nyanza, Western and Coast Provinces with intensive production in Nyanza and Western regions (Kaguongo et al., 2012). The entire sweet potato plant has varied uses such as human food, feed for livestock and as industrial raw material (Githunguri and Migwa, 2007; Claessens et al., 2008; Duvernay et al., 2013). Sweet potato yields in Kenya have steadily declined from 125,050 hg ha-1 in 2014 to 94,220 hg ha-1 in 2017 (FAOSTAT, 2019). Production is limited by various biotic constraints including weevils, aphids, and nematodes (Kivuva et al., 2014; Echodu et al., 2019).
Nematodes are distributed in all environments as both parasites and free-living organisms, and they influence crop production in different ways. Free-living nematodes play crucial roles such as decomposition, mineralization of organic materials and regulation of parasitic nematodes (Xiao et al., 2010; Neher et al., 2012; Ferris et al., 2012a). In addition, they are useful bioindicators of soil health status in agro-ecosystems (Bongers, 1990; Neher, 2001). Free-living nematodes occupy various trophic levels and are classified as bacterivores, fungivores, omnivores and predators based on their feeding habits (Yeates et al., 1993). Plant-parasitic nematodes (PPN) on the other hand negatively affect crop productivity. Worldwide, PPN cause an annual estimated yield loss of up to $118 billion in many crops (Atkinson et al., 2011).
Sweet potato is characterized by three distinct growth phases with variations in the time for each phase based on the cultivar (Woolfe, 1992). Different parasitic nematodes affect sweet potato at particular growth stages. For instance, the reniform nematode, Rotylenchulus reniformis attacks sweet potato during the first two stages and reduces the root system through a “pruning” effect (Clark and Wright, 1983) while Meloidogyne incognita affects the second and third growth stages (Lawrence et al., 1986; Agu, 2004). Concomitant infection by multiple PPN species cause varied responses in sweet potato (Thomas and Clark, 1983; Agu, 2006). Root galls, necrosis and root cracking are key symptoms of sweet potato infected by parasitic nematodes (Coyne et al., 2014). Root galls limit absorption and transport of nutrients and water while cracking and necrosis reduce quality and marketability of storage root (Lawrence et al., 1986). The infection sites create entry points for secondary pathogens such as fungi and bacteria which further lowers yields (Cervantes-Flores et al., 2008; Nicol et al., 2011). Economically important PPN associated with sweet potato include Meloidogyne, Pratylenchus, Rotylenchulus and Ditylenchus species (Niere and Karuri, 2018). Most PPN are controlled using nematicides and soil fumigants. Despite these chemicals being fast-acting and effective (Adegbite and Agbaje, 2007; Dubey and Trivedi, 2011; Abbas et al., 2015), their use is limited due to their negative effects on the environment, non-target organisms, human and animal health (Udo and Ugwuoke, 2010; Renco and Kovacik, 2012). Other nematode management strategies include use of resistant cultivars, crop rotation, use of cover crops, soil solarization and use of microorganisms which also have their own limitations (Mcsorley, 2011; Hajji and Horrigue-Raouni, 2012; Suzuki et al., 2012; Gregory et al., 2017). Resistant cultivars are not available for most crops and those that exist are nematode species-specific and hence not suitable for PPN control in agro-ecosystems with multiple species (Hockland et al., 2012; Briar et al., 2016). Crop rotation is complex in its design and implementation especially where many nematode species are present. Soil solarization on the other hand is expensive and hence inapplicable in large scale farming (Briar et al., 2016).
Use of organic amendments presents a promising strategy which not only increases overall yields and improves plant growth and soil parameters, but also control soil borne pests and diseases including parasitic nematodes (Akhtar and Malik, 2000; Oka, 2010; Renco et al., 2011). Commonly used organic amendments with suppressive effects on PPN include animal manure, green manure and composted materials (Kimenju et al., 2008; Rivera and Aballay, 2008; Hu and Qi, 2010; Renco et al., 2011; Renco, 2013; Amulu and Adekunle, 2015; Osunlola and Fawole, 2015). A decrease in PPN which results from application of different organic amendments is associated with production of nematicidal compounds such as ammonia, different secondary metabolites and organic acids (Thoden et al., 2011). However, nematicidal activity depends on physico-chemical parameters of soil such as pH and temperature (Oka, 2010). In addition, to the toxic effect of these materials on PPN, the natural ability of soil to regulate PPN is increased by addition of organic matter due to the increase in predatory and omnivorous free-living nematodes and the general improvement of the soil food web (Sánchez-Moreno and Ferris, 2018). Effectiveness of organic amendments on PPN infecting sweet potato in Kenya has not been evaluated. We sought to determine the effect of intercropping sweet potato and maize, application of Tithonia diversifolia, cow and goat manure on PPN abundance, nematode metabolic footprints, ecological and functional indices.
Section snippets
Experimental site
The study was carried out in Nyangati sub-location in Mwea, Kirinyaga County, Kenya. It lies between Latitude 0° 37′ 11.1″ S and Longitude 37° 21′ 41.2″ E and it is in the Low Midland (LM) agro-ecological zone, at an altitude of 1204 m above sea level. The soil in the area is classified as vertisols (black cotton soil) with clay content that is higher than 30% (Jaetzold et al., 2009). Soil physicochemical properties of the site are given in Table 1. During the study period, temperature ranged
Effects of different treatments on relative abundance of nematode genera
A total of 47 nematode genera belonging to five trophic groups (bacterivores, herbivores, fungivores, predators and omnivores) were identified in the two seasons (long rains; LR) and short rains; SR)); 47 during LR and 46 in SR. Bacterivores and herbivores were the most dominant nematode feeding guilds followed by omnivores while fungivorous and predatory genera were less frequent in the two seasons (Table 3, Table 4). Aorolaimus, Criconemoides, Hoplolaimus, Mesorhabditis and Dorylaimus were
Discussion
A high diversity of PPN and free-living nematodes was observed across treatments during this study. Meloidogyne, Pratylenchus, Rotylenchulus and Ditylenchus which are the major parasitic nematodes of economic importance (Niere and Karuri, 2018) in sweet potato were recorded. These PPN genera have also been reported in sweet potato fields in Uganda, Niger and Kenya (Coyne et al., 2003; Haougui et al., 2011; Karuri et al., 2017). They attack sweet potato at different growth stages resulting in
Conclusion
Economically important PPN in sweet potato were observed in association with sweet potato. Application of organic amendments in sweet potato plots had varying effects on abundance and diversity of nematodes, metabolic footprints, ecological and functional indices. Goat manure showed the highest potential in management of PPN during the two seasons. Based on the functional metabolic footprints, most treatment plots were classified as degraded in both seasons but MS was structured in SR. The CM
Funding
This work was supported by the Department for International Development under the Climate Impact Research Capacity and Leadership Enhancement programme as part of research uptake fund.
CRediT authorship contribution statement
Hellen Maina: Writing - original draft, Investigation. Hannah Karuri: Conceptualization, Supervision, Methodology, Writing - review & editing. Felix Rotich: Supervision, Writing - review & editing. Franklin Nyabuga: Supervision, Writing - review & editing.
Declaration of competing interest
The authors declare that they have no conflict of interest.
Acknowledgement
The authors acknowledge the assistance of Peter Kithinji during field trials.
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