Research paper Effects of planting density on branching habit in common and Tartary buckwheat Shinya KASAJIMA*, Wakana WATANABE, Maki KITADE, and Hirotake ITOH Faculty of Bioindustry, Tokyo University of Agriculture, Yasaka 196, Abashiri, Hokkaido 099-2493, Japan * Corresponding author: Shinya Kasajima (s3kasaji@nodai.ac.jp) E-mail addresses: s3kasaji@nodai.ac.jp (SK); 49518102@nodai.ac.jp (WW); 49518032@nodai.ac.jp (MK); h-ito@nodai.ac.jp (HI) DOI https://doi.org/10.3986/fag0030 Received: April 24, 2023; accepted May 7, 2023. Keywords: branching habit, common buckwheat, flower cluster, ideotype, planting density, Tartary buckwheat ABSTRACT This study investigated the effects of planting density on the branching habit of common and Tartary buckwheat. A field experiment was conducted using a split-plot design with three replicates, comparing sparse (67 plants m−2), mod- erate (111 plants m−2), and dense (222 plants m−2) planting densities in ‘Kitawasesoba’ (common) and ‘Manten-Kirari’ (Tartary) buckwheat varieties. Growth characteristics were examined at the stages of flower bud appearance, full flow- ering, and maturity. Results revealed that planting density had no significant effect on main stem length but showed significant effects on the number of branches in both species. As planting density increased, the number of branches decreased, with Tartary buckwheat exhibiting more significant changes in response to planting density than common buckwheat. The number of flower clusters on branches also decreased with increasing planting density. These findings suggest a potential role for branching plasticity in adapting to different planting densities while maintaining stable yields and indicate the importance of developing ideotypes with optimal branching habits for both species. This research contributes to a better understanding of buckwheat ideotypes and can contribute to future breeding efforts and crop management practices. Fagopyrum 40 (1): 15-21 (2023) 15 INTRODUCTION Two species of buckwheat are mainly used for food: common buckwheat (Fagopyrum esculentum Moench) and Tartary buckwheat (F. tataricum (L.) Gaertn.). Although common buckwheat is grown throughout the world, mainly in Russia and China, Tartary buckwheat produc- tion is primarily limited to China, Bhutan, and Nepal. Tar- tary buckwheat seeds contain rutin, a major polyphenol, at levels approximately 100 times higher than that found in common buckwheat seeds (Kitabayashi et al., 1995a, b). Consequently, both common and Tartary buckwheat have recently gained attention due to their potential health benefits (Kreft et al., 2020; Kasajima, 2021). Although common buckwheat produces abundant flowers, its seed set is generally low due to self-incompat- ibility, resulting in poor seed yield (Woo et al., 2016). At present, the seed yield of common buckwheat in Japan is less than 1,000 kg ha−1. In contrast, Tartary buckwheat is an autogamous plant with high fertilization efficiency, leading to higher seed yield than common buckwheat. However, pre-harvest shattering, aborting loss, and threshing caused by combine harvester lower the seed yield of both common and Tartary buckwheat (Funatsuki et al., 2000; Morishita and Suzuki, 2017). Additional- ly, Tartary buckwheat is more susceptible to excess soil moisture and salinity than common buckwheat (Matsuu- ra et al., 2005a, b). Buckwheat breeders have been developing varieties by improving plant type, including short plant height and determinate type, to increase yielding ability (Funatsuki et al., 1996; Honda et al., 2009; Morishita et al., 2013). Recently, a useful semi-dwarf common buckwheat line and a Tartary buckwheat cultivar with lodging resistance were developed (Shimizu et al., 2020; Morishita et al., 2015). In terms of cultivation technique, maintaining the ideal plant density through effective crop management is crucial for stabilizing seed yield (Donald, 1963). Plant- ing density is a critical factor that affects the agronomic traits and yield of both common and Tartary buckwheat (Matsui et al., 1974; Xiang et al., 2016; Fang et al., 2018). Previous research on soybeans has shown that yield re- sponse at different planting densities is related to branch development, which is closely related to the ideotype of crops (Agudamu et al., 2016; Peng et al., 2008). Howev- er, an understanding of the effects of planting density on the branching habit in these species is limited. The objective of the present study was to investigate the effect of planting density on the branching habit of common and Tartary buckwheat. This information will be useful for developing optimal crop management prac- tices and ideotypes for these important food crops. MATERIALS AND METHODS The present study used the leading buckwheat vari- eties in Hokkaido, Japan, namely ‘Kitawasesoba’ and ‘Manten-Kirari’ that were developed by the NARO Hok- kaido Agricultural Research Center (Inuyama et al., 1994; Suzuki et al., 2014). The seeds of ‘Kitawasesoba’ were purchased from a commercial source. The cultivation was carried out in an unused field on a local farm, located in Yobito district, Abashiri, Hokkaido, which is the north- ernmost region of Japan, from June to September 2021. The field had peat soil with a slightly acidic texture (pH: 6.1). The study employed a split-plot design with three replicates, resulting in 18 subplots, where the main plots were assigned to two buckwheat species, namely, com- mon buckwheat cv. ‘Kitawasesoba’ and Tartary buck- wheat cv. ‘Manten-Kirari.’ The subplots were assigned to three planting densities, including sparse (67 plants m−2), moderate (111 plants m−2), and dense (222 plants m−2) planting. Each subplot measured 4.8 m2, consisting of four 4 m-long rows, spaced 1.2 m apart, and with a with- in-row spacing of 0.3 m. Border rows were excluded from any investigation. The seeds were planted using seeder tapes (Nippon Plant Seeder Co., Ltd.) at intervals of one every 5 cm (sparse planting), 3 cm (moderate planting), and 1.5 cm (dense planting). The seeding was conducted manually in rows on June 10, 2021. The fertilizer was ap- plied only as a basal dressing for all plants, at the rate of 2 g m−2 of N, 8 g m−2 of P2O5, and 4.7 g m−2 of K2O. The study investigated the main stem length and number of primary branches for 10 individuals of aver- age growth from each plot at the stages of flower bud appearance and full flowering. The stage of flower bud appearance was defined as the day on which flower buds were observed in 40–50% of all plants. The stage of full flowering was defined as the day on which the apical in- florescence of the main stem bloomed in 40–50% of all plants. Owing to the differences in flower bud appearance and full flowering stages between common buckwheat and Tartary buckwheat, the investigation was conducted on an intermediate date, with respective investigations performed on July 12 and 23, 2021. Harvesting was con- ducted on September 3 and 6, 2021 for common and Tar- tary buckwheat, respectively. After harvesting, 10 indi- 16 Kasajima et al. (2023): Effect of planting density on buckwheat branching viduals of average growth from each plot were collected, and their main stem length, number of primary branch- es, number of nodes on main stem, and number of flower clusters on the main stem and branch were measured. All small flower clusters were counted as one cluster. Sub- sequently, the plant samples were examined for seed yield after oven-drying at 80 °C for 48 h, and threshed. In this study, we only investigated the seed yield of Tar- tary buckwheat, as feeding damage by birds was observed in common buckwheat during the seed ripening period. Analysis of variance (ANOVA) was used to evaluate the effect of planting density on the growth and yield charac- teristics in the present study. RESULTS AND DISCUSSION Tables 1 and 2 show the main stem length and num- ber of branches in common and Tartary buckwheat grown under different planting densities at the stages of flower bud appearance and full flowering, respectively. In both cases, the main stem length was significantly longer in common buckwheat compared to Tartary buckwheat, with no significant difference between planting densities. Conversely, the number of branches showed significant differences between planting densities and between the two species (only at the stage of flower bud appearance) but no significant interaction between planting density and species was observed. As planting density increased from sparse to dense, the number of branches decreased for both common and Tartary buckwheat. These findings indicate that while planting density did not significantly impact the main stem length, it had a significant effect on the number of branches at the stage of flower bud ap- pearance. Table 3 shows the results of the main stem length and number of nodes and branches in common and Tartary buckwheat grown under different planting densities at the stage of maturity. The main stem length was significantly shorter in common buckwheat compared to Tartary buck- wheat, but there was no significant difference between planting densities. The number of nodes on the main stem showed significant differences between planting densities and between the two species. The number of branches also showed significant differences between planting den- sities and between the two species. As the planting densi- ty increased from sparse to dense, the number of branches decreased from 3.87 to 1.73 for common buckwheat and from 7.00 to 4.27 for Tartary buckwheat. There was no significant interaction between planting density and spe- cies for both the length of the main stem and the num- ber of nodes and branches. Results presented in Table 3 show that the main stem length of Tartary buckwheat was shorter than that of common buckwheat until flowering, but it became longer at maturity. This result is consistent Species Planting density Main stem length (cm) Number of branches (/plant) Common buckwheat Sparse planting 50.7 3.17 Moderate planting 51.5 2.07 Dense planting 48.8 0.87 Tartary buckwheat Sparse planting 25.7 1.90 Moderate planting 31.1 1.17 Dense planting 32.0 0.13 ANOVA Planting density ns ** species ** ** Interaction ns ns Species Planting density Main stem length (cm) Number of branches (/plant) Common buckwheat Sparse planting 114.9 4.13 Moderate planting 102.6 3.43 Dense planting 104.2 1.93 Tartary buckwheat Sparse planting 87.7 5.00 Moderate planting 96.0 3.70 Dense planting 92.3 2.30 ANOVA Planting density ns ** species ** ns Interaction ns ns * and ** represent significance at 5 and 1%, respectively. ns indi- cates not significant Table 1. Main stem length and number of branches in common and Tartary buckwheat grown under different planting densities at time of flower bud appearance. * and ** represent significance at 5 and 1%, respectively. ns indi- cates not significant. Table 2. Main stem length and number of branches in common and Tartary buckwheat grown under different planting densities at time of full flowering. Fagopyrum 40 (1): 15-21 (2023) 17 with the reports by Kasajima et al. (2012) and Kasaji- ma (2021). Furthermore, the extremely high number of branches (seven) in the sparsely planted Tartary buck- wheat, compared to other planting densities, revealed that the effect of planting density was more significant in Tartary buckwheat than in common buckwheat. Fig. 1 shows the numbers of flower clusters on the main stem and branches of each planting density in common and Tartary buckwheat. The number of flow- er clusters on both the main stem and branches tended to be lower in common buckwheat compared to Tartary buckwheat. Tartary buckwheat is an autogamous plant and has been noted to have a higher yield potential than common buckwheat because of the ease of its seed set (Kasajima et al., 2021). The large number of branches and flower clusters also suggests that these factors contrib- ute to its high yield. Further research is needed to exam- ine the high-yielding traits of Tartary buckwheat from a Species Planting density Main stem length (cm) Number of nodes on main stem (/plant) Number of branches (/plant) Common buckwheat Sparse planting 132.0 13.1 3.87 Moderate planting 122.7 11.8 2.87 Dense planting 120.6 11.9 1.73 Tartary buckwheat Sparse planting 162.4 20.9 7.00 Moderate planting 162.6 20.1 4.43 Dense planting 152.7 18.8 4.27 ANOVA Planting density ns * ** species ** ** ** Interaction ns ns ns Table 3. Main stem length and number of nodes and branches in common and Tartary buckwheat grown under different planting densities at the stage of maturity. * and ** represent significance at 5 and 1%, respectively. ns indicates not significant. Species Planting density Seed yield per plant (g/plant) Seed yield per square meter (g/m2) Tartary buckwheat Sparse planting 1.79 120 Moderate planting 0.93 104 Dense planting 0.82 183 ANOVA ns ns Table 4. Seed yield in Tartary buckwheat grown under different planting densities. ns indicates that the differences among the planting densities are not significant (one-way ANOVA). The seed yield values per square meter were calculated based on the row and hill spacing for each planting density. Number of flower clusters on main stem Number of flower clusters on branches Sparse planting Moderate planting Dense planting Sparse planting Moderate planting Dense planting Common buckwheat Tartary buckwheat 45 40 35 30 25 20 15 10 5 0 N um be r o f fl ow er c lu st er s (/p la nt ) Fig. 1. Numbers of flower clusters on main stem and branches of each planting density in common and Tartary buckwheat. Vertical bars represent standard errors based on three replicates. 18 Kasajima et al. (2023): Effect of planting density on buckwheat branching plant type perspective. There were no considerable differ- ences in the number of flower clusters on the main stem among planting densities in both common and Tartary buckwheat. However, the number of flower clusters on the branches decreased from 18.6 to 5.17 for common buckwheat and from 38.4 to 20.6 for Tartary buckwheat as the planting density increased from sparse to dense. Agudamu et al. (2016) defined branching plasticity in soybean as the ability to adapt to varying planting densi- ties while maintaining a stable yield. This is achieved by controlling branch growth under dense planting condi- tions and promoting more branches under sparse plant- ing conditions to compensate for the reduced main stem yield. In Tartary buckwheat, the seed yield per plant in sparse planting was higher than that of moderate and dense planting, although the seed yield per square meter was the highest in dense planting (Table 4). Branching plasticity may also play a role in the adaptation to differ- ent planting densities while maintaining stable yields in buckwheat as well. Further research is needed to explore the relationship between branching plasticity and yield stability in buckwheat cultivars. Fig. 2 displays the harvested plants obtained from different planting densities of both common and Tartary buckwheat. Sparse planting conditions exhibited more branches and flower clusters compared to those grown under moderate and dense planting conditions, indicat- ing a close relationship with the buckwheat ideotype. The numbers of flower clusters and bloomed florets are con- sidered important factors that govern seed yield (Ujiha- ra and Matano, 1975). Thus, the ideotype of buckwheat is considered a plant type with numerous branches and flower clusters; however, there has been limited research on this topic. In the past, plant types with many branches and flower clusters in buckwheat carried the risk of lodg- ing. Nevertheless, with the introduction of semi-dwarf Fig. 2. Photograph of harvested plant of each planting density in common and Tartary buckwheat. Photograph of common buckwheat (left) on September 3, 2021. Photograph for Tartary buckwheat (right) on September 6, 2021. 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Poljski poskus je bil izveden z uporabo razdeljene ploskve s tremi ponovitvami, pri čemer so primerjali redko (67 rastlin na m2), zmerno (111 rastlin na m2) in gosto (222 rastlin na m2) gostoto setve pri ‚Kitawasesoba‘ ( navadna) in ‚Manten-Kirari‘ (tatarska) ajda. Rastne značilnosti so preverjali v fazah pojava cvetnih popkov, polnega cvetenja in zrelosti. Rezultati so pokazali, da gostota setve ni pomembno vplivala na dolžino glavnega stebla, vendar je pokazala pomembne učinke na število stranskih vej pri obeh vrstah. Z večanjem gostote setve se je število vej zmanjšalo, pri čemer je tatarska ajda pokazala pomembnejše spremembe glede na gostoto setve kot navadna ajda. Z večanjem gostote setve se je manjšalo tudi število socvetij na vejah. Te ugotovitve kažejo na potencialno vlogo plastičnosti razvejanja pri prilagajanju različnim gostotam setve ob ohranjanju stabilnih pridelkov in kažejo na pomen razvoja ideotipov z optimalnimi lastnostmi razvejanja za obe vrsti. Raziskava prispeva k boljšemu razumevanju ideotipov ajde in lahko prispeva k prihodnjim prizadevanjem za žlahtnjenje in način pridelovanja. Fagopyrum 40 (1): 15-21 (2023) 21