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Floral Biology and Pollination Efficiency In agriculture Review Floral Biology and Pollination Efficiency in Yam (Dioscorea spp.) Jean M. Mondo 1,2,3 , Paterne A. Agre 1,* , Alex Edemodu 1 , Patrick Adebola 4 , Robert Asiedu 1, Malachy O. Akoroda 5 and Asrat Asfaw 1 1 International Institute of Tropical Agriculture (IITA), Ibadan 5320, Nigeria; [email protected] (J.M.M.); [email protected] (A.E.); [email protected] (R.A.); [email protected] (A.A.) 2 Institute of Life and Earth Sciences, Pan African University, University of Ibadan, Ibadan 200284, Nigeria 3 Department of Crop Production, Université Evangélique en Afrique (UEA), Bukavu 3323, Congo 4 International Institute of Tropical Agriculture (IITA), Abuja 901101, Nigeria; [email protected] 5 Department of Agronomy, University of Ibadan, Ibadan 200284, Nigeria; [email protected] * Correspondence: [email protected] Received: 21 October 2020; Accepted: 12 November 2020; Published: 20 November 2020 Abstract: Yam (Dioscorea spp.) is a monocotyledonous herbaceous vine cultivated for its starchy underground or aerial tubers in the tropics and subtropics. It is an allogamous and polyploid species that reproduces by both sexual and asexual mechanisms. However, many of the landrace cultivars, including most of the popular varieties, reproduce exclusively by vegetative propagation (planting the tubers). These varieties are either sterile or produce sparse and irregular flowering with high flower abortion rate, low fruit and seed set. Production of crossbreed seeds for genetic improvement and for maintaining genetic diversity in yams is, therefore, mainly achieved through natural or managed pollination. Flowering in yam is mostly dioecious and, in some instances, monoecious. Flowering asynchrony, sticky nature of the pollen grains, and cross incompatibility are among the challenges in making genetic progress in yam breeding. There are many limitations in basic and applied knowledge of yam flower biology and pollination. This paper, therefore, reviews the flowering biology, pollination, and methods of improving pollination efficiency in yam breeding programs. Keywords: profuse flowering; cross incompatibility breakage; pollen viability and storage; West Africa 1. Introduction Yam is a common name for diverse species in the genus Dioscorea. This genus is composed of ~600 species, of which eleven are the main ones cultivated globally and eight of these are grown in Africa [1–4]. In Africa, yam is extensively cultivated in a region from West Cameroon to central Côte d’Ivoire, also referred to as the African “yam belt” [5,6]. The yam belt comprises six countries: Nigeria, Ghana, Côte d’Ivoire, Benin, Togo and Cameroon. This zone accounts for ~92% (67.1 million tons) of the annual global yam production [7]. Yam is an important source of dietary nutrients [8,9] with a low glycemic index, which gives better protection against diabetes and obesity [10,11]. Yam value chain is a direct income-generating activity for more than five million people in West Africa [12]. Thus, it provides an opportunity for poverty alleviation and food security. Breeding is key to release the potential of yam for food security and poverty reduction. Yam breeding uses both sexual and asexual reproductive mechanisms. Sexual reproduction involves an artificial hand or natural open pollination during flowering to generate genetically variable offspring for selection [13]. This step starts with selecting parents with desirable traits. The mating design’s choice is based on a breeding program’s objective, technical and financial capacity. Progenies with desirable Agriculture 2020, 10, 560; doi:10.3390/agriculture10110560 www.mdpi.com/journal/agriculture Agriculture 2020,, 1010,, 560x FOR PEER REVIEW 2 of 2123 choice is based on a breeding program’s objective, technical and financial capacity. Progenies with traitsdesirable such traits as yield such stability, as yield superior stability, resistance superior toresistance disease andto disease pests, andand higherpests, and tuber higher quality tuber are thenquality selected are then and selected advanced and vegetatively advanced for vegetatively several generations for several and generations across locations and across before releasinglocations asbefore new releasing varieties as [3 ,new13]. varieties Therefore, [3,13]. plant Therefore, breeding plant programs’ breeding success programs’ relies onsuccess genetic relies variation on genetic for traitsvariation of interest for traits and of successful interest and transfer successful of the genetictransfer allele of the of traitsgenetic from allele source of traits germplasm from source to the agronomicallygermplasm to preferredthe agronomically background. preferred The transfer background. of genes The or alleles transfer depends of genes on flowering,or alleles depends pollination, on fruitflowering, seed set, pollination, and seed fruit germination. seed set, and seed germination. Pollination isis the the movement movement of of pollen pollen from from the the male male sexual sexual parts parts of a plant of a toplant the receptiveto the receptive surface ofsurface a plant’s of a female plant’s sexual female parts sexual [14 parts]. Pollination [14]. Pollination is central is to central the transfer to the of transfer genes inof plantgenes breeding. in plant Itbreeding. involves It three involves phases: three the phases: release the of pollen, release transfer of pollen, of pollen, transfer and of placementpollen, and of placement pollen (followed of pollen by germination)(followed by [ 14germination)]. The pollination [14]. successThe pollination is associated success with manyis associated factors: pollenwith many quantity factors: and viability, pollen stigmaquantity receptivity, and viability, cross stigma compatibility, receptivity, the cross efficiency compatibility, of pollinator the agentsefficiency and of the pollinator prevailing agents weather and conditionsthe prevailing [3,15 weather] (Figure conditions1). [3,15] (Figure 1). Viable pollen Stigma Weather receptivity Pollination success Pollination Parents agents compatibility Figure 1. Factors linked to plant pollination. Pollination effiefficiencyciency at aat breeding a breeding program program level can level be defined can asbe the defined number ofas viablethe /numbergerminable of seedsviable/germinable over the number seeds of over ripe the female number flowers of ripe which female received flowers viable which pollen. received This ratioviable is pollen. low in yamThis plantratio is and low varies in yam with plant studies and fromvaries 1 towith ~40% studies [3,15 from–17]. 1 At to the ~40% International [3,15–17]. At Institute the International of Tropical AgricultureInstitute of (IITA),Tropical Ibadan, Agriculture Nigeria, (IITA), the percentage Ibadan, fruitNigeria, and seedthe percentage sets are estimated fruit and at 20.3 seed and sets 10.5%, are respectively,estimated at 20.3 for D.and rotundata 10.5%, respectively,and 28.0 and for 9.3%, D. rotundata respectively, and 28.0 for D.and alata 9.3%,[3 ].respectively, This low success for D. alata rate is[3]. partly This linkedlow success to yam’s rate long-term is partly vegetative linked to propagation yam’s long since‐term its vegetative domestication propagation [18]. Some since of theits domestication e[18].ffects Some on yam of the reproduction domestication are sparse, effects irregular,on yam reproduction or absent flowering, are sparse, high irregular, flower and or ovuleabsent abortion flowering, rates, high low flower fruit and and ovule seed abortion set, and lowrates, seed low viability fruit and [ 19seed]. Besides,set, and low the synchronizedseed viability flowering[19]. Besides, time the between synchronized male and flowering female individuals time between is uncertain male and [3,15 female]. The sticky individuals nature ofis theuncertain pollen grains[3,15]. The renders sticky wind nature pollination of the pollen impossible. grains Therenders ineffi windciency pollination of insects interveningimpossible. inThe pollen inefficiency dispersal, of crossinsects incompatibility, intervening in and pollen suboptimal dispersal, weather cross conditionsincompatibility, contribute and tosuboptimal low pollination weather and conditions seed set in yamcontribute under to natural low pollination conditions and [20 ,seed21]. Even set in though yam under the pollination natural conditions efficiency [20,21]. has remained Even though low over the time,pollination the topic efficiency is currently has neglected.remained low Most over relevant time, publications the topic is in currently yam floral neglected. biology and Most pollination relevant arepublications 30–50 years in old.yam There floral is,biology therefore, and apollination need to deepen are 30–50 knowledge years old. on yam There floral is, therefore, biology and a need update to recommendationsdeepen knowledge on on practices yam floral linked biology to pollination and update efficiency recommendations to match with theon currentpractices realities. linked to pollination efficiency to match with the current realities. Agriculture 2020, 10, 560 3 of 21 We hypothesized that yam pollination efficiency is improvable by integrating a range of practices discussed throughout this review. These include quality pollen collection and preservation, adequate pollination methods, identification of and rearing insect pollinators, agronomic and biotechnological manipulations. Therefore, this review explored available knowledge on yam floral biology, pollination practices, constraints limiting pollination
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